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ABSTRACTS

ABDULLA, AMEER A.1,2,3,* and SEAN CONNOLLY.2,4 1 CRC Reef Research Centre, James Cook University, Townsville, Queensland, Australia;
2
Centre for Coral Reef Biodiversity, James Cook University, Townsville,
Queensland, Australia; 3 School of Tropical Environment Studies and Geography, James Cook University, Townsville, Queensland, Australia;
4
School of Marine Biology and Aquaculture, James Cook University,
Townsville, Queensland, Australia. The effects of predation risk on social
coral reef fish at Lizard Island, Great Barrier Reef.
Little emphasis has been placed on non-lethal effects (risk) of predators
that may influence prey behaviour and fitness in marine environments. This
study investigated such non-fatal interactions, which have the potential to
affect the fitness of coral reef fish. Observational data collected from Lizard
Island reefs on the Great Barrier Reef (GBR) show a correlation between
predator abundance and group size of habitat-associated prey. We adopted
a foraging model that predicts predators will significantly reduce the growth
of a prey individual at low prey density, but will have a smaller effect at
higher densities. We tested these predictions in the field using experimental
manipulations of group densities of Pomacentrus moluccensis, a tropical
reef damselfish, and a system of artificial coral reefs and cages at Lizard
Island. Results indicate that the risk of predation affects both the behaviour
and growth rate of the prey. This suggests that suppression of growth due
to predation risk may affect population dynamics of adult coral reef fish
by regulating the rate of individuals reaching reproductive maturity and/or
by increasing the probability of size-selective mortality on juveniles.
ABRAHAM, JOEL K.,1,* JEFFERY D. CORBIN1 and CARLA M.
DANTONIO.1,2 1 University of California, Berkeley, Berkeley, CA;
2
USDA-ARS, Reno, Nevada. Phenology and seedling competition affect
regeneration of perennial grasses in California exotic annual grasslands.
The conversion of California grasslands from perennial-dominated to exotic
annual-dominated ecosystems has been well documented. More recently,
exotic perennial grasses have invaded both grassland types. One of the
theorized reasons for the success of exotic species at the expense of natives
is competitive asymmetry caused by earlier germination and more rapid
seedling growth of exotic species. The rapid emergence and high densities
of annual grasses are thought to reduce resources available to native perennial seedlings, making native restoration difficult. We hypothesized that
decreasing the density of annual grasses and delaying their seedling emergence would increase native perennial performance, while nutrient availability would interact with these conditions. Exotic perennial grasses by
contrast, have rapid seedling growth much like exotic annual grasses so
we hypothesized that they would be less affected by annuals than native
perennials grasses would be. To test these hypotheses, we set up a greenhouse experiment in which an exotic annual grass Bromus diandrus, was
grown in competition with native perennial grasses (Nassella pulchra or
Festuca rubra) or an exotic perennial grass (Holcus lanatus). We maintained Bromus at three competitive densities, manipulated N levels, and
manipulated emergence time by delaying planting of Bromus. As predicted,
increasing the density of annual competitors decreased perennial aboveground productivity and this effect was not as strong for Holcus. Also,
aboveground productivity of Bromus was lower in competition with Holcus
than with natives. Delayed annual emergence increased aboveground productivity in Nassella and Holcus, but had no effect on Festuca. N additions
in non-competition pots increased aboveground productivity in Holcus and
Festuca, while Nassella had no response. However, in competition, Festuca
no longer responded to N addition. These results support other studies
showing that seedling competition between annuals and perennials may be
a critical stage in the regeneration of native grasses in California and suggest that delaying or reducing competition from annuals can increase perennial seedling growth.
ACKERLY, DAVID D.,* WILL CORNWELL and DYLAN SCHWILK.
Stanford Unversity, Stanford, CA. Community assembly, niche conservatism and adaptive evolution in changing environments.
The correspondence between phenotypic variation and environmental conditions (the fit of organisms to their environment) reflects the adaptive
value of plant functional traits. In relatively saturated communities, plants

will establish and regenerate in environments to which they are well adapted, so their distributions (and the distributions of associated functional
traits) will reflect the distribution of environmental conditions. The corollary of this process is that traits related to habitat occupancy (e.g., environmental tolerances) are expected to be under stabilizing selection, leading
to conservatism of niche parameters and related traits over evolutionary
time. Theoretical support for this proposition is provided by habitat selection and community assembly theory. Examples of evolutionary trait conservatism in a phylogenetic, community and biogeographic context are presented, drawing on recent work on woody plants of coastal California.
Based on Jackson and Overpecks (2000) concept of the realized environment, we present three scenarios in which a species distributional responses to environmental conditions will lead to a mismatch between its environmental tolerances and the environments it occupies, thus creating opportunities for adaptive evolution: 1) the colonization of environmental
islands (habitats that are discontinuous in niche space) that require large
adaptive shifts in tolerance of one or more environmental factors; 2) the
persistence of trailing-edge populations in species migrating in response
to changing climate, if barriers to dispersal of competitors prevent competitive exclusion in the deteriorating conditions; and 3) responses to
changes in the realized environment in multi-dimensional niche space, in
which species are predicted to track environmental factors for which they
exhibit narrow tolerances and exhibit adaptive evolutionary response along
axes where they exhibit greater niche breadth. These three scenarios provide a conceptual framework that emphasizes the role of ecological sorting
processes and stabilizing selection as the context for adaptive evolution in
heterogeneous and changing environments.
ADAMS, HENRY D.* and THOMAS E. KOLB. School of Forestry,
Northern Arizona University, Flagstaff, AZ. Using radial growth and
foliar d13C as measures of drought sensitivity in trees at ecotone forest
communities in northern Arizona.
Climate change has the potential to seriously impact forest ecosystems in
northern Arizona. An increase in temperature and changes in the amount
and distribution of precipitation may alter forest growth and tree species
composition. Past tree responses to climatic variability may provide insight
about future forest responses. We used two approaches to compare drought
sensitivity among tree species in ecotone forests of northern Arizona: 1)
the ratio of radial growth during dry years to growth during wet years (D/
W) for selected years between 1950 and 2000, 2) leaf d13C for 2001 (average precipitation) and 2002 (severe drought). At the pinyon-juniper
woodland - ponderosa pine forest ecotone, we compared drought sensitivity
between Pinus ponderosa and P. edulis on three soil types based on different parent materials representing a gradient in water availability: sedimentary, flow basalt, and basalt cinders. At the ponderosa pine - mixed
conifer forest ecotone we compared sensitivity among P. ponderosa, P.
flexilis, and Pseudotsuga menziesii. We expected that trees growing in soils
with greater water availability and higher in their elevational range would
be less sensitive, and sensitivity would vary among co-occurring species
and would be similar based on D/W and d13C. At low elevations P. ponderosa was more sensitive to drought based on D/W than P. edulis and
was more sensitive on cinder soils. In contrast, response of d13C to drought
suggested no difference between these species at the leaf level. P. ponderosa at the ponderosa pine - mixed conifer ecotone was less sensitive than
P. flexilis and Pseudotsuga menziesii based on both D/W and d13C, and P.
ponderosa at high elevations was less sensitive than at low elevations for
both measures of sensitivity.
ADKISON, GREG1,* and SCOTT GLEESON.2 1 Western Carolina University, Cullowhee, NC; 2 University of Kentucky, Lexington, KY. Distributions and light adaptations in a set of forest violets (Viola).
A species9 distribution is a pattern of abundance that reflects the match
between trait values and the environment. Limiting trait theory predicts
that a species9 abundance peaks in habitats where traits that potentially
limit fitness are optimal and that it declines as these traits become suboptimal. This prediction was studied in the understory of an eastern deciduous
forest by examining four traits in several violets (Viola), a group that stood
out in prior community-level work because of their striking range of distributions. V. blanda and V. sororia span much of the primary gradient in

Abstracts

this forest, whereas V. hirsutula, V. triloba, and V. canadensis have rather

narrow distributions. V. pubescens and V. rostrata were particularly interesting because their distributions overlap along much of the gradient but
peak at opposite ends. Individuals were randomly sampled in six distinct
sites along a transect that spanned the forest9s primary environmental gradient. Several intriguing patterns were detected, especially in V. pubescens
and V. rostrata. First, the trait values for these two species fall fairly close
to the community mean trait values along the gradient. Second, in four
cases (V. pubescens SLA; V. rostrata leaf size, etiolation, leaf:shoot) the
traits are fixed across the gradient. In the other four cases (V. pubescens
leaf size, etiolation, leaf:shoot; V. rostrata SLA) trait values change across
the gradient, presumably due to phenotypic plasticity or ecotypic differentiation. In each case the shift is in the same direction as the community
mean and generally with lower slope. Interestingly, the one trait that is
fixed in V. pubescens is the only trait that is variable in V. rostrata. Third,
except for leaf:shoot in V. pubescens, all patterns of deviation from the
community mean across the gradient are consistent with trait limitation:
deviation increases away from the peak niche position.
ADLER, LYNN S.* Virginia Tech, lsadler@vt.edu, Blacksburg, VA. Induced susceptibility to leaf herbivores via floral induction.
Traits that are attractive to mutualists may also be attractive to antagonists,
and the evolution of phenotypic traits will depend upon the combined selection pressure due to multiple interactions. Many species of Lepidoptera
are pollinators as adults and herbivores as larvae, raising the possibility
that adults use floral or nectar traits to make oviposition decisions and
evaluate plant quality. This may be particularly true for species that have
secondary compounds in nectar if levels of nectar defenses are correlated
with levels of leaf defenses. I tested the hypothesis that leaf damage by
the tobacco hornworm, Manduca sexta, and fertilizer level in domestic
tobacco, Nicotiana tabacum, would affect floral morphology, nectar composition, and oviposition choice by adult M. sexta moths. Damaged leaves
were removed from plants prior to moth choice tests to eliminate the possibility of decisions based directly on the presence of damage. I found that
fertilizer and damage both affected several floral morphological traits, including corolla length, width and diameter. Adult moths preferred to oviposit on high-fertilizer plants, and oviposited more on damaged than control
plants when fertilizer levels were high but not when fertilizer was low.
Thus, it appears that damage provides some indirect cue to moths making
oviposition choices, but only when resources are available. Nectar composition has not yet been analyzed but I hypothesize, based on results from
leaf induction, that damaged plants will have higher concentrations of nicotine in nectar and that fertilizer will enhance this effect. In future work I
will experimentally manipulate nectar nicotine in field plants to determine
if moths use this cue explicitly to make oviposition decisions. This research
demonstrates that selection on the production of defensive compounds such
as nicotine may be mediated via floral as well as leaf interactions.
ADLER, PAUL R.,* MATT A. SANDERSON and SARAH C. GOSLEE.
USDA-ARS, University Park, PA. Survey of CRP and other grasslands
in the northeastern USA.
Grassland ecosystems are important wildlife habitat and have the potential
to be a significant component of the new biobased economy. Most currently
established grasslands in the Northeastern USA are on land with marginal
crop production potential. Little is known about the plant composition or
amount of biomass produced on these grasslands. To assemble a database
for the resource assessment of warm season grasslands in the Northeastern
USA we determined plant species composition at multiple scales using the
modified Whittaker plot technique, measured various soil properties, and
quantified biomass yield on CRP, WHIP, mine reclamation, and other grasslands. A total of 22 grasslands were sampled in New York, Pennsylvania,
New Jersey, Maryland, and Virginia during September and October 2002.
We identified over 180 different plant species across the study region. Species richness was about 35 and biomass was about 6000 kg/ha, but both
were quite variable by site. Although biomass yields were substantially
lower than have been reported for monotypic switchgrass fields, minimal
inputs may substantially increase the yields.

Abstracts

ADLER, PETER B.,1,* WILLIAM K. LAUENROTH,1 OSVALDO E.

SALA,2 DANIEL G. MILCHUNAS1 and INGRID C. BURKE.1 1 Colorado
State University, Fort Collins, CO, USA; 2 Universidad de Buenos Aires,
Buenos Aires, CF, Argentina. Functional traits and grazing resistance
in graminoids of sagebrush steppe, USA and Patagonian steppe, Argentina.
We measured grazing tolerance and avoidance traits of graminoids collected at sites in the Patagonian steppe of Argentina and the sagebrush
steppe of the northwestern U.S. to evaluate the role of evolutionary history
of grazing in determining plant traits and to generate predictions about the
vulnerability of these ecosystems to grazing impacts. To measure grazing
tolerance, we conducted a greenhouse defoliation experiment on dominant
grasses from each region. The poor performance of species that increase
with grazing in the field suggests that either tolerance is less important than
avoidance in determining plant population responses to grazing, or greenhouse conditions fail to reproduce field responses. To compare avoidance
traits, we measured both morphological traits and leaf chemical composition on common species from each region, then performed a Principal
Components Analysis on the species by traits matrix. The dominant species
from the arid Patagonia site scored significantly lower than the sagebrush
steppe dominants on the first axis, correlated with measures of forage quality such as N and cellulose content. Plants from the wetter Patagonia site
were intermediate in forage quality. Sagebrush steppe species scored significantly lower on the second axis, indicating greater plant height and leaf
length. These differences are consistent with evidence indicating a more
intense evolutionary history of grazing in Patagonia, though subtle environmental differences offer alternative explanations. Regardless of their
origin, these differences in traits can explain why the introduction of domestic livestock has had different effects on vegetation in these ecosystems.
AGRAWAL, ANURAG A.1 and NORA UNDERWOOD.2 1 University of
Toronto, Toronto, Ontario, Canada; 2 Florida State University, Tallahassee,
Florida. Population growth rate as a general predictor of the strength
of density dependence.
Although ecological studies of density dependence have yielded highly
variable results, little is known about the conditions that may promote
strong versus weak density dependence. Thus, a critical hurdle in developing a predictive framework for population dynamics is understanding the
causes of this variation. Here we present experimental evidence for positive, negative, and no density dependence from 12 independent density
manipulations of milkweed aphids in laboratory and field experiments. A
general pattern emerged from these highly variable results: greater experiment-wide per capita growth rate of aphids was associated with stronger
density dependence (a more negative slope of the relationship between
density and per capita growth rate). This pattern was robust across a broad
range of aphid densities; the initial or final mean aphid density for experiments did not predict the strength of density dependence. Experimental
manipulation of temperature demonstrated that shifts in population growth
rates were associated with changes in the strength of density dependence.
These results suggest a general hypothesis: as density independent factors
increase mean per capita growth rates, the slope of density dependence
decreases. Given certain assumptions, this prediction follows from the definition of density dependence that is included in many widely used models
of population dynamics. Thus, the relationship between population growth
rate and density dependence is likely to be broadly applicable as a determinant of variation in the strength of density dependence.
AHUMADA, JORGE A.,1,* MICHAEL SAMUEL,1,4 ANDREW P. DOBSON,2 DENNIS LAPOINTE3 and CARTER ATKINSON.3 1 USGS/National Wildlife Health Center, Madison, WI; 2 Princeton University, Princeton,
NJ; 3 USGS/Pacific Islands Ecosystems Research Center, Volcano, HI;
4
Univerity of Wisconsin-Madison, Madison, WI. A simple individualbased model for the transmission of avian malaria in Hawaii.
We present an individual based model that simulates the transmission of
avian malaria (Plasmodium relictum) by Culex mosquitoes (Culex quinquefasciatus) on honeycreeepers in the big island of Hawaii. We explore
the properties and predictions of this model as we vary the intensity and
statistical distribution of biting rates on hosts, the numerical ratio of vector

and hosts, the differential attraction of vectors to infected hosts and the
population dynamics of vectors along an elevational gradient. We also suggest ways to estimate some of the transmission parameters (particularly
bitting rates on hosts) using field measurements of parasite prevalence on
hosts and vectors and their population abundance.
AIKIO, SAMI. University of Helsinki, www.helsinki.fi/;saikio/, Helsinki,
Finland, Finland. Asymmetric competition between relatives.
Hamiltons rule predicts that individuals attempt to maximize their inclusive fitness. The number of offspring produced by an individual is typically
a non-linear function of its gain of a limiting resource. Individuals resource
gain depends on the availability of the resource, the number of individuals
in the population competing for the resource, and the distribution of the
resource among them. I carried out a theoretical analysis, asking whether
related individuals should distribute a resource in a different way than a
population of unrelated individuals does. I also studied the optimal distribution of a resource from the view-point of different sized individuals and
the effects of optimal resource distribution on population dynamics. I found
that the individuals which are most efficient in converting the gained resources into offspring production, are predicted to get the largest share of
the resource when population consists of related individuals. The optimal
resource distribution was rather symmetric in low population densities, but
asymmetric in high population densities, which stabilized population dynamics. The results suggest that altruistic resource share can evolve also
in viscous populations of equally related individuals, which was earlier
considered not to be possible. The difference is due to the assumption of
non-linear resource use efficiency in offspring production made in the present study, which contrasts to the earlier assumption of altruistic behaviour
and offspring production being directly related to each other.
ALABACK, PAUL. University of Montana, Missoula, Montana. A multiscale model for monitoring biodiversity.
Biodiversity is an inherently complex multi-scale phenomenon. The characterization of biodiversity therefore requires a clear conceptual model in
which disparate sources of data can be carefully integrated together. Many
plot-based approaches to monitoring species and ecosystems have been
successful in characterizing common elements and their dynamics, but often miss the rarer species. We propose an application of hierarchy theory
to the biodiversity problem whereby data sets of different scales are brought
in to characterize key processes in maintaining biodiversity. Recent studies
have emphasized the role of species pools in constraining biodiversity patterns at regional scales. We explore the use of large-scale and regional
species lists as a means to estimate gamma diversity, and more traditional
plot data to estimate alpha diversity for key habitats. Remote sensing and
landscape modeling approaches are used to define general patterns of species richness to identify biodiversity hot spots. These models can then be
used to guide more detailed field sub-sampling to estimate medium to rare
species elements with mega-transects. In many instances regional data are
available for the reconstruction of a historical or baseline reference condition 20 years or more ago. Re-sampling of rare species distributions and
consideration of contemporary changes in landscape structure can then be
used to establish models of potential changes from environmental stressors
over this time period. The multi-scale approach provides a rich tool set
from which it may be possible to more clearly disentangle the effects of
region-wide patterns from localized phenomena and identify likely processes to effect these changes. By emphasizing relatively rare species, using
broad-scale inventories that emphasize distribution rather than abundance
it may be possible to develop monitoring programs that are more sensitive
to detecting subtle environmental change than more traditional plot-based
sampling approaches
ALBANI, MARCO* and PAUL R. MOORCROFT. Harvard University,
Cambridge, MA. Modeling the relative importance of land use history
and CO2 fertilization in forest carbon dynamics of the eastern U.S.
Atmospheric measurements and inventory data suggest that since the
1980s the forests of North America are storing between 0.15 and 0.30 Pg
C year-1, a substantial component of the continental carbon sink. Two fundamental mechanisms for the accumulation of carbon are 1) the recovery

of forest ecosystem from previous land uses; and 2) the enhancement of

growth by CO2 fertilization. Assessing the magnitude of C accumulation
that is due to each has fundamental implications for the long-term future
of the North American carbon sink. In this study we use a mechanistic
individual-based terrestrial ecosystem model, the Ecosystem Demography
(ED) model, to estimate the magnitude of the two terms. Unlike conventional global vegetation models, ED is capable of mechanistically account
for both the effect of CO2 fertilization on plant growth and the impact of
land use changes on plant demography, stand composition and size structure, stand age structure, and ecosystem productivity, all variables that can
be directly compared to inventory data. We use the ED model to simulate
the historical pattern of land cover and carbon accumulation in the eastern
portion of the conterminous U.S at 18 resolution, forcing the ecosystem
model with an historical dataset of land use change and atmospheric CO2
concentration. We obtain for each grid cell different scenarios of current
rates of Above-ground Net Ecosystem Productivity (ANEP) and current
levels of Aboveground Biomass (AGB) as a function of forest stand age.
Modeling results are compared to the data from the US Forest Inventory
and Analysis, and we assess the ability of each mechanism to predict the
observed patterns of ANEP and AGB variation with stand age. While land
use history affects the age-class distribution within each simulation cell,
CO2 fertilization modifies the historical rates of ANEP and thus directly
impacts the current relationship between AGB and stand age, enabling us
to assess the magnitude of carbon accumulation due to each mechanism.
ALBER, MERRYL. University of Georgia, malber@arches.uga.edu, Athens, GA. A River Runs To It: The effects of changing inflow to estuaries.
An often-overlooked consideration in the management of freshwater flow
in rivers is the eventual effect that different flow regimes could have on
coastal ecosystems. Changes in freshwater inflow (in terms of the quantity,
quality, and timing of water delivery) affect estuarine conditions (such as
salinity and concentrations of dissolved and particulate material), and, in
turn, estuarine resources (such as the distribution and abundance of organisms). This paper will provide an overview of the scientific evidence for
the consequences of changing freshwater inflow to estuaries, and then present a conceptual model that explores the roles of scientists, citizens, politicians, and managers in developing management approaches to address
this issue. The premise of the conceptual model is that the goal of estuarine
freshwater inflow policy is to protect those resources and functions that we
as a society value in estuaries, and that management measures use scientific
information about the relationships among inflow, conditions, and resources
to establish inflow standards that can meet this goal. The model will be
used to explore the ways in which freshwater inflow criteria were established for Texas, Florida, and California, as well as to describe progress
towards the development of freshwater inflow relationships for Georgia
estuaries.
ALBERS, GAYLE L.1 and MERRYL ALBER.2 1 University of Georgia,
Athens, GA, USA; 2 University of Georgia, Athens, GA, USA. Vascular
plant composition and soil characteristics of undeveloped back-barrier
islands near Sapelo Island, Georgia.
This study was designed to describe vascular plant composition and soil
characteristics among undeveloped back-barrier islands of different sizes
near Sapelo Island, Georgia. Known colloquially as marsh hammocks,
back-barrier islands are erosional remnants nested within estuarine salt
marshes. As the population of coastal Georgia continues to grow, there is
increasing interest in developing these areas for residential use. The state
has identified over 1200 hammocks along the Georgia coast, totaling approximately 6900 hectares of upland, but little information is available
regarding their ecological significance. In the summer and fall of 2002, we
surveyed vegetation on 11 hammocks having minimal recent anthropogenic
impacts. We found that overall vascular plant diversity is low, but in keeping with the theory of island biogeography, species richness tends to increase with back-barrier island size. Mean species richness values ranged
from 6.5061.64 for back-barrier islands less than 1.0 ha to 9.9462.10 for
those greater than 4.0 ha. The upland woody plant community is dominated
by yaupon holly (Ilex vomitoria) and is found in association with representative maritime forest species of Georgia. Soil analyses indicate rela-

Abstracts

tively low nitrogen (#2%) and highly variable carbon values (1.7-48%).
Planners, resource managers, and conservation organizations may use these
findings to help designate important natural areas and develop projects or
policies that promote their sustainable use.
ALBRECHT, MATTHEW A.* and BRIAN C. MCCARTHY. Ohio University, Athens, Ohio, United States. Habitat specificity and temporal
trends in harvesting of medicinal herbs in the Wayne National Forest
(Ohio, USA).
The burgeoning demand for wild harvested herb products has exerted considerable pressure on deciduous forest plant populations. Since effective
conservation and management strategies are constrained by the lack of
baseline ecological information, we employed a strip transect sampling
scheme to describe the distribution patterns of economically important forest herbs in the Wayne National Forest (WNF). This study also examined
temporal variation in medicinal plant harvesting by evaluating plant collection permit data over a six-year period (1995-2001). Our permit analysis
revealed a linear increase in the frequency of harvesting black cohosh (Actaea racemosa L.), bloodroot (Sanguinaria canadensis L.) and blue cohosh
(Caulophyllum thalictroides L. Michx.) (P , 0.001). Overall, the total
number of plant collection permits issued in the 2001 harvesting season
increased by 180% when compared to the 1995 harvesting. Our sampling
scheme found that American ginseng (Panax quinqefolius L.) and Virgninia
snakeroot (Aristolochia serpentaria L.) were rare, but broadly distributed
in a variety of microhabitats. Slope aspect was a significant predictor of
black cohosh occurrence (P , 0.05). The frequency of blue cohosh was
greater on north facing aspects than east facing aspects (P , 0.05). Forest
stands designated as protected natural areas, did not harbor greater medicinal herb abundance than unprotected areas (P . 0.05). Our study suggests
that the patchy distribution and high interspecific variation in habitat specificity complicates the efficient management and sustainable harvest of key
non-timber forest resources. Projecting our plant collection permit analysis
into the future indicates that immediate action should be taken to develop
ecologically based sustainable models to prevent overharvesting.
ALERIC, KATHERINE M.1,2,* and L. KATHERINE KIRKMAN.1 1 J.W.
Jones Ecological Research Center, Newton, GA; 2 University of Georgia,
Athens, GA. Growth responses of the federally endangered shrub, Lindera melissifolia (pondberry), to varied light environments.
Lindera melissifolia, pondberry, is a federally endangered shrub that occurs
in seasonally flooded wetlands. Although pondberry populations occur in
dense shade and full sun environments, the optimal conditions for growth
are unclear and differences in observations have led to conflicting management interpretations. Increasing vigor of existing populations is critical
for species persistence, particularly due to low sexual reproduction. The
goal of this study is to determine the optimal light requirements for growth
by comparing morphological and photosynthetic responses of plants growing under natural and controlled light environments. In the first year of this
study, growth responses and light conditions were measured in three natural
populations in Georgia and South Carolina. Preliminary findings indicate
that pondberry has the ability to acclimate to varied light conditions by
displaying typical sun-shade morphological and photosynthetic responses.
Photosynthetic capacity was consistent with other shade tolerant species
with maximum rates of photosynthesis ranging between 2-8 mmol
CO2m22s21. Leaves became saturated at relatively low light levels of 250400 mmol m22s21 PAR. The low photosynthetic capacity and saturation
irradiance found even in high light environments suggests that this species
is adapted to low light conditions. Currently, plants propagated by cuttings
are being grown in three light treatments using shade cloth over support
frames. Cuttings have also been outplanted in contrasting canopy cover in
four wetland sites at Ichauway, the J.W. Jones Ecological Research Center.
The information from both field and controlled experiments will be used
to develop management strategies for the removal of competing vegetation,
if necessary, and in identifying suitable sites for new population establishment.

Abstracts

ALERS-GARCIA, JANICE,* JAMES D. BEVER and KEITH CLAY. Indiana University - Bloomington, jalersga@bio.indiana.edu, Bloomington,
IN. Effect of the parasitic plant, Cuscuta gronovii, on size inequality
in populations of Pilea pumila.
Parasitic plants are characterized by obtaining water and nutrients from one
or a few host individuals causing some harm but not their immediate death.
Given the antagonistic fitness effects present in this parasite-host interaction, we predict that host choice and its effects on individual host plant
sizes can affect the development of size inequality in host populations. We
used a matched pair design (n536) to examine whether Cuscuta gronovii
(dodder) a holoparasite, affects the development of size inequality of its
host Pilea pumila. Cuscuta gronovii did not affect the mean population
size of its host(Fdf351.63, P50.193). Nevertheless, parasitized populations
had a significantly high size inequality for height (F523.87, P50.0001)
and shoot biomass (F56.57, P50.0126) throughout the growing season
compared to non-parasitized populations. This increase in size inequality
was accompanied by an increase in negative skewness for height and a
decrease in positive skewness for shoot biomass by the end of the season.
This indicates that parasitized populations contained a small group of extremely small individuals (short and low biomass). On the other hand,
Cuscuta exhibited a significant increase in biomass (Fdf254.646, P50.0167)
during the growing season, but did not increase in number of hosts infected.
These data indicate that parasitic dodder selects its hosts early in the growing season, which are then intensively parasitized throughout the growing
season. Indeed, examination of mean size differences among parasitized
and unparasitized individuals on population assemblages with Cuscuta
showed that taller individuals of Pilea pumila are preferentially parasitized.
This preferential parasitism strongly depressed their height and biomass
throughout the season. These results suggest that size dependent parasitism
by highly selective holoparasites such as Cuscuta gronovii could have
strong effects in the development of size inequality and thus potential competitive interactions of its host population.
ALEXANDER, HELEN M.,1,* JANIS ANTONOVICS2 and PETER H.
THRALL.3 1 University of Kansas, Lawrence, Kansas; 2 University of Virginia, Charlottesville, Virginia; 3 CSIRO, Canberra, A.C.T., Australia. Spatial patterns of disease incidence in plants: metapopulations and beyond.
The majority of host-pathogen studies are focused on small spatial scales,
yet plants and pathogens exist in patches, populations, metapopulations,
and regional assemblages. At large spatial scales, both abiotic factors and
biotic factors (such as host density and isolation of host patches) will affect
disease patterns. We will discuss three types of methods for studying disease processes at large spatial scales. First, for some plant hosts, we can
define discrete host patches, and disease incidence can be studied between
and among patches. Second, for plant hosts with generalist habitats, discrete
host patches cannot be easily defined, but large scale processes can be
studied by superimposing a grid over favorable plant habitat, and studying
plants and their diseases in arbitrary units of space. Third, herbarium collections can provide a surprising source of information on spatial, as well
as temporal, patterns of disease incidence. We will illustrate these methods
with three host-pathogen systems. For studies of discrete host patches, we
will describe studies of Alternaria infection of the beach plant, Cakile
maritima, where discrete plant populations exist on beaches that are separated from other beaches by rocky intertidal areas. To illustrate research
on diseases of plants with habitat generalists, we will discuss infection of
Silene alba by the anther smut pathogen, Microbotryum violaceum, and
infection of Helianthus annuus by the rust Puccinia helianthi. Finally, we
will illustrate research with herbarium specimens with studies of anthersmut infection of Silene species. For all methods and systems, research on
larger spatial scales provides perspectives that are not apparent from smallscale studies.
ALLEN, CHRISTOPHER B.,1,* RODNEY E. WILL,1 DAVID R. COYLE2
and MARK D. COLEMAN.2 1 Warnell School of Forest Resources, University of Georgia, Athens, GA, USA; 2 USDA Forest Service, Savannah
River, New Ellenton, SC, USA. The effect of resource availability on
canopy dynamics and biomass accumulation in fertigated hardwood
stands.
To determine how resource availability affects canopy dynamics, leaf-level
physiological parameters, and biomass accumulation, we measured the im-

pacts of water and nutrient additions on intercepted photosynthetically active radiation (IPAR), leaf gas exchange parameters, foliar nitrogen concentration, specific leaf area, and xylem water potential (C) during the 2001
growing season. Four genotypes (Liquidambar styraciflua, Platanus occidentalis, and two Populus deltoides clones with differing drought tolerances) planted in the spring of 2000 received a factorial combination of irrigation and fertilization. The four treatments were: control (0.5 cm water
wk21), irrigation only (3.0 cm water wk21), fertilization only (0.5 cm water
wk21, 120 kg N ha21 yr21), and irrigation with fertilization (3.0 cm water
wk21, 120 kg N ha21 yr21). Beginning in March 2001, IPAR measurements
for each genotype were made every 6 weeks during the growing season.
Leaf gas exchange, foliar nitrogen concentration, specific leaf area, and C
were measured 4 times during the growing season on L. styraciflua and P.
occidentalis. The addition of water and nutrients increased biomass accumulation for all species. When between-block variability was removed,
IPAR was well correlated with biomass accumulation for all genotypes (r2
range from 0.54 to 0.99). While there was no impact of treatment on photosynthetic capacity (Amax), irrigation significantly increased stomatal conductance (gs) (p 5 0.001; p 5 0.009) and leaf internal CO2 concentration
(Ci) (p 5 0.003; p 5 0.02) for L. styraciflua and P. occidentalis, respectively. Foliar nitrogen concentration was not affected by treatment and was
a poor predictor of biomass accumulation. There were no treatment effects
for either specific leaf area or C. These results indicate that IPAR may
play a large role in the accumulation of biomass under a broad range of
resource availabilities.
ALLEN, CRAIG R.,1,* JAN SENDZIMIR2 and GARRY PETERSON.3
1
South Carolina Cooperative Fish and Wildlife Research Unit, Clemson,
South Carolina; 2 International Institute for Applied Systems Analysis, Laxenburg, Austria; 3 McGill University, Montreal, Quebec. Cross-scale
structure and resilience in animal communities.
Species interact with scale-dependent sets of ecological structures and processes that determine their functional opportunities. The cross-scale resilience model proposes that animal communities are compartmentalized by
scale. It suggests that strong interactions within a range of scale lead to a
diversity of functions within a scale, because members of the same functional group are likely to strongly interact and tend to coexist only by
operating at different scales, reducing competitive interactions. It further
suggests that ecological resilience is generated by diverse overlapping functions within a scale and by apparent functional redundancy at different
scales, thereby reinforcing function across scales. To test those propositions
we, 1) determined if the body mass distributions from a large number of
diverse ecosystems were discontinuously distributed, 2) determined if body
mass distributions were over-dispersed at the community level and at the
within-aggregation (scaled) level, and 3) determined if functional groups
were distributed evenly across scales. We utilized 33 terrestrial vertebrate
data sets from 22 ecosystems. We tested for discontinuities in animal body
size distributions with null models and cluster analysis. Spacing between
species (segment length along the body mass axis) within each body mass
aggregation was calculated and the evenness of spacing was quantified by
determining the variance of segment lengths within aggregations. Observed
variances were compared with simulations, with low variances indicating
more even spacing. We utilized feeding guilds as surrogates for functional
groups, and determined the variance in the distribution of guilds among
body mass distributions, and compared that with the output of simulations.
Observed segment length variances tended to be small relative to the variance distributions generated by simulation. There was also low variance
in the distribution of feeding guilds. Our results suggest that body mass
distributions are discontinuous, and support the predictions of the crossscale resilience model.
ALLEN, ELIZABETH A.* and ROBERT NOWAK. University of NevadaReno, Reno, Nevada. The immediate effect of fire on the soil seed bank
of pinyon-juniper woodlands.
Fire is a common agent of disturbance that can vegetatively denude an
area. Once a site is denuded, the soil seed bank is a major source for new
plant recruitment and therefore important to understand. I am studying the
immediate effect of fire on the seed bank in the P-J woodlands of central
Nevada. My study site is comprised of three medium density P-J plots that

contain three microsites: under-tree, under-shrub and interspace. On May

11, 2002 a prescribed burn was applied to the plots. I obtained pre-burn
soil samples a week before the fire and post-burn soil samples one week
after the fire. Because different microsites have differing abilities to trap
seeds, I obtained 10 samples from each of the three microsites. I also took
the soil samples at different depths: litter (under-tree only), 0-1cm, 1-5 cm
and 5-10 cm. These soil samples were placed in flats in a greenhouse and
induced to germinate. Surprisingly, the data have shown that the number
of seedlings emerging from the post-burn samples increased by 67%. I had
hypothesized that there would be a loss of viable seeds in the soil for two
reasons. First, heat from the fire would have deleterious effects on the seed
bank. Second, in May there would be no new influx of seeds to the soil,
and the number of viable seeds in the seed bank should have decreased
between the pre- and post-burn samples due to natural germination at the
site. The under-tree microsite is driving the increase, especially the 0-1 cm
layer where the number of seedlings increased 4.5 times. The two most
common species within this microsite and at this depth are the native annual mustard Descurainia pinnata and quaking aspen, Populus tremuloides.
Various mechanisms to account for the increase in seed germination are
discussed.
ALLEN, TIMOTHY F.H. and AMANDA J. ZELLMER. Botany Dept.,
University of Wisconsin, 430 Licoln Drive, Madison, WI, USA. Complexity and Sustainability, Part II: Ecological Complexity.
Complexity, less a matter of material nature, is more a question of normative values. With a paradigm there is agreement on structure vs. behavior, continuity vs. discreteness, meaningful change vs. dynamics, all reflecting a level of analysis. Paradigms get rid of ambiguity by fiat; the system
may remain complicated, but it is simple, not complex. Without a paradigm, the above distinctions remain unspecified, and complexity emerges
from the infinite possibilities for defining experience of whatever is beyond
our observational choices and conceptual frame. So non-linearity, emergence, uncertainty, difficulty predicting, chaotic behavior, and multiplicative ambiguous relationships are not directly material considerations, but
are a consequence of not having decided completely on how to look at and
specify the system. In complex organized structures, there is a loop between
observable structures, such as salmon, and a context that generates those
structures. There is another loop that also passes through those same structures, and defines them into a class, say endangered species, that amounts
to a model. Returning from the class closes the modeling loop in verification of membership. Both loops involve change: in the models of those
structures; in the context of the structures themselves. If the contextual
loop breaks, then without a viable context the structures we seek to sustain,
be they social, economic or biogeophysical, begin to fail. On the modeling
side, if the scientist/manager fixates on some level of toxicity or rule such
as the 50/500 rule for endangered species, then the science/management
gets stuck, and is left behind by the other loop involving context. Supplyside sustainability says manage from the context. Sustainability invokes
long term views and models, and so is about maintaining both loops as
functional. In all this it is crucial to have a firm grasp of the layers of
subjectivity in the modeler/manager.
ALLISON, GARY W.,1,2,* MARIA KAVANAUGH,2 SHERI ETCHEMENDY,2 CARL SCHOCH,3 JANE LUBCHENCO2 and BRUCE A. MENGE.2
1
The Ohio State University, Columbus, OH; 2 Oregon State University,
Corvallis, OR; 3 Kachemak Bay National Estuarine Research Reserve, Homer, AK. A latitudinal gradient of diversity in the near-shore: assembling the pieces across scales.
Latitudinal gradients in diversity have provided important patterns for ecological debate. Such patterns are often estimated from species ranges or
other indirect means. Because patterns span such large areas, they are rarely
verified with on-the-ground measurements. Because of this lack of verification, the reliability of macroecological datasets and their usefulness in
both quantifying pattern and evaluating mechanisms can be uncertain. We
have begun to address this and related issues by establishing an extensive
and intensive monitoring program of rocky intertidal community structure.
Here we use this dataset to evaluate a purported gradient of macroalgal
diversity across the west coast of the continental US. This survey quantifies
abundance of 250 taxa in 5000 quadrats annually. Sampling is spatially

Abstracts

nested and includes both biotic and physical measurements to help determine the primary physical predictors of community structure. Across this
large area, the direction of the gradient we find in macroalgal diversity
contradicts what was expected based on range data: higher diversity is
found at higher latitudes. Among the primary explanations for this unexpected diversity pattern are, with increasing latitude: increasing tidal excursion, increasing wave force (both potential proxies of or influence on
total habitat area), decreasing temperature, and increasing nutrient availability. Although many of the potential factors co-vary across the survey,
we matched scales of variation in diversity with the variation in these
factors to help us identify the best predictors. Preliminary analyses suggest
that area is a major determinant of the diversity pattern. Although funding
agencies avoid supporting "monitoring" programs, our results suggest that
such programs are critical for the development of rigorous databases that
will be useful in accurately quantifying macroecological patterns, and for
pointing the way towards evaluation of likely mechanisms.
ALLISON, STEVEN D.* and PETER M. VITOUSEK. Stanford University, Stanford, CA, USA. Decomposition and nutrient dynamics in native and exotic Hawaiian understory plant litter.
We studied decomposition rates and nutrient dynamics in 15 understory
plant litter types from the Hawaiian islands in control and N + P fertilized
plots. The 15 litter types included 6 exotic species, 4 types of stem material,
and 5 fern species. Using measurements of initial litter chemistry, litter
nutrient dynamics, and mass loss over time, we investigated the potential
for changes in litter inputs to affect ecosystem nutrient cycling. We found
a 100-fold variation in litter decay rates, with native leaf litter decaying at
rates of 0.2-2.3 yr-1 and exotic leaf litter at rates of 1.4-9.4 yr-1. Angiosperm
litter decayed more rapidly than fern litter, and leaves generally decayed
more rapidly than stems. Fertilization increased the decay rates of exotic
litter types, but left most native litter decay rates unchanged. Only litter
from 3 native ferns and litter from an exotic ginger was able to retain or
immobilize N after 100 days of decomposition. All litter types except the
recalcitrant native ferns lost . 50% of initial P during decomposition.
Fertilization increased nutrient immobilization or delayed the onset of nutrient loss for most litter types. If native understory plants are displaced by
invasives, nutrient cycling rates could increase dramatically due to rapid
decomposition and nutrient release in exotic litter. Rapid decay rates and
acceleration of exotic litter decomposition in response to nutrient additions
could create a positive feedback between invasion, decomposition, and nutrient cycling rates.
ALLISON, VICTORIA J.,* MIKE MILLER, JULIE D. JASTROW and
ROSER MATAMALA. Argonne National Laboratory, Argonne, IL, USA.
Characterization of environmental and edaphic factors affecting soil
microbial communities using a tallgrass prairie restoration chronosequence.
We examined environmental factors regulating soil microbial community
structure, using a tallgrass prairie restoration chronosequence located at the
Fermi National Accelerator Laboratory in Batavia, Illinois. This chronosequence allows us to examine microbial community structure over a wide
range of soil and biotic conditions. Soil microbial community structure was
determined using phospholipid fatty acid (PLFA) patterns, and summarized
by correspondence analysis. We found a clear difference between agricultural and restored prairie plots. The prairie plots had considerably higher
relative amounts of the fungal signature PLFA 18:2:omega:6, and the arbuscular mycorrhizal fungal signature 16:1:omega:5c. In contrast, agricultural plots had higher relative amounts of cy17:0 and cy19:0; markers for
gram-negative bacteria. Further, the ratio of cyclopropyl fatty acids to their
precursors was considerably lower in prairie than in agricultural plots indicating higher substrate availability in the restored prairie soils. Regression
of ordination plots against environmental variables indicated that the chronosequence represents an aggrading soil system, with microbial composition
related to a suite of environmental variables, most notably increased production of root biomass, surface litter accumulation, and a widening of
plant tissue and soil C:N ratios. These changes in biotic and edaphic factors
encountered along the chronosequence appear to be related to the termination of tillage and fertilizer inputs, and especially to the associated increases in plant biomass (both above and belowground), all associated with
the cessation of agriculture.

10

Abstracts

ALONSO, CONCHITA and CARLOS M. HERRERA. Estacion Biologica

de Donana, CSIC., Sevilla, Sevilla, Spain. Gender-specific effects of the
solar irradiance environment on pollination success of a winter-flowering gynodioecious shrub.
Daphne laureola L. (Thymelaeaceae) is a gynodioecious, winter-flowering
shrub inhabiting the understory of mixed montane forests in southeastern
Spain. The small, inconspicuous green flowers are pollinated by ectothermic insects whose activity is mainly confined to infrequent spells of sunny
weather. We tested the hypotheses that (1) the solar irradiance environment
of individual D. laureola plants should influence their pollination success,
and (2) the effect of solar irradiance on pollination success should be gender-dependent, given the difference between male-sterile and hermaphrodite flowers in conspicuousness and floral rewards. We estimated pollination success for female function (mean number of pollen tubes/flower and
proportion of unfertilized flowers) and assessed the irradiance environment
(using hemispherical fisheye canopy photographs) for individual D. laureola plants in five different populations. Hermaphrodite flowers were pollinated more often, and received threefold more pollen tubes/flower than
male-steriles, in all the populations. There existed a highly significant interaction effect of gender 3 canopy cover on pollination success (F1,50 5
9.75, P 5 0.003; F1,50 5 5.61, P 5 0.02 for pollen tubes/flower and unfertilized flowers, respectively). Among hermaphrodite individuals, the
number of pollen tubes/flower increased and the proportion of unfertilized
flowers decreased from closed to more open sites, while male-steriles
showed the opposite trends. The contrasting sign of the relationship between cover and pollination success for the two genders is expected to
generate a fine-scale, within-habitat mosaic in the magnitude of the pollination advantage of hermaphrodite individuals, which would be greater in
the more open, sunny locations where ectothermic pollinators should be
more active.
ALTIZER, SONIA* and ANDREW K. DAVIS. Emory University, Atlanta,
GA. Individual infection risk and epidemics of mycoplasmal conjunctivitis in wild House Finches.
Mycoplasmal conjunctivitis is a recently emerged bacterial disease affecting wild House Finches (Carpodacus mexicanus) in eastern North America.
This novel strain of a poultry pathogen has caused dramatic reductions in
host abundance, and follows highly seasonal epidemics with rapid increases
in prevalence during the fall and winter months. We examined the epidemiology of conjunctivitis in a local population of House Finches in Atlanta,
GA, a region where this pathogen reaches unusually high prevalence from
August through November. To understand what factors affect changes in
host susceptibility and the timing of outbreaks, we monitored monthly
prevalence of conjunctivitis among wild-captured House Finches within a
20 km radius of Emory University over two successive years. We examined
individual traits associated with high infection risk (including age, sex,
molting status, condition, and co-infection with other parasites), and also
compared how the timing of epidemics covaried with seasonal changes in
flock sizes. Our results showed that during outbreaks, infection risk was
significantly higher among juveniles than adult birds, suggesting that annual pulses of juvenile recruitment may affect the timing and severity of
epidemics. Infection risk also depended on sex and molting status, although
the directions of these associations ran counter to our predictions. House
Finches infected with Mycoplasma were in poorer condition, had elevated
counts of two types of WBCs, and were more likely to harbor hemoparasites and feather mites than uninfected birds. Collectively, these results
point to several factors that are likely triggers of rapid changes in prevalence in this and other host-pathogen systems, including seasonal shifts in
age structure and host social behavior.
ALTWEGG, RES,* KERRY B. MARCHINKO and BRADLEY R. ANHOLT. University of Victoria, Victoria, BC, Canada. Inducible morphological defense and community dynamics in protists.
Inducible defenses are expressed according to current risk situation and
incur a fitness cost that may decrease competitive ability. Thus, predators
affect food webs both directly, through consumption, and indirectly by
changing the interaction strength between their prey and other community
members. Depending on which of these effects prevail, the dynamics of

communities are predicted to be quite different. In response to live or dead

predators, the hypotrich ciliate Euplotes aediculatus develops lateral projections that effectively protect it from being eaten by the gape limited
predator Stenostomum sphagnetorum, at the cost of slower reproduction.
Paramecium aurelia shares resources with Euplotes, but does not alter its
morphology in response to predators. In an attempt to disentangle direct
from indirect effects, we exposed Euplotes, alone or together with Paramecium, to either live, dead or no predators and followed the dynamics for
twenty generations. The morphological defense of Euplotes varied over
time and loosely tracked predator density. Simple models predict destabilizing lagged effects on population dynamics for such systems. However,
population dynamics of Euplotes, characterized by the coefficient of variation in population size, was not affected by the predator treatment in any
detectable way. Paramecium depressed the population growth rate of Euplotes, and the effect was stronger when Euplotes expressed defenses in
response to dead predators. Thus, we present a system in which inducible
morphological defenses affect community processes. Its experimental tractability and the short generation time may make this system an important
tool to bridge the gap between theory and empirical evidence and significantly improve our understanding of community dynamics.
AMMANN, REBECCA L.* and DENNIS W. NYBERG. University of Illinois-Chicago, Chicago, IL. Seeded prairies are taller and grassier than
original Illinois prairies.
Vegetation height structure was studied in six original Illinois tallgrass
prairies and eight reconstructions, two planted (RP) and six seeded (RS).
Robel pole visual obstruction (VO), species contributing to VO, stem density at 0.5, 1.0 and 1.5 m and the height of the tallest individual stem near
the pole were measured at 15 points within each prairie. The mean VO of
original prairies was 0.4660.13 m, significantly shorter than the RP,
0.5460.16 m, and RS, 0.6760.18 m. The number of forb species contributing to VO was significantly greater in original prairies (14) than in RS
prairies (9). At 0.5 m all prairies had about 160 stems m-2 with a 3:1 grass
to forb ratio. At 1.0 m original prairies dropped to 19 total stems m-2 and
RP to 13 m-2, both significantly less than the 41 m-2 of RS, with 73, 42
and 87% as grasses, respectively. At 1.5 m there were only 0.2-0.4 forb
stems m-2, but grass had much higher values, 6.3, 1.5 and 16.6 stems m-2,
for original, RP and RS, respectively, with RS significantly greater than
the others. The height of the tallest nearby individual (grand mean 1.33 m)
did not differ significantly among types, but the range in original prairies
(0.99,1.75) was much less than in RS (0.67,2.01) because two RS sites
were dominated by short non-native species. When Andropogon gerardii
was the nearby tall individual, 35% of the cases, its height was significantly
greater on RS than original prairies. The taller stature of reconstructed
prairies appears to be due to both greater heights of individuals as well as
species composition differences. Restoration efforts through seeding have
not yet mimicked vegetation height structure of original prairies, which is
important for animal usage.
ANAND, MADHUR and BRIAN C. TUCKER.* Department of Biology,
Laurentian University, Sudbury, Ontario, Canada. Matrix modelling of
natural and pollution-induced forest vegetation gradients.
We sought to compare the efficacy of the stationary Markov model and
conventional ordination techniques in describing compositional and structural changes in forest vascular plant communities along natural and manmade spatial gradients at two scales, local and regional. Stationary Markov
models have been used extensively in ecology to study community change
with time but estimation of transition matrix probabilities in the absence
of individual-based data has been problematic. We applied a method of
estimating species transitions for the Markov model put forward to deal
specifically with coenosere ecological data. Vegetation abundance (understory percent cover, overstory frequency) and structure (overstory height)
data are from six sites spanning a pollution gradient in the Great LakesSt. Lawrence forests near Sudbury, Ontario, Canada. The manmade gradient varies from barren locations near smelter stacks to diverse, minimally
impacted forests 40 km away. At each site, parallel transects extend through
suspected natural spatial gradients down south-facing slopes where topographic factors were thought to affect plant communities. Both ordination
techniques and the Markov analyses detected the strong regional pollution-

induced gradients in abundance and structure. Ordination did not detect

slope-related local gradients despite the general trend that, as pollutionlevel decreases along the regional gradient, vegetation along the slopes
begins to display Markovian spatial dynamics. This is due to information
loss resulting from static ordination analyses: information regarding transitions between observations along the natural ordering of quadrats is not
maintained.
ANDERSON, DEAN P.,1,* THOMAS P. ROONEY,1 MONICA G. TURNER,1 DONALD M. WALLER,1 ADRIAN P. WYDEVEN,2 JANE E. WIEDENHOEFT,2 DEAN E. BEYER3 and WILLIAM S. ALVERSON.4 1 University of Wisconsin-Madison, Madison, WI, USA; 2 Wisconsin Department of Natural Resources, Park Falls, WI, USA; 3 Michigan Department
of Natural Resources, Marquette, MI, USA; 4 The Field Museum, Chicago,
IL, USA. Tri-trophic interactions of wolves, deer, and plant communities in the Northern Wisconsin and Michigan.
The effect of high-density deer populations on plant communities has been
well documented. However, few studies have examined whether top predators may mitigate this effect spatially. We used data from Northern Wisconsin and Michigan to address the following questions: (1) is there an
inverse relationship between deer (Odocoileus virginianus) and wolf (Canus lupus) densities across the landscape?; (2) does species richness of forbs
vary as a function of the density of wolves in the area?; and (3) does the
biomass of grasses, forbs, and shrubs vary with wolf density? Radio-telemetry and ground-tracking data of wolves were used to identify locations of
wolf packs. GIS analysis was used to determine the center point of wolf
territories, and the inverse of the distance from the center point was used
as an indicator of wolf density. Deer fecal-pellet surveys were conducted
at 46 locations to estimate deer densities relative to wolf pack locations.
Vegetation was sampled in the Chequamegon National Forest, Wisconsin
to quantify the species richness of forbs, and the biomass of forbs, graminoids and shrubs. Results indicate a significant inverse relationship between deer and wolf densities. In coniferous forests and cedar swamps,
habitats used extensively by deer, forb species richness had a hump-shaped
relationship with wolf density, indicating a potential intermediate-disturbance-hypothesis mechanism. In cedar swamps, biomass of all forb species
and shrub species that experience high rates of deer browsing was positively related to wolf densities. In contrast, graminoid biomass was negatively related to wolf densities. These results suggest that wolves mitigate
spatially the effect of an over-abundant deer population on vegetation communities.
ANDERSON, KRISTINA J.* University of New Mexico, Biology department, Albuquerque, NM. Latitudinal trends in energy expenditure of
endotherms.
Energy expenditure (field metabolic rates) of endotherms scales allometrically with body mass, but the remaining variation is not well understood.
The purpose of this study is to examine how energy expenditure varies
with environmental factors and species characteristics after accounting for
mass. When plotted against latitude, mass-corrected field metabolic rates
of approximately 200 bird and mammal species fall within a constraint
triangle consisting of a flat upper bound and a lower bound that increases
with latitude. Thus, endotherms are constrained to increasing energy expenditure (relative to body size) with increasing latitude, but are free to
vary between this constraint and an upper limit of sustainable energy expenditure. An examination of some correlates of latitude suggests that the
lower bound of this relationship is controlled by temperature and that ecosystem productivity does not predict energy expenditure. Species characteristics such as diet and life history play a role in determining where an
animal falls within the constraint triangle. These macroecological patterns
provide insight toward an integrated, theoretical understanding of energy
expenditure of endotherms.
ANDERSON, KURT E.,1,* ROGER M. NISBET1 and SEBASTIAN
DIEHL.2 1 Department of Ecology, Evolution, and Marine Biology, University of California, Santa Barbara, Santa Barbara, CA; 2 Department fur
Biologie II, Ludwig-Maximilians-Universitat, Munchen, Bavaria, Germany.
The spatial length scale of the effects of environmental variability on
spatial population dynamics in systems with unidirectional dispersal.
In environments where organisms passively disperse in a unidirectionally
biased media, such as in streams and along coastlines, dispersal can play

Abstracts

11

a large role in linking upstream to downstream habitats. The effects of

localized environmental variability, for example, increased light input
into a stream due to gaps in shoreline vegetation can propagate rapidly
downstream, affecting the dynamics of and interactions between downstream populations. We will present a spatial model of a stream community
that consists of a sessile algal producer and an aquatic insect grazer that
disperses passively downstream in the water column. We have developed
a simple technique based on Fourier analysis which allows us to predict
how far downstream localized environmental variability defined in our
model as local changes in parameter values will cause changes in downstream algae and grazer abundances. This downstream length of effect we
define as the spatial length scale of the impact of the variability. Values
for model parameters were obtained from observations of algal periphyton
and the mayfly larva Baetis tricaudatus in an eastern Californian stream.
The model predicted that environments with high average per capita baetid
dispersal rates, such as in areas with high flow and low algal biomass
growth, exhibit longer spatial length scales than those with low baetid per
capita dispersal. These predictions of our parameterized model matched the
observed spatial scale of population responses, and, in turn, the degree of
spatial patterning, by Baetis to both locally isolated as well as spatially
extensive and irregular environmental disturbances in current speed and
algal primary production. We suggest that our model can be altered and
applied to many aquatic and marine systems.
ANDERSON, LYNN L.,* FENG SHENG HU and KEN N. PAIGE. University of Illinois, Champaign/Urbana, IL, USA. Using morphology to
infer environmental conditions: Genetics, environment, or both?
Recent studies have used morphological traits of plant fossils to reconstruct
past environments. This approach is compromised if morphological features
of different individuals of the same species vary with genetic history. We
measured a suite of morphological traits of extant needles collected from
several populations for both black spruce (Picea mariana) and white spruce
(P. glauca) across northwestern North America. Using discriminant function analysis, we found that 98% of plants correctly classified back to their
respective species designation. More important, withinspecies analyses
showed that plants correctly classified back to their populations of origin
89% of the time. The underpinnings of such variation could be populational
genetic variation (G), environmental induced plasticity (E), or G X E interaction. To assess the relative importance of these factors, we compared
needle morphological data from four natural whitespruce populations and
four populations of the same geographic origins grown in a common garden
experiment. Results revealed that 84% of plants correctly classified back
to their common garden populations, not to their natural areas of origin,
supporting G X E interaction. These results underscore the essential role
of genetic history in plantmorphological response to changing environmental conditions. Because morphological traits of spruce needles are dictated by G X E interaction, it is problematic to use them to infer past
environmental conditions.
ANDERSON, M. REBECCA1,* and SABINE S. LOEW.2 1 banderso@
ilstu.edu, Illinois State University, Normal, IL; 2 ssloew@ilstu.edu, Illinois
State University, Normal, IL. Low allozyme diversity in small populations of a commercially valuable plant, American ginseng.
American ginseng, Panax quinquefolius L., is an herbaceous species of
highly fragmented forest habitat in eastern North America that is harvested
from small wild populations for export, primarily to Asian markets. In
general, small populations (, 500 individuals) of species exhibit lower
genetic diversity than larger ones (1000 to 10,000 individuals), which may
reduce their evolutionary capability to adapt to environmental change. We
tested the hypothesis that genetic variation is positively correlated with
population size. Neutral genetic diversity was assessed using electrophoretic allozyme variation in individuals selected from seven Illinois populations of American ginseng that vary in size from fewer than 10 to several
hundred individuals. Allozymes are codominant markers inherited in Mendelian fashion that are frequently used to detect population differentiation.
A large body of literature exists for comparison that describes allozyme
variation in numerous species characterized by a variety of reproductive
modes and threat status. We used cellulose acetate gel membrane electrophoresis to separate and visualize allozyme variants. We analyzed up to 30

12

Abstracts

individuals from each of the seven populations for eight allozymes representing five enzyme systems. As a high proportion of those individuals
were homozygous for the same allele, most populations analyzed so far are
monomorphic. Two possible reasons explain these results: (1) these markers are not sufficiently sensitive to reveal differences among populations,
or (2) significant loss of genetic diversity has already occurred due to small
size and isolation of populations and long history of harvest pressure. In
order to determine the more likely explanation we will expand our genetic
analysis to include more variable molecular markers, such as AFLPs.
ANDERSON, SEAN S.* Center for Conservation Biology, Department of
Biological Sciences, Stanford University, Stanford, CA, USA. Incorporating adaptive management into restoration design: lessons from California grasslands.
The past decade has seen a marked proliferation of "adaptive management"
in the design and monitoring of restoration projects. Typically, "adaptive
management" has been an afterthought, often not central to project design,
and frequently the first casualty of logistic or budgetary constraints. To
demonstrate the value of active adaptive management, we have developed
a phased, experimental approach to restoring degraded communities wherein the results from previous phases inform the design and implementation
of subsequent phases. Both examples are from a 300 ha restoration of oak
savanna in the coastal foothills of the San Francisco Bay Area. The first
project is an attempt to modify soil seed banks to restore native grasslands.
The second project is an effort to augment amphibian populations with a
series of seasonal wetland breeding sites. By beginning at small spatial and
temporal scales, both projects have been able to rapidly incorporate sitespecific results into the design of subsequent restoration phases and dramatically increase the likelihood of a successful restoration.
ANDERSON, WENDY B.,1,* CORY BETHMANN,1 ALEXANDER
WAIT2 and LESLIE ALLFREE.3 1 Drury University, Springfield, MO,
USA; 2 Southwest Missouri State University, Springfield, MO, USA; 3 National Center for Ecological Analysis and Synthesis, Santa Barbara, CA,
USA. Boundary permeability to marine subsidies affects arthropod
community structure on Gulf of California islands.
Cross-ecosystem flux of energy, nutrients, detritus, and prey can have substantial impacts on population and community dynamics in recipient systems. Such fluxes may be inhibited or facilitated by the physical structure
of the interface between the two systems. Differential permeability of ecosystem boundaries to mobile resources may partially explain distribution
and abundance of species in the recipient systems. We assessed the arthropod community composition and relative contribution of marine-derived
resources in their diets along 16 shore-to-inland transects established from
beach- or cliff-type shorelines on desert islands in the Gulf of California.
Total density of arthropods was highest along transects originating from
low-angled beaches, which receive much marine macroalgae. Amphipods
and spiders dominated these communities, with high densities maintained
up to 25 m from the shore, whence they declined to typical inland densities.
Beetles and ants became relatively more common further inland. Amphipods and spiders exhibited enriched 13C (indicative of marine-derived diets), but these signatures tapered off in individuals trapped further inland.
In communities along transects originating at high-angled cliff shorelines
inhabited by seabirds, total densities were lower than that on the beach
transects, and the communities were dominated by beetles and ants. All
guilds exhibited high 15N values, which indicate either direct or indirect
consumption of seabird products. Neither density nor diet patterns varied
along the 50m transects on bird inhabited cliffs. On transects from cliffs
not inhabited by seabirds, total densities were lowest, and communities
were dominated by amphipods and beetles in the first 2 meters, and by
ants further inland. 13C and 15N isotopes indicated very little consumption
of marine-derived resources. These results suggest that communities living
on islands or other isolated habitats, or those living near the interface of
two ecosystems may be distributed unevenly in response to differential
permeability of ecosystem boundaries to allochthonous resource subsidies.

ANDREWS, JEFFREY S.,* MICHAEL J. VANNI and WILLIAM H.

RENWICK. Miami University, Oxford, Ohio, USA. The influence of
storm subsidies on nutrient and light limitation of reservoir phytoplankton growth.

ANDRUSKIW, MARK C.,1,* JOHN M. FRYXELL1 and IAN D. THOMPSON.2 1 University of Guelph, Guelph, Ontario, Canada; 2 Canadian Forest
Service, Sault Ste. Marie, Ontario, Canada. Prey abundance, availability,
and anxiety in logged and unlogged boreal forest.

Phytoplankton in most lakes are nutrient-limited during summer. During

storms, reservoirs erratically receive pulses of nutrients and sediment from
their watersheds. These pulses may alleviate nutrient-limitation by providing nutrient subsidies and by inducing light limitation. However, the dynamics of nutrient- and light-limitation in relation to storm events have not
been rigorously quantified. The goals of this study were to (1) determine
the magnitude of nutrient-limitation before, during, and after storm events;
(2) evaluate the time scale over which storms alleviate nutrient limitation
and over which nutrient-limitation returns; and (3) determine if storms
alleviate nutrient limitation by providing excess nutrients, inducing lightlimitation, or both. Nutrient and light-limitation were evaluated in spring
and summer 2002 with a series of algal growth bioassays that utilized
varied nutrient combinations (control, N, P, N+P) and light intensities (200,
100, 50, 25 umol m2/sec) on natural phytoplankton assemblages from Acton Lake (SW Ohio, USA), a reservoir in an agricultural watershed. We
found that phytoplankton were P-limited during relatively calm periods,
and that storms alleviated P-limitation. During storms, phytoplankton became light-limited. In addition, during large storms phytoplankton were not
P-limited even when grown at saturating light intensities, showing that
storm events deliver quantities of nutrients sufficient to saturate growth.
Phytoplankton became nutrient-limited within a few days after a storm
subsided, and the magnitude of nutrient limitation was negatively correlated
with discharge into the lake over preceding time periods. Maximal correlations occurred with discharge over the preceding 10-14 days; correlations
over shorter and longer time scales were not as strong. These results suggest that phytoplankton are light-limited due to storm-induced turbidity and
then shift to nutrient limitation as storm inputs subside, with severity of
nutrient limitation being linked to flow over a time period equal to several
phytoplankton generations.

Most studies of prey vulnerability come from laboratory aquatic systems.

The common result is reduced vulnerability in structurally complex environments due to impeded predator search and physical refuge for prey.
Here we present a terrestrial predator-prey system with the opposite result.
We tested the hypothesis that American marten suffer reduced foraging
efficiency in structurally simpler forests generated by clearcut logging.
From fall 2001 to spring 2002 marten foraging dynamics were studied in
logged and unlogged boreal forest of northwestern Ontario. Coarse woody
debris and availability of subnivean access were censused along linear transects. Small mammals were censused by live trapping. Marten hunting
behavior was examined from snowtracking 34 marten over 100km, and
diets were quantified by scat analysis. Behavioral response of voles to
predation risk was measured with giving-up densities. While coarse woody
debris levels were higher in unlogged forest, marten selected subnivean
access in both forest types. Despite equal small mammal densities, marten
encountered and killed prey twice as often in unlogged forest. Giving-up
densities of voles were higher in unlogged forest indicating greater fear of
predation risk. We conclude that, rather than providing refuge for small
mammals, forest structure creates an aggregated and predictably located
resource for marten. Second-growth forest therefore represents reduced
hunting efficiency. We are calculating prey preferences to test the hypothesis that marten use alternate prey in logged forest where small mammals
are less vulnerable.

ANTEN, NIELS P.1,2,* 1 Dept. Plant Ecology, Utrecht University, P.O.Box

800.84, Utrecht, Utrecht, The Netherlands; 2 Chair Group of Plant Production Systems, Wageningen University, Haarweg 333 6709 RZ, Wageningen,
Gelderland, The Netherlands. Costs and benefits of mechanical stability
in competing and non-competing plants.

H
D

A
W

We examined the fitness costs and benefits of mechanical hardening in

plants. Tobacco (Nicotiana tabacum) plants were flexed 40 times each day
(flexed plants) or not flexed (control plants) and grown either in isolation
or in a dense Tobacco stand where flexed and control plants were mixed.
Compared to control plants, flexed plants produced shorter and thicker
stems, which were more flexible (lower elastic modulus) but had equal
maximum break stress. It was estimated that as a result, they would be
able to resist 40-70% greater lateral forces (Fmax) and that they had 60 to
500% greater buckling safety factors. For all plants Fmax values exceeded
wind forces measured on whole plants in a wind tunnel at wind speeds of

IT

Current soil testing, usually of chemical fertility interpreted on the basis of

crop nutrient requirements, provides an imcomplete picture of soil function.
Meaningful information for agroecosystem management could be gained
from a tool that suggests chemical, biological and physical indicators, offers site-specific interpretations of those indicators, and provides an overall
assessment of those interpretations in relation to soil function. Designing
this assessment tool as a framework allows researchers to continually update and refine the interpretations for many soils, climates, and land use
practices. We designed and assessed a framework consisting of three steps:
indicator selection, indicator interpretation, and integration into an index.
We tested transferability of the framework using data from four case studies
in GA, IA, CA, and the Pacific Northwest (NRI) that differed in climate,
management, spatial extent, and soil type. Applying decision rules in the
selection step successfully identified indicators that were present in the
existing data sets. The interpretation step resulted in site-specific and scientifically defensible differences in indicator scores. We found four main
patterns of indicator results, including significantly different scores when
no difference was detected in the measured data (before scoring). As a
check of the efficacy of this framework approach, we performed stepwise
regressions using the scored (and observed) indicators as independent variables and endpoint data as iterative dependent variables for each case
study. The scored indicators usually had coefficients of determination (R2s)
that were similar or greater than those of the observed indicator values.
The R2s between indicators and endpoints were higher when examined for
one treatment at a time rather than the entire data set. For instance, when
using all NRI data, there was no relationship between indicators and nematode diversity but when only the data for no-tilled Xerolls cropped to
continuous small grains were examined alone, the R2 was 0.92. The results
of this study show significant progress toward development of an assessment framework for adaptive management that is transferable to a variety
of climates, soil types, land uses and management systems

Although species distribution limits are universal, little is known about the
evolutionary constraints that hinder range expansion. To examine the factors creating and maintaining range limits, I am studying the altitude ranges
of sister species of monkeyflowers, Mimulus lewisii and M. cardinalis,
across their elevation ranges in the Sierra Nevada Mountains of California.
Although both species occur in disturbed, moist riparian habitats, they differ in their altitude distributions; M. cardinalis is found from 0 to 1500 m
whereas M. lewisii occurs from 1400 to 3000 m. I am using a combination
of reciprocal transplants and demographic census plots to determine whether populations are maladapted at range boundaries. Both species survive
and reproduce better in transplant gardens within their native distributions
than beyond their current elevation limits. However, within each species,
populations from different elevations do not differ predictably in garden
performance across the altitude gradient. Population matrix analysis of census plot data reveals asymptotic population growth rates that vary with
elevation and year. To isolate the traits and environmental factors that affect
survival and reproduction at range limits, I am using growth chambers with
varying temperature and UV-B light regimes. These controlled environment
experiments demonstrate the importance of high and low temperature tolerance and UV-B tolerance to performance at different elevations. These
results will be used to study selective constraints at range boundaries.

ANDREWS, SUSAN S.1,* and DOUGLAS L. KARLEN.2 1 USDA-NRCS

Soil Quality Institute, 2150 Pammel Drive, Ames, IA, 50011; 2 USDA-ARS
National Soil Tilth Lab, 2150 Pammel Drive, Ames, IA, 50011. A framework for soil management assessment: Four case studies.

ANGERT, AMY L.* Michigan State University, East Lansing, MI. Adaptation at geographic range limits of monkeyflowers (Mimulus lewisii
and M. cardinalis).

Abstracts

13

up to 13 m s-1. For isolated plants the increase in stability had no consequences for either growth or seed production. By contrast, in the dense
stand flexed plants were progressively shaded by control plants and had
considerably lower rates of seed production and survival. These results
show that the fitness costs of mechanical stability increase strongly when
plants compete for light and suggest that a partial suppression of thigmomorphogenesis (growth response to mechanical disturbance) by the shade
avoidance response would be a beneficial trait.
ANTHONY, NICOLA M.,1,* STEPHEN CLIFFORD,2 MIREILLE JOHNSON-BAWE,2 KATE ABERNETHY,2,3 CAROLINE E.G. TUTIN,3 LEE J.
WHITE,4 JEAN E. WICKINGS2 and MICHAEL BRUFORD.5 1 University
of New Orleans, New Orleans, LA 70148, USA; 2 Centre International de
Recherches Medicales, Franceville, Gabon; 3 University of Stirling, Stirling, U.K.; 4 Wildlife Conservation Society, New York, NY, U.S.A.; 5 Cardiff University, Wales, Cardiff, Wales, U.K. Mitochondrial phylogeography of western lowland gorillas.
Gabon is an important centre of biodiversity and harbors the largest remaining populations of western lowland gorillas (Gorilla gorilla gorilla).
Understanding population genetic structure and patterns of gene flow at
different spatial scales is essential to the formulation of an effective longterm management strategy. We surveyed mitochondrial DNA variation in
hair and faecal samples collected from wild gorillas across much of their
current range. Preliminary results indicate high levels of genetic diversity
and considerable phylogeographic structure within western gorillas. Phylogenetic analysis recapitulate the major evolutionary split between eastern
and western gorillas and indicate two major haplogroups within western
gorillas. One group is made up of sequences from Nigeria and Cameroon.
The other is comprised of populations from the remaining western gorilla
range and includes three discrete subgroups of sequences from Gabon and
adjacent Congo, Equatorial Guinea and Central African Republic. The geographic distribution of DNA variation suggest that Pleistocene forest refugia may have played an important role in shaping patterns of genetic
structure within western lowland gorillas.
ANTONSEN, HILDE,* ROGER C. ANDERSON and STEVEN A. JULIANO. Illinois State University, Normal, Illinois, United States. Effects
of supplemental nutrients on Schizachyrium scoparium (little bluestem)
with high and low AM colonization.
An experimental field study was conducted to evaluate the impacts of supplemental nutrients (N, P, K and Ca+Mg) on little bluestem grass (Schizachyrium scoparium) with a high (21.0 6 0.7%) or low (3.2 6 0.4%)
levels of colonization by arbuscular mycorrhizal fungi (AMF). Low-colonized plants were grown in autoclaved soil in a glasshouse and transplanted
into fumigated soil in the field, while high-colonized plants were started in
unsterilized soil and planted into unfumigated field soil. Originally, data
were analyzed by multiple univariate analysis of variance. Previous conclusions drawn from these analyses were that bases (Ca+Mg) were the
limiting nutrients in this sandy prairie system. Since tools for performing
multivariate statistics are now more accessible and userfriendly, a reanalysis of the data was performed. New interpretations suggest that phosphorus and bases were limited nutrients in this system, and enhanced little
bluestem growth occurred when mycorrhizal colonization was low. However, none of the added nutrients had significant effects on growth for plants
with high levels of colonization. This interaction is explained by the cost
of having a fungal partner. Even if levels of P and Ca+Mg are high enough
to sustain increased growth, available carbon sources are not sufficient for
increased growth due to the drain of the fungal component. Thus, in this
sandy prairie system the AMF acts as a parasitic organism.
APPLE, JENNIFER L.,1,* JOHN G. BISHOP1 and WILLIAM F. FAGAN.2
Washington State University-Vancouver, Vancouver, WA; 2 University of
Maryland, College Park, MD. Successionally driven changes in leaf nutrients and spatial patterning of herbivory on lupines at Mount St.
Helens.
1

Previous work demonstrated that several lepidopteran herbivores of lupines

(Lupinus lepidus) in primary successional habitat on Mount St. Helens
exhibit similar distributions with respect to host plant density, with damage

14

Abstracts

concentrated at the sparse margins of high-density (core) patches or in

lower-density (edge) patches. We focus on one of these herbivores, the
cutworm Euxoa sp. (Noctuidae), to determine whether host plant tissue
quality is responsible for the observed negative density-dependent pattern
of herbivory. Larvae hatched from eggs laid by field-caught females were
reared on leaves collected from either core or edge lupine patches. Caterpillars fed leaves from core plants were significantly delayed in their development: after 8 weeks, they were about 33% smaller than larvae fed
edge tissue. Eventually, larvae on core diets achieved comparable maximum weights as larvae on edge diets. Euxoa larvae also exhibited fourfold higher mortality on diets of core leaves during the first 2 weeks of
development. Comparisons of C, N, and P content of the core vs. edge
leaves comprising the experimental caterpillars diets revealed that edge
plants provide a higher quality food source for larvae, with elevated %P
and lower C:P and N:P elemental ratios. In addition, we found that the
nutrient content of both core and edge leaves declined significantly over
the course of the summer. These results suggest that the nutritional content
of plants available to early instar Euxoa larvae is critical in determining
their growth trajectory and survivorship, especially since late instar larvae
will be subject to deteriorating food quality. Thus, successionally driven
bottom-up processes that produce density-dependent variation in host plant
quality are likely responsible for the spatial distribution of Euxoa at Mount
St. Helens.
APPLE, MARTHA E.,2 VICKIE L. SMITH-LONGOZO,1,* RENEE RICHARDS1 and ROBERT S. NOWAK.1 1 University of Nevada, Reno, Environmental and Resource Sciences, Reno, NV, 89557; 2 University of Central
Washington, Biological Sciences, Ellensburg, WA, USA. Root respiration
and elevated CO2 in Mojave Desert shrubs at the Nevada Desert FACE
Facility in drought and rainy years.
In a long term study of root physiology in response to elevated CO2 at the
Nevada Desert FACE (Free Air Carbon Enrichment) Facility (NDFF) in
the Mojave Desert, (where CO2 levels are 550 ppm in three plots and
ambient at three others), we previously found that root respiration and
specific root length (m/g dry weight) were significantly greater with elevated CO2 in two co-dominant native shrubs creosote bush, Larrea tridentata, and white bursage, Ambrosia dumosa. Therefore, both physiology and
structure are influenced by elevated CO2 in these two desert shrubs. However, root growth as observed through root observation boxes was minimal
during the drought year of 2001-02. In contrast, 2002-03 is predicted, and
to date is yielding greater quantities of precipitation. Thus, we are again
measuring root respiration and specific root length as well as branching
patterns and nitrate and ammonium uptake by fine roots in these two shrubs
with the goal of comparing the effects of variations in precipitation and
elevated CO2 on root physiology and carbon use as part of this on-going
investigation of the possible effects of climate change. Greenhouse experiments compliment our field study: In a 2x2 factorial experiment on L.
tridentata with the following treatments: ambient (360 ppm) or elevated
(550 ppm) CO2, and addition of 2 or 20 ppm phosphorus (n 5 10 plants/
treatment), we found that 10-month-old L. tridentata plants had a significantly larger root: shoot ratio at elevated CO2 than at ambient CO2. Interestingly, although the root systems had greater dry weights at elevated CO2,
the ratio of fine: coarse roots were similar between the CO2 levels. Analyses
of minirhizotron images, respiration rates, specific root length, uptake of
nitrate and ammonium in fine roots labeled with stable isotopes, and elemental analyses for carbon, hydrogen, and nitrogen will be instrumental in
determining how these desert shrubs allocate carbon in response to elevated
CO2.
AQUILINO, KRISTIN M.,* BRADLEY J. CARDINALE and ANTHONY
R. IVES. University of Wisconsin-Madison, Madison, Wisconsin, USA.
Predator diversity and the biological control of agricultural pests: A
test of the host-plant heterogeneity hypothesis.
Recent theoretical and experimental evidence suggests that consumer diversity may influence how efficiently limited resources are captured
through resource partitioning and/or changes in species interactions. Resource partitioning and species interactions are known to be sensitive to
habitat diversity; yet few studies have examined the potential for diversity
effects of consumers on resources to change between homogeneous and

heterogeneous environments. We used a dual replacement series experimental design to simultaneously manipulate the species richness of predators (consumer diversity - Harmonia axyridis, Coleomagilla maculata, and
Nabis) of the pea aphid Acyrthosiphon pisum (the resource) in laboratory
microcosms having one or three host plant species (habitat heterogeneity Medicago sativa, Trifolium pratense, and Vicia faba). We then compared
how predator richness influenced aphid consumption in plant monocultures
and polycultures. We found no interaction between predator and host plant
richness on the consumption of aphids (F1,100 5 0.01, P50.92). There were,
however, main effects of both predator diversity and habitat heterogeneity
treatments. Consumption of aphids was significantly higher (by 21%) in
microcosms containing three predator species versus those having an equal
density of just one predator species (F1,100 5 4.46, P50.04). This was due
to either reduced intraspecific competition in the more diverse assemblages
and/or interspecific facilitation between predator species. The consumption
of aphids was also significantly reduced in plant polycultures (by 31%)
compared to monocultures having equal plant density (F1,100 5 4.60,
P50.03). These results were driven by a reduction in predator efficiency
on the host plant Vicia faba, and were potentially exacerbated by aphids
moving to Vicia faba in polycultures. Our study demonstrates that diversity
at multiple trophic levels may simultaneously impact the capture of limiting
resources. Furthermore, in our study, the diversity of trophic levels acted
independently as some theory has predicted.
ARCHER, STEVEN R.1 and THOMAS W. BOUTTON.2 1 School of Renewable Natural Resources, Biological Sciences East 325, University of
Arizona, Tucson, AZ, USA; 2 Rangeland Ecology & Management, Texas
A&M University, College Station, TX, USA. Wooded grasslands: functional consequences of an ecological oxymoron.
Changes in vegetation cover in drylands have the potential to modify atmospheric chemistry and climate processes. Proliferation of woody vegetation in grasslands and savannas over the past century represents a class
of land cover change widespread in the world9s rangelands. In some instances, this vegetation change results in ecosystem simplification, as a
single woody species, either native or exotic, may come to dominate landscape function; in other instances biodiversity may be unaffected or even
enhanced. Although woody plant proliferation in rangelands has long been
a concern of land managers, research has focused primarily on its impacts
on grass production and approaches for reducing the abundance of established trees and shrubs. Little is known of the ramifications of this vegetation change for ecosystem function, atmospheric chemistry and climate
processes. Studies from various bioclimatic zones suggest a broad range of
potential responses. Results from a case study in southern Texas will be
used to illustrate how historical increases in woody plant abundance have
altered C and N storage, dynamics, and trace gas fluxes in ways that can
affect tropospheric chemistry and greenhouse gas concentrations. With
greenhouse gas mitigation and carbon sequestration issues gaining national
and international prominence, biogeochemical, land surface-atmosphere
perspectives suggests the need for new approaches for evaluating the functional role of woody plants in rangelands. A carbon-accounting perspective
suggests cost/benefit trade-offs that have not been traditionally considered
in managing vegetation on rangelands. It will be argued that perverse incentives may develop whereby societies may become increasingly willing
to tolerate or even subsidize land use practices that promote proliferation
of woody plants historically regarded as undesirable. Such scenarios would
have an adverse affect on the conservation of grassland and savanna plants,
animals and ecosystems.
ARMITAGE, ANNA R.* and PEGGY FONG. University of California Los
Angeles, Los Angeles, CA. Trophic-level effects of habitat-modifying
crabs in muddy and sandy tidal flats.
We evaluated the influences of an omnivorous decapod on habitat characteristics and trophic structure in a tidal mudflat and a sandflat in southern
California. In enclosures that contained subadult and adult herbivorous
snails (Cerithidea californica), we added or excluded crabs (Pachygrapsus
crassipes). Crab predation on the snails was intense, with up to 90% consumption of the large snails in the sandflat and 75% consumption in the
mudflat, but there was no cascading effect detectable on the benthic microalgal community. In fact, crab burrowing activities decreased benthic

chlorophyll a concentration, particularly in the sandflat, suggesting a reduction in benthic microalgal availability for the snails. Crabs altered sediment characteristics by unexpectedly increasing anoxia and, in the sandflat,
reducing tensile shear strength. Crab bioturbation also increased nutrient
flux out of the plots. In the mudflat, crabs decreased small snail growth
rates, possibly via alterations of habitat characteristics or by inducing snail
burial or burrowing as an escape behavior. The wide range of effects that
crabs exerted on the community included direct consumption and prey
behavioral alterations, indirect reduction of primary producer biomass, and
ecosystem-level impacts on sediment characteristics and nutrient cycling.
However, the strengths of each of these interactions varied between habitat
types.

ARMSWORTH, PAUL R.* and JOAN E. ROUGHGARDEN. Stanford

University, Stanford, California. Biodiversity consequences of directed
versus random movement patterns.

Most ecological and evolutionary theories of spatial dynamics assume organismal movements are unconditional, non-directional and occur at constant rates. We explore an alternative representation, in which movement
is viewed as an adaptive response of a phenotype to varying environmental
conditions. In this representation of directed movement, we assume movement stimuli derive from local gradients in fitness and that movements are
oriented along these gradients. We compare the implications of the two
movement assumptions for speciation and biodiversity patterns. Random
movement results in high alpha-diversity but minimal beta-diversity. Too
much random movement results in a loss of regional diversity. Directed
movement maintains biodiversity through a spatial partitioning of species.
Therefore, with directed movement, alpha-diversity is low, and beta-diversity is much higher. There is no cost of vagility for regional diversity with
directed movement. We consider speciation by reinforcement. An intermediate dispersal principal applies for speciation with random movement.
Directed movement precludes speciation in undisturbed environment, but
speciation is possible with directed movement when there are infrequent
environmental disturbances.

ARNOLD, HEATHER G.* and LONNIE W. AARSSEN.* Queens University, Kingston, ON, Canada. The influence of local and regional parameters on species richness in riparian vegetation.

Streams provide a linear, continuous habitat along which community patterns and dynamics have long been examined. The discrete nature of riparian habitat creates a convenient study system in which the influence of
elusive regional factors such as dispersal and effective regional species pool
size can be estimated. If hydrochory, or diaspore transport by water, is the
principal means of dispersal along streams then seeds are preferentially
dispersed downstream. Models predict that the available species pool is
constantly increasing downstream as seeds and species accumulate from
upstream communities. We extend this model by predicting that the more
community-types the stream traverses, the greater the number of species
that can potentially accumulate at the downstream sink. Our hypotheses
were that: 1) both local and regional parameters can explain more of the
variation in site richness than can local variables alone; 2) streams flowing
through highly heterogeneous landscapes have the largest species pool
available for recruitment, and consequently the highest species richness at
the downstream study site. We established study sites on 100 streams that
represented a range of upstream heterogeneities and lengths in SE New
Brunswick. At each study site we inventoried 60m2 of streamside vegetation, and recorded data for local variables hypothesized to influence plant
richness: canopy cover, channel width, bank slope, and ground cover (rock,
moss, soil and litter). Regional factors hypothesized to influence richness
(i.e. habitat heterogeneity, stream tortuosity and length) were extracted
from GIS data. Richness ranged from 21 to 109 species per site. Despite
the range in plant richness observed, and the number of local and regional
variables considered, we were able to explain very little of the variation in
riparian plant richness.

Abstracts

15

ARNOLD, JENNIFER M.,1,* SOLANGE BRAULT2 and JOHN P. CROXALL.3 1 Auburn University, arnolj1@auburn.edu, Auburn, AL, USA; 2 University of Massachusetts, Boston, MA, USA; 3 British Antarctic Survey,
Natural Environment Research Council, Cambridge, UK. Population
models in a conservation setting: A case study of the albatross and
long-line fishing.

Natural resource managers are regularly faced with making regulatory decisions for long-lived species that spend most of their life in remote places.
Even if accurate information on vital rates is available, understanding the
sensitivity of these populations to anthropogenic or environmental stresses
is difficult. Population modeling, as a tool for management, addresses these
issues; we present a stage-classified model designed to elucidate causes of
fluctuations in a population of black-browed albatross over the past 3 decades. As with many albatross species, the black-browed albatross population at Bird Island, South Georgia, is currently declining. This may reflect
fishing activities around their breeding and wintering habitats and the availability of krill, their primary food source. The model incorporates details
of the life cycle, including variation in age at maturity and annual breeding,
and behavioral differences between successful and unsuccessful breeders;
it is tested with 23 years of demographic data. When using field-measured
values for survival, reproductive success and breeding probabilities, our
model trajectory closely follows the observed changes in breeding population size. The model projection gives a 98% chance pseudoextinction (N
, 25 pairs) within 81 years given current conditions. Comparing pre-1988
and post-1988 demography reveals a 6.1% decrease in lambda, the population rate of increase. This decline is correlated with changes in fishing
activities in the home range of this population. A life table response experiment (LTRE) shows that the decline in lambda is caused mostly by
declines in return rates of breeders. These results provide clear evidence
that management actions should focus on sources of adult mortality. We
argue that the correlation between fishing activities and changes in adult
survival in this albatross species supports increased regulatory effort for
long-line fishing activities, including increased monitoring of illegal fishing.

ARNONE, JOHN A.,1,* DAVID SCHORRAN,1 DALE W. JOHNSON,2

LINDA L. WALLACE,3 YIQI LUO,3 JAMES S. COLEMAN,1 DAVID S.
SCHIMEL4 and PAUL VERBURG.1 1 Desert Research Institute, Reno, NV;
2
University of Nevada, Reno, NV; 3 University of Oklahoma, Norman, OK;
4
National Center for Atmospheric Research, Boulder, CO. NSF-I.R.C.E.B.
Interannual climate variability and ecosystem processes in tallgrass
prairie: Effects on ecosystem CO2 fluxes.

Mechanistic numerical models currently in use can reliably estimate the

effects of intra-annual, or seasonal, effects on terrestrial ecosystem CO2
exchange. However models become less robust when estimating interannual, or year-to-year, effects because the response times and lagged behavior of key ecosystem processes are poorly understood. In a 4-year experiment using the EcoCELL whole-ecosystem controlled-environment gas exchange facility at DRI, we are quantifying the responses to an anomalously
warm year of interlinked ecosystem processes that are involved in modulating the ecosystem CO2 fluxes. We excavated, containerized, and installed
twelve 13,000 kg intact tallgrass prairie soil monoliths with their respective
plant communities into four separate EcoCELLsthree monoliths per
EcoCELLin the fall of 2001. Starting in February 2002, have been exposing all monoliths to an "average" Oklahoma climate. In February 2003,
we increased the air temperature in two of the EcoCELLs by 48C to simulate an anomalously warm year while maintaining an "average" climate
(air temperature) in the other two EcoCELLs. Mean daily net ecosystem
CO2 exchange (NEE) in 2002 increased from wintertime lows (Feb. 2002
NEE: 0.39 mmol CO2 m-2 s-1 uptake) to relatively large rates of uptake (June
2002: 4.34 mmol CO2 m-2 s-1) as plant communities developed, followed by
immediate and dramatic declines in NEE after aboveground harvest (September 2002: 0.22 mmol CO2 m-2 s-1). Results to date indicate "normal"
responses of NEE to plant phenology, and significant initial responses to
experimental warming (Feb. 2003: -0.12 mmol CO2 m-2 s-1, CO2 efflux).

16

Abstracts

ARNOTT, SHELLEY E.,1,* BILL KELLER,2 CARRIE HOLT3 and NORMAN D. YAN.4 1 Queens University, Kingston, Ontario, Canada; 2 Cooperative Freshwater Ecology Unit, Sudbury, Ontario, Canada; 3 Simon
Fraser University, Burnaby, British Columbia, Canada; 4 York University,
Toronto, Ontario, Canada. Local and regional-scale assessment of zooplankton recovery from acidification.
There is overwhelming evidence that many of the worlds ecosystems have
been damaged by human activities. This has prompted international agreements to alter human activities and reduce their impact on ecosystems (e.g.,
Kyoto Protocol, Great Lakes Water Quality Agreement, Clean Air Act).
Now, some of the most pressing challenges facing scientists involve understanding the process by which damaged ecosystems recover. Lakes in
Killarney Wilderness Park, Canada, provide a rare opportunity to study
mechanisms associated with the recovery of aquatic communities after a
prolonged period of acidification. Since the 1970s, local sulphur emissions
have been reduced an order of magnitude, resulting in widespread chemical
recovery of many damaged lakes. Limited evidence of biological recovery
associated with improvements in water quality, suggests that biotic communities may not be as resilient as water chemistry. We analysed data from
three years (1972, 1990, and 2000) for zooplankton communities in 44
Killarney Park lakes to examine biological recovery trajectories at both
local and regional scales. Twenty-two reference lakes in south-central Ontario, an area less influenced by acid deposition, were compared with Killarney Lakes. Regional richness for lakes in Killarney Park was higher than
in the reference area, primarily resulting from the presence of rare species.
Mean individual lake richness, however, tended to be lower in the recovering lakes than in the reference lakes. Species turnover rates were high in
recovering lakes, reflecting community changes associated with recovery.
High regional diversity and species turnover rates suggest that colonists
have access to lakes and that zooplankton recovery is probably limited
primarily by local conditions. Assessing diversity at both local and regional
scales provides additional insights into factors that limit biological recovery
in stressed ecosystems.
ASCHENBACH, TODD A.,* KELLY KINDSCHER and BRYAN FOSTER. University of Kansas, Lawrence, KS. Dispersal limitation in an
early successional grassland.
It is suggested that local species richness is limited by seed availability in
low productivity environments and that this limitation becomes less important as productivity increases. To evaluate the role of propagule availability in regulating species richness along a natural nitrogen gradient, 30
prairie forb species were added to 80 plots in an early successional grassland in eastern Kansas over a two-year period. Species were seeded in
spring 1999 and spring 2000. Treatments 1 and 4 received all 30 species,
treatments 2 and 3 received 20 forb species, and the control plots (N520)
did not receive any forb species. Species richness of individual treatments
and the control were paired along the nitrogen gradient and compared to
evaluate the effect of seed additions relative to productivity. Data from fall
and spring 2001 and 2002 are reported separately. Analyses indicate that
seed additions result in significant increases in species richness (one-way
ANOVA; P,0.001), however, the magnitude of this affect decreases with
increasing soil nitrogen. In the absence of seed additions, species richness
is positively correlated with soil nitrogen indicating that the range of productivity is characteristic of the left side of the classic hump shaped curve
formed by the relationship between species richness and productivity. Results agree with the hypotheses that dispersal limitation is important in
regulating rates of succession and species accumulation during early succession.
ASHTON, ISABEL W.* and MANUEL LERDAU. Stony Brook University, Stony Brook, NY. Exotics do NOT always invade: Mechanisms
underlying the success and failure of vine invasions.
Some species of exotic woody vines are aggressive invaders in temperate
forests, yet others do not invade these ecosystems. Invasion success in vines
may be related to differential growth and response to light environment.
To test the hypothesis that invasive vines are better adapted to both low
and high light environments than their non-invasive relatives, I performed
a greenhouse experiment in Long Island, NY with temperate vines from 3

families (Caprifoliaceae, Celastraceae, and Vitaceae). I grew plants of 3

invasion categories (invasive exotic, non-invasive exotic and native) within
each family. Plants were grown under either shade cloth (75% ambient) or
ambient light conditions. Invasive species grew significantly faster than
either native or exotic non-invasive species under both treatments. These
results suggest that invasive vines have the capacity to grow rapidly even
in the forest understory. To examine the physiological underpinning of this
response, light response curves and maximum photosynthetic rates were
measured. The exotic non-invasive vines respond to increases in light availability differently than native or invasive vines. Native and invasive vines
respond to increases in light availability with increases in photosynthetic
capacity. In contrast, non-invasive vines do not increase their photosynthetic capabilities. These results suggest that exotic vines that have not
established in the United States may be limited in part by their physiological properties.
ATHY, ERIN R.1 and CAROLYN H. KEIFFER.2 1 Miami University,
athyer@muohio.edu, Oxford, OH; 2 Miami University, Keiffich@muohio.
edu, Oxford, OH. Effects of mulch on edaphic conditions and seedling
survival on a closed Ohio landfill.
Restoration of degraded lands, including closed landfills, is urgently needed; however, many restoration attempts fail due to poor soil conditions
(e.g., compacted, low in nutrients and organic matter). Current agricultural
strategies include applications of mulch, which are known to increase soil
organic matter, increase soil moisture, moderate rhizosphere temperatures,
and reduce herbaceous competition. Employing such practices on sites with
similar soil limitations may increase tree survival and growth. This study
evaluates the effect of mulch type and depth on tree seedling survival and
edaphic conditions on Center Hill landfill, Cincinnati, OH. The area, a
former gravel mine in the 1950s, was filled to capacity with urban and
construction refuse until it was capped with 1.5m of compacted clay and
0.5m of fill dirt in the 1970s. A subsequent reforestation attempt with
hybrid poplars failed (17.5% survival). In Spring 2002, five species of tree
seedlings (Fraxinus pennsylvanica, Robinia pseudoacacia, Prunus serotina, Quercus macrocarpa, Populus spp.) were planted as bare root seedlings
(n51750) on each of 35 replicate plots, including non-mulched controls.
Three types of mulch (hardwood chips, leaf, and mixed) were distributed
in three depths (0, 5, 15 cm). Gravimetric soil moisture and temperature
were taken monthly (May-September). Seedling survival varied by species,
from Fraxinus (97%) to Prunus (11%). Populus, Prunus, and Robinia benefited by mulch additions; 15cm hardwood and mixed mulch improved
survival of Prunus (p,0.005) while leaf mulch had little effect. Populus
responded positively to the 15 cm application of any mulch (p,0.004),
while Robinia was sensitive to specific treatments (p,0.004). Fraxinus and
Quercus showed no significant differences in survival on mulched plots
compared to the control. Data suggests that the 15 cm mulch treatment
negatively impacted herbaceous growth (p,0.017) and lowered soil temperatures (p,0.0001). Soil moisture was not significantly different per
treatment, attributed to drought conditions.
AUGSPURGER, CAROL K.,* JOHN M. CHEESEMAN and CARL F.
SALK. University of Illinois, carolaug@uiuc.edu, Urbana, IL. Potential
for carbon gain by understory plants via phenological avoidance of
canopy shade.
Understory individuals that avoid canopy shade by leafing out early in
spring and/or maintaining leaves into late autumn have the potential to
enhance carbon gain. We evaluated the extent to which leaves of five species are phenologically, developmentally, and physiologically capable of
exploiting these periods of high light within the understory. Adults of three
species of shrubs/treelets and juveniles of two canopy species were sampled. Measurements were made throughout leaf lifespan of phenological
stage, leaf area and mass, chlorophyll, nitrogen, and light- and CO2-saturated net C assimilation (Asat). Dates of canopy closure in spring and opening in autumn were quantified based on phenological observations of canopy tree species weighted by their relative basal areas. For leaves of all
species, Asat (ymol CO2 m-2 sec-1) peaked and plateaued during and immediately following expansion and then slowly declined until senescence.
The period of highest photosynthesis paralled highest levels of leaf nitrogen
(mg/cm2), but not of chlorophyll (mg/cm2), which did not peak until well

after leaf expansion. Species differed markedly in whether sufficient phenological avoidance occurred to capitalize on the potential for high carbon
gain in spring. Prior to canopy closure, Aesculus glabra juveniles had the
highest potential for carbon gain, whereas Asimina triloba had no potential
for carbon gain because its major leaf expansion postdated canopy closure.
Lindera benzoin, Carpinus caroliniana, and Acer saccharum juveniles were
intermediate in response, with minor avoidance of shade and minor potential for carbon gain in spring. In contrast, potential for carbon gain in
autumn was non-existent or negligible for all species either because leaf
senescence occurred prior to canopy opening or because of low Asat of
senescing leaves. Species differences in carbon gain by phenological avoidance could contribute to variation in resource availability for survival,
maintenance, and reproduction of these understory plants.
AUKEMA, JULIANN E.* and ANDREW B. CAREY. Pacific Northwest
Research Station, USDA Forest Service, Olympia, WA, USA. Restoring
biocomplexity in Douglas-fir forests: Understory development after inducing canopy heterogeneity and underplanting tree seedlings.
Ancient forests have become prominent in conservation issues in North
America, in part because of their value as habitat for threatened species
and reservoirs of biological diversity. In the Pacific Northwest, the northern
spotted owl is a symbol of the conflict between conservation values and
economic values associated with old-growth Douglas fir forests. Today,
early successional forests make up nearly half of Federal forested lands in
the Pacific Northwest. The Forest Ecosystem Study is an experiment in
using variable density thinning (VDT), to induce spatial heterogeneity in
forest canopies, and underplanting of shade tolerant tree seedlings, to restore tree species diversity, with the goal of accelerating development of
second-growth forests into forests with some of the characteristics (e.g.
structural, biological diversity) of old, natural forests. We conducted this
study in two forests with different management histories (commercial and
legacy). We examined the survival and growth of underplanted Abies grandis (grand fir), Thuja plicata (western redcedar), Pinus monticola (western
white pine), and Alnus rubra (red alder) in experimentally thinned stands,
and composition and cover of understory vegetation in both thinned and
unthinned stands. We found that forest management history had the greatest
effect on seedling survival and growth and on understory vegetation. Survival of seedlings and understory vegetation was not directly correlated
with thinning intensity or overstory canopy. This is probably because the
influence of canopy gaps is displaced due to the low sun angles in this
region. Variable density thinning in conjunction with other conservation
measures may accelerate biocomplexity in second-growth forests and holds
promise for providing a full range of ecological services and economic
goods.
AULD, JOSH R.* and RICK A. RELYEA. University of Pittsburgh, Department of Biological Sciences, Pittsburgh, PA, USA. Having the guts
to compete: Competitor-induced plasticity in tadpole intestines.
Phenotypic plasticity is the environment-specific expression of a genotype.
In amphibians, there has been a growing interest in the plasticity of external
morphology in response to predators and competitors and how these changes affect amphibian performance. However, there has been little attention
paid to how predators and competitors simultaneously affect the internal
morphology of amphibians. If critical shifts in organ morphology occur in
different environments, then this could be an important, yet neglected area
of plasticity. We examined the relative gut length of wood frog tadpoles
(Rana sylvatica) in the presence of intra- and interspecific competition
(Pseudacris triseriata). Intra- and interspecific competitors both reduced
wood frog growth, although intraspecific competition had a larger negative
effect. At the same time, wood frogs facing higher competition developed
relatively longer intestines. This phenotypic change appears to be an adaptive response because it results in increased wood frog growth. The ability
to alter relative intestine length could be the underlying mechanism that
tadpoles use to improve their fitness and may alter competitive outcomes
in aquatic communities. This more integrated view of morphological plasticity is shedding new light on the way we conceptualize amphibian plasticity and the underlying mechanisms of competition.

Abstracts

17

AUSTIN, AMY T.* and LUCIA VIVANCO. Faculty of Agronomy and

IFEVA-CONICET, University of Buenos Aires, Buenos Aires, austin@
ifeva.edu.ar, ARGENTINA. Abiotic and biotic controls on litter decomposition in the semiarid Patagonian steppe.
Controls on decomposition and nutrient turnover in water-limited ecosystems are elusive, as many studies of litter decomposition have shown contradictory results and little correlation with annual precipitation. We carried
out two different factorial experiments designed to disentangle the abiotic
and biotic controls on litter decomposition in water-limited ecosystems. In
the first experiment, we decomposed litter of mixed native grasses under
different light regimes (full sunlight, reduced ultraviolet-B (UV-B) radiation, and blocked total radiation), in combination with a treatment of soil
sterilization. In the second experiment, we decomposed litter of mixed native grasses treating underlying soils with additions of labile carbon and
inorganic nitrogen. After one year of decomposition in the field, photodegradation was the dominant control on mass loss (P,0.001) with a 50% and
250% reduction in litter mass loss in the reduced UV-B and blocked total
radiation treatments, respectively. Soil sterilization had no effect on mass
loss. In the substrate addition experiment, increased inorganic nitrogen accelarated litter mass loss by 15% (P50.05), while carbon or combined
carbon and nitrogen additions did not affect decomposition. Nutrient release in early stages of decomposition, however, was significantly affected
by soil substrate availability, with increased nitrogen immobilization in
litter from carbon addition plots (P,0.01). We conclude that photodegradation is the dominant control on litter mass loss in this ecosystem, and
that surprisingly, UV-B radiation may account for a large fraction of these
abiotic losses. Biotic controls are secondary to photodegradation for litter
mass loss, but may become critical for soil organic matter turnover and
nutrient cycling.
AWADA, TALA,1,* MICHEAL E.L. PERRY2 and WALTER W.H.
SCHACHT.2 1 School of Natural Resource Sciences, University of Nebraska, Lincoln, NE, USA; 2 Department of Agronomy and Horticulture, University of Nebraska, Lincoln, NE. Responses of Bromus inermis and Andropagon gerardii to various canopy levels of Fraxinus pennsylvanica.
Agroforestry systems are designed to improve efficiency of use of available
resources and to increase potential site productivity. The capability of a
plant to acclimate to shade when cultivated beneath trees is important in
determining success of agroforestry projects. The objectives of this study
were to determine the morphological, physiological and growth responses
of C4 big bluestem (Andropogon gerardii Vitman.) and C3 smooth bromegrass (Bromus inermis Leyss) to various canopy levels of green ash (Fraxinus pennsylvanica Marsh) in the field, and to examine the impacts of these
responses on grass yield. Net photosynthesis (Anet), stomatal conductance
(gs), and dark respiration (Rd) declined in response to shade in both species,
but the decline was steeper in big bluestem than in smooth bromegrass.
Total chlorophyll content (Tchl), specific leaf area (SLA), and nitrogen (N)
content of the leaves increased with shade in both species. In addition,
Tchl, SLA, N, and gs were significantly higher in smooth bromegrass than
in big bluestem at all canopy levels. Lower gs and N, and higher Anet in
big bluestem resulted in a higher water and nitrogen use efficiencies in this
species than in smooth bromegrass. Yield of big bluestem sharply declined
with increased canopy cover, whereas yield of smooth bromegrass was not
affected by canopy cover. Our results indicated that while both species were
productive under various levels of green ash canopy, and showed similar
ecophysiological responses to shade, smooth bromegrass acclimated better
to shade than big bluestem. Therefore, the selection of species should be
based on specific management goals and environmental conditions. For
example the higher water and nitrogen use efficiency in big bluestem compared to smooth bromegrass may become advantageous in drier climates
and where N is a limiting factor.
AYRES, MATTHEW P.,1 SHARON J. MARTINSON,1 TIINA YLIOJA,1
RICHARD W. HOFSTETTER1 and KIER D. KLEPZIG.2 1 Biology, Dartmouth College, Hanover, NH, USA; 2 Southern Research Station, USDA
Forest Service, Pineville, LA, USA. Causes and consequences of dynamic feedback in population dynamics of Dendroctonus frontalis (Coleoptera: Scolytidae).
High impact herbivores are frequently those that display intermittent outbreaks. Outbreaks can result from dynamic feedback systems that include

18

Abstracts

delays and/or nonlinearities in per capita growth rates as a function of

density. Dendroctonus bark beetles have large impacts on forests during
outbreaks that can kill vast areas of pine. Dendroctonus typically employ
pheromones to organize mass attacks of individual trees - apparently as a
means of overwhelming tree defenses. This could generate destabilizing
positive feedback (Allee effect) in which per capita reproduction tends to
increase with increasing abundance of beetles to participate in attacks. We
tested for an Allee effect in D. frontalis by comparing growth rates of local
infestations that began at a range of population sizes. As predicted, the
relative growth of infestations was positively related to initial size. To evaluate whether this was due to density-dependent interactions with tree defenses, we (1) developed a model that predicts dynamics of the tree resin
system during attacks and eventual reproductive success of the beetles, and
(2) tested model predictions by measuring attack rates, resin dynamics, and
reproduction within natural infestations. Results supported predictions that
resin flow declines most rapidly in trees sustaining high attack rates, that
per capita reproduction declines with resin flow as a negative exponential,
and that reproductive success is maximized at intermediate attack rates (due
to increasing negative feedback from competition at high densities). However, the surprising additional result was that average beetle attack rates
were unrelated to population size except at very low local abundance. This
was because even low to moderate populations of beetles were efficient at
concentrating their attacks on just a few trees. Thus, although beetle attack
rates influence reproductive success, and vary greatly among trees, this
does not seem to produce density-dependence among infestations of different sizes because attack rates are uncoupled from local population size.
D. frontalis displays properties of a positive dynamic feedback, but the
presumed mechanism seems to be wrong. Bark beetle demographics appear
to involve feedback systems that operate at multiple spatial scales.
BACKUS, VICKIE L.1,* and JOAN M. HERBERS.2 1 Middlebury College,
Middlebury, VT; 2 Ohio State University, Columbus, OH. Testing alternative models of social evolution.
The decisions an organism makes when allocating energy will affect every
aspect of its behavior. This process is especially complicated for eusocial
species because the parties in the nest may disagree over the decisions. In
ants hapliodiploid sex determination means that queens and workers have
different optimum sex allocation ratios. Differences in life span for these
two parties means they also differ in their optimum allocation ratios between growth and reproduction. Natural selection is superimposed on these
conflicts which can lead to different sets of allocation rules as being optimal
depending on the environment. Path analysis was used to determine the
magnitude and direction of internal conflict over allocation decisions, and
to determine the rules used by nests in making those decisions, for the
small forest ant Leptothorax longispinosus. The data show that conflict over
allocation decisions is common in this species; queens and workers frequently disagreed over the amount of energy to allocate to reproduction
versus growth and how to package that allocation. Which party won the
conflict varied both temporally and spatially. Path analysis also showed
that selection would lead to different rules being used by nests while making these important decisions. Nests located in Vermont used a set of rules
that operate at a proximate level; the parties in the nest first packaged
resources into diploid or haploid offspring and then later decided how much
of the diploid resources became growth versus reproduction. On the other
hand nests in New York used rules that evolved under an ultimate level
selection pattern where nests first decided how much to allocate to growth
or reproduction and later decided how to package those resources.
BADE, DARREN L.,1,* STEPHEN R. CARPENTER,1 JONATHAN J.
COLE2 and MICHAEL L. PACE.2 1 University of Wisconsin-Madison,
Madison, Wisconsin, USA; 2 Institute of Ecosystem Studies, Millbrook,
New York, USA. Exogenous and endogenous origins of DOC in lakes:
Surprising results from whole lake 13C additions.
Dissolved organic carbon (DOC) is the largest reservoir of organic carbon
in the water column of many lakes. DOC can originate from primary production within the lake itself (endogenous) or in the terrestrial watershed
(exogenous), and both sources are often thought to have distinctive dynamics and roles in lakes. Because DOC consists of a highly complex mixture
of organic compounds, little is known about the sources of the standing

stock of DOC. To determine the sources and fates of DOC in lakes we

conducted four whole ecosystem inorganic carbon stable isotope additions.
In one of these manipulations (Peter Lake, 2002) we also added inorganic
N and P to increase primary production; in the other experiments, no nutrients were added (Peter and Paul lakes, 2001; Tuesday Lake 2002). Prior
to 13C enrichment, background isotope signatures of DOC were similar to
terrestrial carbon and consequently it was difficult to discern the contribution of each source of DOC. Following 13C addition, the DOC isotope
signature increased between 1.5 and 9.5 per mil, due to DOC inputs from
13
C labeled primary producers. The largest change in the DOC isotope
signature was observed in Peter Lake (2002; the year of nutrient enrichment) and the smallest change was observed in Tuesday Lake, which had
the highest concentration of DOC. Using carbon isotope results to constrain
models of carbon fluxes, algal inputs of DOC was 10 to 30 % relative to
terrestrial carbon inputs in the unfertilized lakes, and higher in the fertilized
lake. Despite the fact that terrigenous DOC is considered to be fairly recalcitrant to microbial respiration and algal DOC quite labile, the loss rates
of each form of DOC were found to be similar.
BAER, NICHOLAS A.* and WILLIAM O. LAMP. University of Maryland, College Park, MD. Detrital foodweb response to dissolved organic
carbon and nutrient changes: Exploring Coastal Plain stream dynamics.
Changes in water quality can greatly affect the detrital foodweb. Rates of
leaf decomposition have been shown to increase with nutrient enrichments.
Other research has suggested that elevated DOC levels may cause a shift
in the bacterial community and therefore changes in the detrital foodweb.
Our study of two Coastal Plain watersheds in Maryland has demonstrated
elevated phosphorus concentrations, ranging from 0.20 mgL21 to 2.75
mgL21, and low nitrate-N concentrations. In addition, Coastal Plain streams
are often characterized by black or brown water containing high levels of
dissolved organic carbon (DOC). Thus, we performed a laboratory experiment examining both microbial and macroinvertebrate response to changes
in nitrogen, phosphorus and DOC. We hypothesized that increased DOC
reduces leaf decomposition by the microbial community but not the isopod,
Caecidotea communis. We also predicted an increase in detrital processing
with elevated nitrogen concentrations. Microbes and Caecidotea communis
were reared in artificial stream environments varying in nitrate, phosphorus
and DOC concentrations. Microbial respiration, leaf decomposition rates,
and isopod assimilation rates were measured to compare treatments. The
results illustrate the relationship between nutrients, DOC, and leaf decomposition.
BAIDYA ROY, SOMNATH, PETER D. WALSH and STEPHEN W. PACALA. Princeton University, Princeton, NJ, USA. Deforestation and the
climate of equatorial African parks.
Congo and Gabon contain some of the largest blocks of intact tropical
forest in the world. However, more than half of the remaining forested area
in each country has now been allocated in timber concession. We used a
regional climate model to simulate the impact of future deforestation on
local climate. Results show widespread increases in temperature and decreases in rainfall, both in regions slated for timber exploitation and in
neighboring parks. Our results suggest that deforestation could undermine
the effectiveness of isolated parks for wildlife conservation.
BAILEY, LARISSA L.,1,* THEODORE R. SIMONS1 and KENNETH H.
POLLOCK.2 1 North Carolina State University, Cooperative Fish and Wildlife Research Unit, Department of Zoology, Campus Box 7617, Raleigh,
NC, USA; 2 North Carolina State University, Departments of Biomathematics, Statistics, and Zoology, Campus Box 7617, Raleigh, NC, USA.
Estimating site occupancy and detection probabilities for terrestrial
salamanders.
World-wide concern for amphibian populations has focused research on
documenting amphibian distributions and habitat associations over large
geographic areas. Most programs involve sampling potential habitats using
a wide variety of techniques to document species occurrence and relative
abundance. Population declines may be inferred by a decrease in the number of sites win which a species is detected. These methods assume constant

(often perfect) species detectability over time, space, and among different
sampling methods. We utilized recent developments in capture-recapture
theory to estimate proportion of sites occupied and species detection probabilities for a suite of salamanders found in Great Smoky Mountains National Park. We included large-scale habitat characteristics (e.g. elevation,
vegetation type, and stream presence) as site and sampling covariates. We
explored differences among area-constrained and coverboard sampling
techniques. We found species-specific differences in detection probability
ranging from 0.26-0.72. Detection probabilities were lower for coverboards
than area-constrained searches for all terrestrial species. Habitat characteristics were important in both site occupancy and detection probabilities but
varied among species. Nevertheless, the identification of important habitat
characteristics was consistent among sampling methods. We strongly recommend incorporating detectability estimation into research studies designed to document presence and spatial distribution of amphibian species.
BAKER, PATRICK J. Institute of Pacific Islands Forestry, USDA Forest
Service, Hilo, HI. Suppression and release during canopy recruitment
of five tropical tree species of differing shade tolerance.
How trees reach the upper canopy of tropical forests is poorly understood
due to the lack of long-term growth data. In temperate forests, ecologists
have identified several different patterns of canopy recruitment using treering analyses. To determine if similar patterns of canopy recruitment occurred in tropical tree species, I analyzed annual growth rings from 92
canopy trees (.20 cm dbh) of five species in a mature stand of speciesrich seasonal dry evergreen forest in western Thailand. The species included Neolitsea cambodiana (Lauraceae) and Vitex peduncularis (Verbenaceae), two relatively shade tolerant species, Toona ciliata (Meliaceae) and
Chukrassia tabularis (Meliaceae), two moderately shade intolerant species,
and Melia azederach (Meliaceae), a highly intolerant gap species. Mean
tree age was 67 y (range: 23-129 y). Mean annual growth rates were positively correlated with relative shade tolerance; however, there was wide
variation in radial growth rates within and amongst trees and species.
Growth release patterns differed among species. The number of minor
growth releases (i.e., growth increases of .50% sustained for 10 y) was
positively correlated to shade tolerance; however, the number of major
releases events (i.e., radial growth increases of .100% sustained for 10 y)
were similar for all species, except Melia which showed no releases. Qualitative assessments of the tree-ring series showed several different patterns
of canopy recruitment that varied in relative abundance amongst species.
These included (1) direct recruitment to the canopy, (2) a period of prolonged suppression followed by growth release into the canopy, (3) multiple periods of suppression and growth release prior to canopy recruitment,
and (4) continuous suppressed growth without growth release. These patterns are similar to ones described for temperate forests and further underscore similarities in stand development patterns between temperate and
tropical forests.
BAKER, PAUL W.* and LAURA G. LEFF. Kent State University, Kent,
OH. Response of bacteria to oligotrophic conditions: Comparison of
enumeration methods and the impact of microgravity.
Under oligotrophic conditions, bacteria can enter into a starvation survival
mode. To examine responses of bacteria to starvation, aquatic microcosms
were inoculated with bacteria that were isolated from an extremely oligotrophic setting, the water storage on the Mir Space Station. Total bacterial
counts were determined using acridine orange (AO), DAPI and DAPI-HCl
(acidification prior to DAPI staining). Bacteria were also enumerated using
Live/Dead BacLight kit (Molecular Probes), fluorescent in situ hybridization (FISH) and cultivation (to enumerate colony forming units). Staining
using AO, DAPI or DAPI-HCl showed that there were no significant differences between any of these techniques. Furthermore, using longer staining times revealed that many bacteria were not stained. Each method of
enumeration revealed different aspects of the bacterial population and the
effectiveness of the methods varied among the three species examined and
with starvation. Having illustrated the differences among detection methods, these bacteria were studied under simulated microgravity conditions.
Simulated microgravity was obtained using a device called a clinostat that
rotates vessels at a slow speed around the horizontal axis thus suspending
the bacteria in the solution. At the same time, control experiments (normal

Abstracts

19

gravity) were performed so that the vessels rotated around the vertical axis.
The experiments were conducted under starvation and low nutrient conditions to investigate cells in the planktonic phase and cells in biofilms.
The development of biofilms was determined by attachment of bacteria to
stainless steel disks mounted inside the rotating vessels. Only minor significant differences were observed for planktonic cells under simulated microgravity compared to normal gravity. However, for most bacteria, biofilm
formation under simulated microgravity was higher compared to normal
gravity. Together, the findings of these experiments suggest that the bacteria
studied are adapted to the oligotrophic, microgravity conditions from which
they were obtained and may persist in the environment because of biofilm
formation.
BAKKER, CHRIS,1,* PETER M. VAN BODEGOM,2 W.H.O. ERNST1 and
HANS J.M. NELISSEN.1 1 Free University of Amsterdam, Dept. of Ecology and Physiology of Plants, Amsterdam, NH, The Netherlands; 2 Free
University of Amsterdam, Dept. of Systems Ecology, Amsterdam, NH, The
Netherlands. Nutrient management and duration of winter flooding determine plant responses to rising groundwater tables.
In a large scale restoration project in the Dutch coastal dunes, groundwater
tables have been raised after 150 years of drinking water extractions. This
rise of groundwater is accompanied by additional nutrient management, i.e.
mowing or sodcutting, with the aim of restoring oligotrophic wet dune
vegetation. Responses in soil parameters, vegetation and planted phytometers were measured along elevational gradients in 8 dune valleys. The
effects are analysed in steps through the chain: management - abiotic
changes - nutrient uptake - plant growth. Interactions between duration of
winter flooding, nutrient management and current spring groundwater level
appear to dominate the plant responses, rather than the separate effects of
these factors. I will discuss which information we need as a minimum to
build a satisfactory model for plant responses to rewetting.
BAKKER, LIESBETH1,2,* and HAN OLFF.3 1 Wageningen University,
Wageningen, The Netherlands; 2 University of Nebraska, Lincoln, Nebraska; 3 University of Groningen, Haren, The Netherlands. Compositional
constancy in grazed plant communities increases with herbivore size.
An important current discussion is how the loss of biological diversity will
affect community structure and ecosystem functioning in grasslands. Several recent experimental and theoretical studies have shown important links
between plant diversity, the stability in species composition of plant communities and the reliability of grassland ecosystems to perform certain critical functions, e.g., to maintain productivity. In most grassland ecosystems,
vertebrate herbivores are key elements without which community structure
and ecosystem functioning cannot be understood. However, herbivores have
hardly been incorporated in studies on the relation between diversity, compositional stability of plant communities and ecosystem functioning. We
experimentally investigated the effects of different-sized herbivores on the
spatio-temporal stability of grassland community composition and biomass
by stepwise exclusion of vertebrate herbivores from large to small body
size at two nutrient levels. We found that large herbivores had a strong
stabilizing effect on plant community composition, whereas excluding them
(leaving only small herbivores) induced strong variability in plant species
composition, both in space and time. Seven years of nutrient addition did
not change the vegetation in the presence of large herbivores, due to selective grazing in the fertilised plots. Small herbivores in contrast, did not
counteract the effects of nutrient addition, resulting in strong compositional
changes in the vegetation. Furthermore, large herbivores suppressed the
expression of small-scale soil differences in the vegetation, resulting in
homogeneous vegetation, while the presence of small herbivores enhanced
vegetation heterogeneity. The temporal stability of the vegetation composition increased towards higher plant species richness, and large herbivores
increased plant species richness more than small herbivores. We conclude
that different-sized vertebrate herbivores have important and unique effects
on plant diversity, compositional stability and thus possibly ecosystem
functioning in grasslands and therefore should be incorporated in studies
that link these aspects.

20

Abstracts

BALBACH, HAROLD E.* and WILLIAM D. GORAN. US Army Engineer Research and Development Center, Champaign, IL. The quest for
measures of site condition in a military installation context.
The SERDP Ecosystem Management Project (SEMP) is a Department of
Defense research program, conducted at Ft. Benning, GA, that includes a
suite of projects, each pursuing a specific hypothesis. Overall, the program
proposes to assist military installation land managers to better understand
and manage their lands for sustainable mission use and proactive stewardship. Adaptive management methodologies and tools will be developed,
based on relating SEMP research findings to management concerns. Field
research was initiated by multiple teams in 1999, after a competitive selection process. Each team originally selected study sites with reference
only to that teams hypothesis. Qualitative designations (e.g., low, medium,
and high disturbance) of relative condition were adequate for each team to
differentiate their sites. When the need arose to be able to relate the site
condition among all sites, these qualitative, relative designations broke
down. The terrestrial and aquatic site condition indices were developed,
with participation from all the research teams, as a framework for comparison across sites from all research teams. Based on this starting point,
elements such as vegetation structure, soil compaction, microfloral populations, plant productivity, soil and sediment carbon, historical land use to
the extent knowable, surface cover, and others, some of which were already
a focus of SEMP study, were identified as a basis for index development.
This presentation will examine these indices, and evaluate their effectiveness for cross-site comparisons, and their applicability beyond this unique
context.
BALICE, RANDY G.1,* and DAVID D. BRESHEARS.2 1 Ecology Group,
Los Alamos National Laboratory, Los Alamos, New Mexico, USA; 2 Atmospheric, Climatic and Environmental Dynamics Group, Los Alamos National Laboratory, Los Alamos, New Mexico, USA. Fuel structures and
fire behavior in pinyon-juniper woodlands: Contrasting low vs. high
density understory scenarios.
Pinyon-juniper woodlands, one of the most extensive forest types in the
western USA, can pose substantial fire risks from hot burning crown fires,
highlighted by large extensive fires in summers of 2002. Despite these
risks, data on fuel structure in pinyon-juniper woodlands is largely lacking,
particularly with respect to changes in the amount of tree canopy cover, as
well as predictions of fire behavior associated with fuel structure. Our objectives were to (1) measure fuel structure in pinyon-juniper woodlands
that ranged from low to high amounts of tree canopy cover, (2) to compare
measured fuel structures with the baseline used in fire models (e.g. BEHAVE), and (3) to predict fire behavior as a function of wind speed and
1-hr moisture the measurements of fuel structure. We estimated fuel pinyon-juniper models from 14 plots in northern New Mexico. These results,
which reflect low density understory conditions, are 50 to 90 percent less
than values for published fuel models, which assume high density understory conditions associated with shrubs. The low-density understory condition produces lower rates of fire spread and probability of crown fire than
the high understory conditions. Using our estimated fuel models as inputs
to fire behavior models resulted in continuous canopy fires for windspeeds
of 50 miles/hour and greater. Lower windspeeds propagated fires on the
ground or fires that burned intermittently in the canopies. These results
assume that fuels are continuous across the landscape. Actual fire behaviors
may be strongly influenced by the interspersion of three pinyon-juniper
fuel elements; trees with litter on the ground surface, intercanopy gaps with
grasses and forbs, and intercanopy gaps with bare ground. Hence, our results highlight the importance of factoring in understory conditions in evaluating fire behavior for extensive pinyon-juniper woodlands.
BALLANTYNE, ASHLEY P.,1,* MICHAEL T. BRETT2 and DOERTHE
MULLER-NAVARRA.3 1 Duke University, Durham, NC; 2 University of
Washington, Seattle, WA; 3 University of California, Davis, CA. Nutrients
disrupt trophic coherence in aquatic ecosystems.
Nutrient availability is a strong determinant of species growth and assemblage. Thus alterations to the biogeochemical cycling of such nutrients as
phosphorus can have major implications for primary productivity, growth
of consumers and ultimately water quality. Herein we present results from

a series of consumer growth experiments and a numerical model that suggest an increase in phosphorus supply may cause a phase shift between
phytoplankton and zooplankton populations in freshwater ecosystems,
thereby reducing the transfer of mass and energy between trophic levels.
Empirical results show that zooplankton growth was correlated with algal
biomass (i.e. chlorophyll-a) in all three lakes of differing states of productivity; however, zooplankton growth was positively correlated in the mesotrophic lakes and negatively correlated in the eutrophic lake. Results from
a coupled phytoplankton-zooplankton model were consistent with observations in that ecosystems with increased phosphorus levels and reduced
zooplankton growth rates experienced greater time lags between maxima
in algal biomass and zooplankton production. Thus coherence between trophic levels is important in governing the flow of biomass through ecosystems and phase changes between trophic levels may contribute to eutrophication in aquatic ecosystems.
BALLANTYNE, FORD* and ANDREW J. KERKHOFF. Masting, variance and the mean.
Masting, or synchronized reproduction in forests may occur as a result of
"communication" between trees in the form of pollen output. Satake and
Iwasa have formulated an energy based tree reproduction model that takes
coupling of trees, through pollen production, into consideration. Depending
on parameter values of their model, simulated forests exhibit synchronized
reproduction or masting as well as completely desynchronized reproduction. We use this model to test predictions about the degree of correlation
among individuals and the slope of the variance to mean scaling relationship known as Taylors power law. We show that populations of independent individuals yield a slope of one for Taylors power law whereas completely dependent individuals yield a slope of two. Desynchronized reproduction in the forest model corresponds to a population of independent
individuals and synchronized reproduction corresponds to a population of
dependent individuals. Our theory predicts a slope of one for regions of
forest model parameter space that result in desynchronized reproduction
and a slope of two for regions that result in masting. Taylor power laws
calculated from the forest model output confirm our theoretical predictions.
This indicates that Taylors power law reflects the degree of correlation
among individuals in a population.
BANCROFT, G. THOMAS* and MIKE MCMILLIAN. Breeding ecology
of Barred Owls on a central Florida cattle ranch.
We followed the breeding ecology of Barred Owls (Strix varia) on a
10,300-acre cattle ranch in central Florida during 1995 through 1999.
Barred Owls nested in palm-oak forest islands spread through a matrix of
pastures and isolated wetlands. The number of nesting pairs on this ranch
appears to be limited by the number of palm/oak forest patches and the
availability of rotten palms for nesting. Twelve to 15 pairs were found on
the ranch. From 1 to 6 pairs apparently did not attempt to nest in a given
year. Ten clutches were started in 1995, 12 in 1996, 9 in 1997, 8 in 1998
and 9 in 1999. Nesting success averaged 75% over the five years and
ranged from 56% to 100%. Success per territory ranged from 42% to 67%.
To examine nesting core area selection in Barred Owls, we calculated the
habitats found within a 25-h circled centered on the general nesting areas
and compared this to habitat components in 50 randomly placed non-overlapping 25-h circles across Buck Island Ranch. Barred Owl core areas had
significantly more area of palm and oak forests (8.1 + 1.2 h) than did
randomly placed circles (1.9 + 0.6 h; Mann-Whitney U 5 48.0, P , 0.001).
Hammocks represented 32% of Barred Owl nesting core areas, whereas
only 8% of available habitat. Core areas contained significantly less pasture
(11.7 + 1.0 h: Mann-Whitney U 5 66, P , 0.001) than did randomly
placed circles (19.2 + 0.8 h). Maintaining forested islands in this ranching
matrix will be critical for maintaining populations of Barred Owls in this
area of Florida. These areas are under increasing development pressures as
more intensive agriculture, particularly citrus, moves into central Florida.
BARNEY, JACOB N.,* ANTONIO DITOMMASO and LESLIE A. WESTON. Cornell University, Ithaca, NY. Growth and invasive potential of
two mugwort (Artemisia vulgaris) populations in contrasting habitats
and under different management practices.
Mugwort (Artemisia vulgaris L.) is an introduced invasive perennial that
exhibits wide variation in morphology and allelopathic properties. We stud-

ied the growth response and spread of mugwort plants from two morphologically distinct populations (ITH-1 and ITH-2) in two habitats typically
colonized by mugwort: a fallow field and a turfgrass field. Important differences in growth and clonal expansion were observed between the two
populations in both habitats over the 2-year study period. Although plants
from the ITH-2 population were shorter than plants from the ITH-1 population, they produced more ramets and had greater rates of expansion.
Plants from the two populations produced nearly three times as many ramets in the fallow field than in turfgrass and spread more than 1 m from
the point of introduction within the two growing seasons. Mowing had a
greater impact on the ITH-1 population than the ITH-2 population especially in the turfgrass habitat, greatly reducing the total number shoots
produced. In addition, a greenhouse study was conducted to evaluate specific morphological characters that differed between mugwort populations.
Differences in morphology, growth, and invasive potential of plants from
these two mugwort populations are discussed in light of differences in their
allelopathic abilities. The synthesis of growth data with morphological
characters and allelopathic potential has implications for preemptive management.
BARRETT, J E.,1,* ROSS A. VIRGINIA,1 DIANA H. WALL2 and WILLIAM B. LYONS.3 1 Dartmouth College, Hanover, NH, USA; 2 Natural
Resource Ecology Laboratory, Fort Collins, CO, USA; 3 The Ohio State
University, Columbus, OH, USA. Stoichiometric evolution of Antarctic
Dry Valley landscapes.
Alfred Redfield9s concept of elemental stoichiometry describes how organisms influence, and are influenced by the chemical composition of their
environment. This general approach has provided a useful framework for
understanding nutrient dynamics across the Earth9s major biomes. In the
polar desert of the McMurdo Dry Valleys, Antarctica, physical processes
are the dominant control over ecosystem development and the affect of
biology on the chemical environment is relatively inconspicuous. The dry
valleys hence provide a unique opportunity to evaluate the stoichiometric
approach and the relative influences of biotic vs. abiotic controls over ecosystem development across landscape units comprising a gradient of biological activity and production, i.e. soils, streams and lakes. We report that
biotic control over C:N:P is poorly expressed in soils of the dry valleys,
while wider C:N and C:P ratios in the aquatic ecosystems reflect important
biological influences over ecosystem stoichiometry. C:N:P ratios widen
across landscape units from soils, to streams to lakes. In the oldest and
most extreme soil environments, C:N:P ratios are not different from the
parent material, while the wide C:P ratios of stream water and some lakes
indicate severe P deficiency and significant biotic control over the availability of nutrient elements. Based upon these data we conclude that the
elemental stoichiometry of dry valley ecosystems is ultimately regulated
by geomorphic processes operating over long time scales, while biotic communities may affect local C and N availability in productive ecosystems
over seasonal to annual cycles. The exchanges of C, N and P across these
ecosystems, facilitated by hydrology and glaciation, link the landscape units
of the dry valleys.
BARRETT, KYLE,1,3,* D. ALEXANDER WAIT1 and WENDY B. ANDERSON.2 1 Southwest Missouri State University, Springfield, MO; 2 Drury University, Springfield, MO; 3 Auburn University, Auburn, AL. Small
island biogeography in the Gulf of California: Lizards, the subsidized
island biogeography hypothesis, and the small island effect.
We used insular lizard communities in the Gulf of California to test predictions of the subsidized island biogeography (SIB) hypothesis and the
small island effect (SIE). The SIB hypothesis suggests spatial subsidies
may alter species richness patterns on small islands, while the SIE suggests
a break-point model more accurately depicts species-area relationships by
representing inter-island variation in the species richness of small islands.
To evaluate the SIB hypothesis, we identified subsidized and unsubsidized
islands based on surrogate measures of allochthonous productivity (i.e.,
island size and bird presence). Subsequently, we created species-area
curves for Gulf island lizards. We used the residuals and slopes from these
analyses to compare species richness on subsidized and unsubsidized islands. To test for a SIE, we used breakpoint regression to model the relationship between lizard species richness and island area. We compared

Abstracts

21

these results to results from the log-linear regression model. Subsidized

islands had a lower, but not significantly different, slope than unsubsidized
islands. In addition to comparing slopes, we tested for differences in the
magnitude of the residuals (from the species-area regression of all islands)
for subsidized versus unsubsidized islands. We found no significant patterns
in the residual values for small versus large islands; however, we found
that islands without seabirds have significantly greater variance from the
species-area linear regression relative to seabird-inhabited islands. We
found the SIE to be a better predictor of lizard species richness than the
log-linear model, although it did not explain a significantly greater amount
of variation. Predictions of the SIB hypothesis were not strongly supported
by the data; however, the absence of a significant SIE may be a result of
spatial subsidies as explained by the SIB hypothesis and data presented
here. We suggest potential scenarios to test for interactions between these
two small island hypotheses.
BARRETT, MARK A.* and PETER STILING. University of South Florida, Tampa, Florida, USA. The effects of Key deer (Odocoileus virginianus clavium) herbivory on forests in the Florida Keys.
This study examined the effects of herbivory on native plant species by
the endangered Florida Key deer (Odocoileus virginianus clavium), which
have increased from approximately 50 animals in the1950s to over 700 at
present. To assess deer impacts, over 250 stratified vegetation transects
were conducted in hammock, pineland, buttonwood, and mangrove habitats
throughout 13 islands (categorized into deer densities of high, intermediate,
and low). Also, six exclosures were established in hammock and pineland
on a high-density deer island. Transect data indicated very low densities
of preferred woody plant species less than 1.2m in height on islands with
high deer densities. Hammock exclosures showed a mean increase in height
of preferred woody species, while pineland exclosures exhibited a mean
increase in percent cover of herbaceous vegetation. Using cut stems of 16
native woody species in cafeteria feeding trials, it was shown that deer
highly preferred the same plant species that were found in low densities in
the transect samples. Furthermore, numerous nursery plants of eight native
species set in hammock and mangrove habitats on six Keys showed very
high herbivory rates on preferred plant species while the non preferred
species remained untouched. The data suggest that on islands of high deer
densities preferred plant species may fail to regenerate while unpalatable
plant species become dominant. Therefore, a balance must be found between increasing Key deer numbers and maintaining their habitat.
BARRON, STACY H.,1,* NOAH LONG,1 JOHN LICHTER1 and WILLIAM H. SCHLESINGER.2 1 Biology Department and Environmental
Studies Program, Bowdoin College, 6500 College Station, Brunswick, ME,
04011; 2 Nicholas School of the Environment, Duke University, Durham,
NC, 27708. Turnover rates of particulate organic matter and chemical
fractions of forest soils under elevated atmospheric CO2.
The Duke Forest Free Air CO2 Enrichment experiment (FACE) offers a
unique opportunity to quantify rates of soil carbon turnover in a deciduous
forest under elevated concentrations of atmospheric CO2. Because the CO2
used in these experiments is derived from natural gas it is strongly depleted
in 13C relative to 12C; this 13C depleted CO2 is assimilated into plant biomass, thereby lowering the 13C:12C ratios of new plant tissue. As soils
incorporate new plant tissue over time the turnover rate of different soil
carbon pools can be estimated from rates of change in 13C:12C ratios. The
estimated mean residence time (MRT) of soil organic matter in the bulk
mineral soil between 0-15 cm depth was 22.0 years and for the bulk mineral
soil between 15-30 cm depth, 62.9 years. To quantify turnover of various
soil carbon pools we fractionated organic matter in the mineral soil by
particle size and chemical methods. Coarse (. 0.250 mm) and sand-sized
(0.053-0.0250 mm) fractions were wet-sieved while the silt and clay-sized
(, 0.053 mm) fraction was centrifuged. In the upper mineral soil, the
coarse, sand-sized, and silt/clay-sized fractions had MRTs of 12.3, 28.2,
and 30.3 years, respectively. In the mineral soil between 15 and 30 cm
depth, these same particulate organic matter (POM) fractions had MRTs of
41.8, 57.4, and 118.7 yr, respectively. A strong acid extraction was used
to isolate clay-bound organic matter and a cold-water extraction was used
to gather labile organic matter. The MRT of clay-bound organic matter was
shorter than expected, 26.6 years, probably due to land use history, while

22

Abstracts

that of labile organic matter was 14.0 years. These results are consistent
with increasing stability of soil carbon with decreasing particle size, and
with the recalcitrant nature of clay-bound organic matter.
BARRON-GAFFORD, GREG A.,1,* KATIE A. GRIEVE,1 TERENCE E.
PAIGE,1 EDWARD G. BOBICH,1 RANDY L. PATTERSON,1 JOHN D.
MARSHALL2 and RAMESH MURTHY.1 1 Biosphere 2 Center, Columbia
University, Oracle, AZ, USA; 2 University of Idaho, Dept of Forest Resources, Moscow, ID, USA. Influence of drought and VPD on net ecosystem carbon exchange and CO2 uptake of cottonwoods.
Under the controlled conditions of a sealed greenhouse system at the Intensive Forestry Mesocosm (Biosphere 2 Center), eastern cottonwoods (Populus deltoides Bartr.) established in 1998 were used to examine the concomitant effects of atmospheric vapor pressure deficit (VPD) and drought
on net ecosystem carbon exchange (NECE), sap flux, carbon dioxide (CO2)
uptake, conductance, and transpiration under ambient and elevated levels
of CO2. Under low VPD (1.0 kPa) and high soil water content (0.3 v/v),
CO2 uptake for plants growing under 400, 800, and 1200 ppm CO2 averaged 18.8, 34.4, and 40.2 mmol m-2 s-1, respectively. Under high VPD (3.3
kPa) and high soil water content (0.3 v/v), CO2 uptake for plants under
400, 800, and 1200 ppm CO2 averaged 17.3, 30.9, and 39.0 mmol m-2 s-1,
respectively. Four weeks into the drought volumetric soil water content
declined to 0.1 v/v, resulting in a 40% decrease in CO2 uptake under low
VPD for all of the CO2 treatments. Under high VPD, CO2 uptake decreased
by an average of 80% for all CO2 treatments. In addition, transpiration,
conductance, respiration, and sap flux all decreased significantly as soil
water content declined for plants in all of the CO2 treatments. Leaf-level
CO2 uptake data were used to estimate stand-level CO2 uptake values using
leaf area measurements. NECE for each treatment was calculated using a
mass balance model within this particular closed system. Results at leaf
and stand levels will be compared and discussed.
BARRY, MATTHEW J.* and JEFFREY M. KLOPATEK. Arizona State
University, Tempe, Arizona. Carbon and nitrogen pools in old- and second-growth Pacific Northwest coniferous forests.
Pacific Northwest (PNW) coniferous forests contain the greatest amount of
belowground carbon (C) storage and flux of any ecosystem in the conterminous United States. Nitrogen is a regulator of C sequestration and release
in these forests. We compared inorganic N and belowground C and N pools
in three recently logged (young; 25 y) and three old-growth (450-750
y) forest stands in south-central Washington at and near the Wind River
Canopy Crane Research Facility. Total ammonium (NH4) and nitrate (NO3)
levels were quantified along a vertical gradient throughout the forest stands
using ion exchange resin lysimeters. Lysimeters were placed at 10 m intervals descending the canopy (in three different tree species), above and
below the forest floor, and 20 cm into the mineral soil. We also measured
C and N pools in forest floor material and soils to a depth of one meter.
Forest canopies absorbed a substantial proportion of inorganic N, and our
results are compared to data from a local NADP monitoring site. Ammonium levels exceeded NO3 in all stands, and the old-growth stands tended
to have more NH4 in soils than young. Within age class, sites with red
alder (a N-fixing species) exhibited more NH4 in soils than those without
N-fixers. In contrast to NO3, ammonium patterns mimicked N pools for all
stands and were maintained throughout the belowground profile. Carbon:
nitrogen ratios increased with age in forest floor material due to both higher
C and lower N concentrations in old-growth compared to second growth
stands. However, C:N ratios decreased with stand age in mineral soils. Total
forest floor C and N pools were larger in the old-growth stands due to
greater biomass. Carbon and N pools were also greater in old-growth soils
at 0-10 cm depths but no differences were observed in deeper soils.
BARTEL, REBECCA A.1,* and FREDERICK F. KNOWLTON.1,2 1 Utah
State University, Logan, UT, USA; 2 USDA/WS/National Wildlife Research
Center, Logan Field Station, Logan, UT, USA. Functional feeding responses of coyotes to fluctuating prey abundances in the Curlew Valley,
Utah, 1977-1993.
Predator-prey interactions were investigated in Curlew Valley, Utah by correlating prey abundances with prey consumption data sets. This extended

data series allowed us to evaluate coyote diet trends that encompassed

substantial fluctuations in abundance of various prey species. Previous studies reported a cyclic trend in jackrabbit density with a period of ten years
and .150-fold amplitude, as well as short-term fluctuations among some
rodent species that exceeded an 8-fold difference in amplitude over twoyear periods. Our results suggest changes in coyote diets mainly reflect the
fluctuations in jackrabbit abundance. Prey switching to rodents during periods of low jackrabbit abundance was also evident. Our main objective
was to assess coyote functional feeding responses to these changing prey
populations. We used the initial feeding pattern analysis to compare prey
consumption data to prey abundance. Our results show coyotes respond to
jackrabbit abundance with a Type II curve. Knowledge of coyote-prey interactions can aid in understanding coyote food preferences, population
density, habitat selection, and depredation rates. This information is useful
in predicting future predator-prey interactions and evaluating management
options.
BARTON, KASEY E.* and M. DEANE BOWERS. Department of Ecology and Evolution, University of Colorado, Boulder, CO. Effects of competition on plant size and secondary chemistry in Plantago.
Plants experience multiple biotic interactions simultaneously, although
most studies examine interactions in isolation. The goal of this study was
to test the effects of competition, both interspecific and intraspecific, on
the anti-herbivore defense in Plantago lanceolata and P. major. The experiment had a 2x3x2 factorial design in which the factors were focal
species (2), neighbor species treatment (3), and harvest time (2). Each
replicate consisted of a single plant of either P. lanceolata or P. major
planted in the center of the pot (focal species) and surrounded by 4 individuals of either P. lanceolata or P. major, or surrounded by no neighbors
(neighbor species treatment). Harvested shoots were dried, weighed, and
analyzed for iridoid glycoside concentrations. Plants grown without neighbors were significantly larger than those with neighbors (F*2,170 5 103.60,
p,0.0001). The two species differed in their competitive ability as neighbors: focal plants with P. major neighbors were, on average, significantly
larger than those with P. lanceolata neighbors (F*1,170 5 8.18,
p50.0048). Focal plant species and harvest date were reliable predictors
of iridoid glycoside concentrations. Plantago lanceolata had significantly
higher concentrations of iridoid glycosides than P. major plants (F*1,141
5 78.68, p,0.0001). Iridoid glycoside concentrations were higher in plants
harvested in week 10 than those harvested in week 7 (F*1,141 5 44.89,
p,0.0001). Iridoid glycoside concentrations were not influenced by the
presence of neighbors. These results are discussed in light of evidence that
nutrient availability influences plant defensive chemistry.
BARTUSZEVIGE, ANNE M. and DAVID L. GORCHOV. Miami University, bartusam@muohio.edu, gorchodl@muohio.edu, Oxford, OH, USA.
Community and landscape features that influence the invasion of Lonicera maackii.
Although invasive plants are recognized as a major ecological problem,
little is known of the role of dispersal agents and landscape structure on
the invasion process. Lonicera maackii (Rupr.) Herder (Caprifoliaceae), a
bird-dispersed shrub introduced to the United States in 1898, has become
invasive in 24 eastern U.S. states. As part of our investigation of how the
invasion of this species is shaped by landscape features and by movement
patterns of the birds that disperse its seeds, we explored the relationship
between its distribution and woodlot characteristics at an invasion front in
southwest Ohio. During summer 2002, 11 woodlots were sampled using
the point-quarter method. Basal areas, densities, and importance values of
shrub, sapling, and tree species were calculated. For each woodlot, we also
calculated mean gap fraction (canopy openness) using a Licor LAI 2000.
Digital Ortho Quarter Quad photos and ArcView GIS were used to collect
landscape parameters from the same woodlots. Stepwise multiple regression analysis was used to determine the community and landscape factors
that best explain L. maackii density. The best predictor of woodlot L.
maackii density was the density of L. maackii on the perimeter of the
woodlot (R2 5 0.897). This perimeter density, in turn, was best predicted
by the gap fraction of the woodlot (R2 5 0.334) and number of corridors
connecting to other woodlots (R2 5 0.212). Additional woodlots will be
sampled in 2003, but the results to date suggest that both community fac-

tors (e.g. canopy openness) and landscape factors (connectivity) are important in the invasion of L. maackii.
BARTZ, TRACI M., TIMOTHY P. DECHANT and KATHLEEN L.
SHEA.* St. Olaf College, Northfield, MN. Size and growth patterns of
trees in a maple-basswood forest restoration after twelve years.
Growth patterns and survival of trees in a restored maple-basswood forest
in southeastern MN, USA, were examined over a twelve year period. Data
were collected on heights and diameters of 1,052 individually tagged trees
planted in old fields. Tree size, growth curves, percent mortality and effects
of plastic tree shelters were compared among species. Trees with tree shelters were significantly taller than trees without shelters after two years of
treatment, but after eleven years there was no significant difference between
trees with and without shelters. Tree mortality rates ranged from 9.9% in
Acer rubrum, 17.6% in Tilia americana and 30.0% in Quercus macrocarpa
to 50.7 % in Acer saccharum. The growth pattern of most species followed
a linear increase in height over time, nearly doubling in height during the
last three years. The mean heights of the tallest species, Quercus rubra,
Fraxinus americana and Tilia americana, were 4-5 m while the shorter
species, Juglans nigra and Acer saccharum, had mean heights from 2-3 m.
Results suggest use of protective tree shelters is not recommended after the
first three years unless herbivory levels are very high. Species with lower
light tolerance, such as Acer saccharum, will become a more important
component of the forest as trees reach reproductive age and the canopy
closes over.
BARUCH, ZDRAVKO,1,* JAFET NASSAR,3 JOSE BUBIS2 and KARIvar, Caracas, Miranda, VeMAR TORREALBA.1 1 Universidad Simon Bol
ficas,
nezuela, Venezuela; 2 Instituto Venezolano de Investigaciones Cient
var, CaCaracas, Miranda, Venezuela, Venezuela; 3 Universidad Simon Bol
racas, Miranda, Venezuela, Venezuela. Genetic, quantitative, environmental and geographical distances among populations of a dominant
C4 grass from Neotropical savannas.
The Orinoco savannas in Venezuela are exposed to land use changes, contamination and biological invasions which compromise their persistence
and function. Trachypogon plumosus dominates the herbaceous matrix of
the savannas under diverse combinations of climate and soils, suggesting
substantial inter-population variation. We quantified isozyme and quantitative trait variation in nine populations of this grass and related estimates
to geographic and environmental features of sampled locations. Isozyme
diversity estimates were based on 10 polymorphic enzyme systems. Also,
16 quantitative traits were evaluated. Distance matrices for isozyme, quantitative traits, geographic and environmental data were subjected to cluster
analysis and ordination. Correspondence between isozyme and quantitative
trait variation and their relationships to geographic and environmental distances were analyzed with Mantel tests. Isozyme and quantitative trait diversity differed significantly among populations. About 28% of total isozyme variation occurred among populations. A significant association was
only detected between genetic and geographic distances; however, dendrograms generated for isozyme and quantitative traits were congruent with
environmental and geographic data and ecologically meaningful. The core
of lowland and highland populations were clearly defined leaving 2 - 3
populations without clear affinities. Our results support the genetic basis
of local population differentiation and the relatively large intra-specific diversity in T. plumosus. They also can be used to decide conservation strategies and select seed sources for future restoration programs.
BASHEY, FARRAH.1,2,* 1 Department of Biology, University of California,
Riverside, CA; 2 Department of Biology, Indiana University, Bloomington,
IN. Local adaptation and plasticity of offspring size in guppies.
Offspring number is closely tied to evolutionary fitness, yet many organisms produce relatively few offspring. Making fewer offspring can be adaptive if, by making fewer offspring, each offspring receives greater investment from the parent resulting in increased offspring fitness. In the guppy,
Poecilia reticulata, offspring size varies genetically across populations and
plastically in response to maternal condition. Across populations, variation
in the competitive environment faced by newborn guppies is potentially an
important source of selection on offspring size. Additionally, resource lim-

Abstracts

23

ited guppies produce larger offspring. I evaluate whether these effects of

the maternal environment increase offspring fitness in low resource environments. Furthermore, these plastic differences in offspring size are of the
same magnitude as the genetic differences between populations. I examine
the genetic basis of plasticity in two populations of guppies and evaluate
potential costs and limitations of plasticity that might favor local adaptation
over plasticity in this system. Current evidence suggests that the direct
benefits to offspring fitness of increasing offspring size may be compromised by links between plasticity and reproductive effort.
BASTOW, JUSTIN L.,* EVAN L. PREISSER and DON R. STRONG.
University of California-Davis, Davis, CA, USA. The effect of an invasive
grass, Holcus lanatus, on the breakdown of litter by Porcellio scaber.
Invasive species may alter ecosystem processes through their impact on
the density and behavior of resident species. Holcus lanatus is an invasive
perennial grass in the coastal prairie of California. At the Bodega Marine
Reserve, the dominant detritivore is the isopod Porcellio scaber. Porcellio
is twice as dense in Holcus, monocultures as in surrounding uninvaded
prairie. We hypothesized that Holcus may alter the rate of decomposition
through its effect on densities of Porcellio. We used a litter bag experiment
to determine the relative rates of litter break down in an invaded and uninvaded section of prairie. Litter was taken from Holcus, mixed grasses
and Lupinus arboreus, the dominant shrub in the system. All litter types
were transplanted into all three habitats in September of 2002. Half of the
litter bags had holes punched in them to allow the entrance of macrodetritivores, such as P. scaber. Half of the litter bags were retrieved in March
of 2003 in order to measure the rate of decomposition during the winter,
while half of the litter bags will be left until September 2003 to measure
annual decomposition. Lupine litter lost 50% more mass during the winter
than either resident grasses or Holcus. Litter loss in the Holcus invaded
prairie was 10% greater, but this habitat effect was independent of Porcellio. Although the isopod exclusion was effective, there was no difference
in over-winter litter loss between bags with and without isopod access.
This may be the result of low isopod activity during the winter.
BATCHELOR, MARGARET E.* and NORMA L. FOWLER. University
of Texas at Austin, Austin, Texas, USA. Consequences of including demographic heterogeneity in population models.
Demographic stochasticity and demographic heterogeneity have different
effects on the behavior of population models. Demographic stochasticity is
variation among individuals not due to differences in the underlying probabilities of survival, of reproduction, and of growth from one size class to
another. Demographic heterogeneity is variation among individuals in the
underlying probabilities of survival, reproduction, and growth. Three models were developed. One model had neither demographic stochasticity nor
demographic heterogeneity. The second model had only demographic stochasticity. The third model had both demographic stochasticity and demographic heterogeneity. The addition of demographic stochasticity alone
increased the mean population growth rate (l) at the smallest of three
(reset) population sizes. The addition of demographic heterogeneity to a
model already containing demographic stochasticity did not further change
mean l, which was a consequence of the particular constraints imposed on
these models. However, the addition of demographic heterogeneity to a
model already containing demographic stochasticity increased the variance
of l, which in turn increased the probability of extinction. This occurred
because the addition of demographic heterogeneity also increased the variation in population size. At very small population sizes, an increase in the
variation in n, from adding either demographic stochasticity or demographic heterogeneity to the model, tended to increase mean l but also to increase the probability of extinction.
BATTAGLIA, LORETTA L.1,* and JULIE S. DENSLOW.2 1 Southern Illinois University Carbondale, Carbondale, Illinois; 2 USDA Forest Service,
Institute of Pacific Island Forestry, Hilo, Hawaii. Sapium sebiferum invasion of a forest-marsh complex in the Louisiana deltaic plain.
Spread of the exotic Sapium sebiferum (Chinese tallow) threatens the integrity of wetland ecosystems along the Atlantic and Gulf coasts of the
Southeast. High fecundity and tolerance to shade, flooding, and salinity

24

Abstracts

enable it to invade wetland forest and marsh alike. Structural and compositional characteristics of native vegetation, underlain by hydrologic and
disturbance patterns, are thought to determine relative invasibility of these
communities. Determination of differences in invasibility will facilitate prediction of spatial spread, early detection, and development of management
plans that prioritize areas at greater risk. In Summer 2001, we conducted
ground surveys for tallow abundance in the wetland forest, freshwater
marsh, and forest-marsh ecotone at Jean Lafitte National Park in coastal
Louisiana to determine the pattern of tallow distribution and its relationships with other species. We also interpreted and analyzed color infrared
aerial photographs (1:8000) taken in Autumn 2001. We used GIS to determine the associations among native vegetation types, anthropogenic features, and tallow occurrences and to identify factors that are predictive of
tallow presence. Tallow was common in swamp forests where understory
cover is sparse. Along midslopes and hardwood ridges, its distribution is
patchy and often associated with canopy and understory gaps. Tallow in
the marsh is restricted to floating shrub thickets, Taxodium distichum (bald
cypress) islands in the marsh, and hummocks at the forest-marsh ecotone.
Spoilbanks along canals and roadsides support dense linear stands of mature tallow and thus provide an abundant seed source with a long invasion
front. Vegetation that is highly invasible and in close proximity to an exotic
seed source should be a priority for control measures. Color infrared aerial
photography is a useful tool for locating patches of tallow in the native
forest canopy and isolated individuals in herbaceous marsh and shrub thickets, where it forms an overstory above the native canopy.
BATTAGLIA, MICHAEL A.,1,2,* WAYNE D. SHEPPERD,1 KURT H.
MACKES3 and LINDA A. JOYCE.1 1 USDA Forest Service, Rocky Mountain Research Station, Fort Collins, CO; 2 Management and Engineering
Technologies International, El Paso, TX; 3 Colorado State University, Fort
Collins, CO. Stocking conditions influence strength of small diameter
ponderosa pine wood harvested in forest restoration treatments.
Treatment to reduce risk of catastrophic fire in ponderosa pine forests of
the Colorado Front Range requires removal of large numbers of small diameter trees. Many of these trees have grown in highly competitive environments and it is unknown what effect these conditions have on strength
properties of these trees. We measured strength properties of 9 to 11 inch
diameter open grown (n515), suppressed (n520), and mistletoe infected
(n521) ponderosa pine trees. Strength properties were compared to the
basal area increment (BAI), heartwood-sapwood ratio and forest density
surrounding each tree. Growth rates of open grown trees were 50% higher
than suppressed and mistletoe trees, while age and heartwood-sapwood
ratio were lower in open grown trees. Modulus of rupture (MOR) was
similar for suppressed and mistletoe trees, but lower for open grown trees.
Modulus of elasticity (MOE) was highest in the suppressed trees and lowest
in open grown trees. MOR and MOE increased with increasing stand basal
area and tree ring density, but decreased with increasing individual stem
BAI. Different competitive environments impact growth rates, which influence the strength of small diameter wood. Small diameter trees removed
from dense Front Range forests appear to be superior in strength than
rapidly growing young trees of the same size.
BATZLI, JANET M.,* ANN BURGESS, EVELYN HOWELL and CURT
CASLAVKA. University of Wisconsin-Madison, Madison, WI. Teaching
ecology through prairie restoration: The Biocore Prairie at University
of Wisconsin-Madison.
As part of an introductory honors biology curriculum at the University of
Wisconsin-Madison (Biocore) students have become actively involved in a
long-term ecological experiment involving a 5-acre prairie restoration begun in 1997 in an old agricultural field on the edge of the campus. Each
fall students gather data, analyze results, make recommendations for further
experiments, and communicate their findings to the next cohort of students
that implement the design. In fall 2002, students collected plant species
richness data for areas that had undergone one of three types of site preparation regimes prior to planting 3 to 5 years ago; herbicide, mulch, or
rototilling. These data were compared to those collected in Curtis Prairie,
a 67 year old established restoration. Students found a very low frequency
of prairie species in the Biocore Prairie and no statistical difference in weed
vs. prairie plant composition among the three site preparation techniques.

While both the Biocore Prairie and Curtis Prairie had the same total number
of species, only 28% of the species were common. These results forced
students to generate an alternative restoration plan focusing on a mixedmethod site preparation. They also proposed studies on the efficacy of
sawdust for soil microsite preparation and a mixed planting regime using
both seed and seedlings. Students in this program have actively applied
their understanding of the complex ecological interactions encountered at
the prairie, have managed and analyzed large data sets, and have considered
the many variables involved in designing a novel restoration plan for the
site. Moreover, students have taken ownership of the site and ownership
of their own learning resulting in great motivation to apply ecological concepts in a concrete way. Introduction of this project to a student body
dominated by pre-medical professionals has been an exciting and effective
way to teach ecology.
BAUERLE, WILLIAM L.1,* and TARYN L. BAUERLE.2 1 Clemson University, Clemson, SC; 2 Penn State University, University Park, PA. Modeling leaf conductance of Acer rubrum L. ecotypes from contrasting
hydrological habitats: Integrating biochemical and hydraulic signaling.
Recent evidence in an herbaceous system indicates that chemical and hydraulic stomata controls are a function of the combined responses of both
above and below ground environments. To date, nearly all woody plant
models used to study atmospheric and subterranean water stress treat the
above and below ground environments separately. In woody plant systems,
however, the significance of the link between chemical and hydraulic control on regulating stomatal conductance (gs) is still unclear. To examine the
effects of atmospheric and subterranean water stress on leaf chemical and
hydraulic signals, we modeled stomatal water loss in response to gas exchange function. We test the hypothesis that chemical, hydraulic, and aerial
gs controls act sequentially. In an attempt to quantify the combined control
of gs in Acer rubrum L., a woody species known to radiate into contrasting
hydrologic extremes, two existing woody plant data sets were used. The
presence of intraspecific variation between wet and dry site red maple
populations in physiological response to atmospheric vapour pressure deficits and soil water stress in a previous study provided the impetus to
compare combined gs control models to more common empirical aerial
environment models in a woody plant system. Coupling leaf water potential
(CL) and bulk leaf abscisic acid (ABAL) in a combined model of stomatal
response to leaf and soil environment improved the g s prediction over
common aerial empirical models. The results support the hypothesis that
ABAL and CL may act sequentially and, based on the results, ecotypic
variation in stomatal responsiveness to air humidity is likely mediated by
ABA accumulation in leaf tissue. The results also suggest that growth environment can influence the gs response. Based on the findings, more research into physiological integration of biochemical and hydraulic signaling
is warranted.
BAXTER, COLDEN V.,1,* KURT D. FAUSCH1 and MASASHI MURAKAMI.2 1 Colorado State University, Fort Collins, CO; 2 Hokkaido University
Forests, Tomakomai, Takaoka, Japan. Nonnative stream fish invasion interrupts reciprocal subsidies that shape stream and forest food webs.
Results of a large-scale field experiment demonstrated the importance of
reciprocal subsidies to stream and forest communities, and showed that
rainbow trout Oncorhynchus mykiss invasion can lead to significant changes in both of these connected systems. During summer 2002, the effects of
invading rainbow trout, and a mesh greenhouse cover that excluded terrestrial prey, were manipulated in a replicated (n54 each) factorial design in
16 fenced reaches of a small stream in Hokkaido, Japan. In control reaches,
native Dolly Varden charr Salvelinus malma foraged substantially on terrestrial as well as aquatic insects. Adding competitively dominant rainbow
trout caused charr to shift to foraging on herbivorous benthic insects, thereby increasing stream periphyton biomass in an archetypal trophic cascade.
Furthermore, decreased biomass of aquatic insects emerging to the forest
resulted in fewer riparian-specialist spiders. Adding the greenhouse reduced
input of terrestrial insects, which led to a similar foraging shift by charr,
and subsequent increased periphyton biomass, decreased emerging insect
biomass, and fewer riparian-specialist spiders. Finally, adding both the
greenhouse and rainbow trout produced the same results. These results
indicate that rainbow trout had as large an effect as excluding terrestrial

prey, and that the effects occurred rapidly, reaching a similar threshold of
low herbivore and high periphyton biomass within about 6 weeks. The
results also suggest that rainbow trout can cause an interaction modification indirect effect, by changing native charr foraging behavior, that cascades to stream periphyton, emerging insects, and riparian consumers.
BAZZAZ, FAKHRI A.* and KELLY S. WOLFE-BELLIN. Harvard University, Cambridge, MA, USA. Assemblages of annuals exhibit nonlinear responses to gradients of elevated air temperature and atmospheric
CO2.
Two abiotic global change factors, air temperature and atmospheric CO2,
are expected to have considerable impact on future ecosystem function by
reducing biodiversity. We conducted a study in which assemblages of nine
annual species, representing three functional groups (C3, C4, and legumes),
were exposed to gradients of either elevated air temperature alone or a
combination of elevated air temperature and atmospheric CO2. A relatively
novel approach was employed, in which greenhouse tunnels were used to
create the two environmental gradients. All plant assemblages were exposed to an air temperature gradient that ranged from ambient to approximately 38C above ambient during daylight hours. Additionally, half the
plant assemblages were exposed to a CO2 gradient that ranged from 370
to 810 ppm. Species-specific, non-linear responses to the two gradient treatments were evident in plant height and biomass. In particular, Ambrosia
artemisiifolia, a C3 species, exhibited little response to increasing air temperature alone, but exhibited a strong positive response that quickly reached
an asymptote when exposed to increasing air temperature and CO2. In fact,
a strong shift in species composition was evident by the end of the study.
Two species of Setaria, both C4 grasses, dominated assemblages under
conditions of ambient CO2 and high temperature, while A. artemisiifolia
dominated assemblages grown under conditions of high CO2 and high temperature. These results demonstrate that it is important to consider both
elevated air temperature and atmospheric CO2 when conducting experiments designed to predict ecosystem responses to global change, and that
plants may exhibit important nonlinear responses to gradients of both factors.
BEARD, KAREN H.,1,* KRISTIINA A. VOGT, DANIEL J. VOGT,2
FREDERICK N. SCATENA,3 ALAN COVICH,4 RAGNHILDUR SIGURDARDOTTIR,5 THOMAS C. SICCAMA5 and TODD CROWL.1 1 Utah
State University, Logan, UT, USA; 2 University of Washington, Seattle,
WA; 3 University of Pennsylvania, Philadelphia, PA; 4 Colorado State University, Fort Collins, CO; 5 Yale University, New Haven, CT. Structural,
functional characteristics and resilience of forest ecosystems.
We conducted an experimental manipulation in the Luquillo Experimental
Forest, Puerto Rico, to test hypotheses on the effect of hurricane disturbance on ecosystem processes. Our experimental treatments consisted of
wood addition, removal and control plots, because wood addition is a major
consequence of hurricane disturbance. Our ecosystem indicators included:
tree growth by species, litterfall, leaf litter decomposition rates, and root
turnover. We found that the effect of coarse wood in ecosystem resilience
varied across the Luquillo landscape. For example, in one site there was
no influence on tree species growth rates by removing or adding coarse
wood while in two other sites the removal of wood resulted in decreased
growth rates and the addition of wood increased their annual productivity.
We found that part of the variability measured in ecosystem response to
disturbances and their recovery rate were controlled by legacies that maintain a stronger constraint on the functioning of the system than that imposed
by the disturbance. For example, stream geomorphology controls the type
of riparian zone produced and created distinct legacies. Legacies are also
produced by soil chemistry, coarse wood, and individual plant life history
strategies.
BEAUCHAMP, VANESSA B.* and JULIET C. STROMBERG. Arizona
State University, Tempe, AZ. Cottonwood-willow stand structure on
regulated and unregulated reaches of the Verde River, Arizona.
Cottonwood (Populus sp.) and willow (Salix sp.), the dominant overstory
species in western riparian forests, are disturbance-adapted species with
narrow germination windows. Changes to flood cycles often lead to a de-

Abstracts

25

crease in recruitment success and survival of these species. This research

investigates the effects of damming on the flow regime of a river managed
for urban and agricultural water supply, and on the structure and composition of riparian cottonwood-willow forests downstream from the dam.
Fifty-five years of stream gage data were used to compare flow regimes on
unregulated and regulated reaches of the Verde River, in central Arizona.
The species composition, stem density and basal area of cottonwood (P.
fremontii) and willow (S. gooddingii and S. exigua) dominated stands were
compared in above and below dam reaches. Dam operation has decreased
peak flows and flow variability, shifted the timing of high flows, and increased summer base flows. However, regulated reaches along the Verde
still experience spring floods in very wet years, allowing for periodic cottonwood and willow regeneration. Cottonwood and willow stem density
was not different between above and below dam reaches in sapling (1-10
year) and mature (11-54 year) stands, but cottonwood stem density in oldgrowth (55+ year) stands was higher in unregulated reaches (P , 0.01).
Flow regulation has altered other attributes of the riparian vegetation. For
example, some measures of tree and shrub richness varied between reach
types. Also, stem density of salt cedar (Tamarix ramosissima), a stresstolerant, reproductively opportunistic species, was higher in regulated
reaches, for sapling classes only (5.82 6 2.15 stems/m2 vs. 0.03 6 0.03
stems/m2, P , 0.001).
BECKAGE, BRIAN,1,* BILL PLATT3 and LOU GROSS.2 1 University of
Vermont, Burlington, Vermont; 2 University of Tennessee, Knoxville, Tennessee; 3 Louisiana State University, Baton Rouge, Louisiana. Modeling
overstory dynamics of pine savannas: responses to climate and largescale disturbances.
Southeastern pinelands can vary from nearly open stands with few overstory trees to dense stands with closed overstories. Fire and hurricane disturbances affect overstory structure, with both the frequency of disturbance
and interactions between disturbances influencing overstory density. Fire
and hurricane disturbance regimes are also likely to change with global
warming, affecting the structure of pine overstories. We explore the sensitivity of overstory density to variation in disturbance regimes and to interactions between disturbances using matrix models and more complex
spatial simulation models. In the matrix model approach, we assume that
stands are in one of three states: densely packed trees with a closed overstory (forested state), widely spaced trees with an open overstory (savanna
state), or completely open with no trees (grassland state). Transitions between these states are determined by disturbance, i.e., fire, hurricane, fire
plus hurricane, or none. The state of the stand can, in turn, effect the
likelihood of fire disturbances. In the second modeling approach, we use
an individual-based model of pine dynamics that follows the fate of every
individual seed, seedling, and tree and that explicitly models demographic
processes as a function of disturbance. We use both approaches to develop
hypotheses regarding mechanisms that regulate pineland overstories. These
mechanisms are useful in developing concepts and methods of restoration,
as well as in predicting changes in stand structure in response to altered
disturbance regimes associated with global warming.
BECKSTEAD, JULIE,1,* INGRID M. PARKER2 and HAIVAN V. NGO.2
Gonzaga University, beckstead@gonzaga.edu, Spokane, WA; 2 University
of California, Santa Cruz, Santa Cruz, CA. Testing the Natural Enemies
Hypothesis: is invasiveness explained by a release from soil-borne pathogens?

The Natural Enemies Hypothesis (i.e., introduced species experience release from their natural enemies) is a common explanation for why invasive
species are so successful. We tested this hypothesis for Ammophila arenaria (European Beachgrass), an invasive plant of the coastal dunes in California. European research on A. arenaria in its native range has established
that soil-borne pathogens, primarily nematodes and fungi, reduce the
growth of A. arenaria. In a series of greenhouse experiments designed to
parallel European studies, seeds and two-week old seedlings were planted
in field collected soil that was sterilized (free of pathogens) and non-sterilized. Our first experiment found that the degree of biomass reduction
caused by pathogens was indistinguishable between the previously reported
native site in the Netherlands and our introduced site in California, failing
to show a demographic release. However, a partial escape from enemies

26

Abstracts

was found; fungal pathogens were present (including Fusarium spp.), but
pathogenic nematodes were functionally absent from the California site. A
second experiment found variability among eight coastal sites in California
with respect to a demographic release, and this variability was accompanied
by differences in the assemblage of pathogens. Because the predictions of
the Natural Enemies Hypothesis depend critically on whether the dominant
enemies are specialists or generalists, we performed studies on the host
range of the dominant fungal pathogens. It appears that soil-borne pathogens in the coastal dunes are host-generalists and exploit A. arenaria as
well as other co-occurring dune species. Thus, although escape from enemies is frequently cited as an explanation for the invasive nature of invasive
species, we found that a release from soil-borne pathogens does not explain
the invasion success of A. arenaria in its introduced range.
BELK, MARK C. and JOSEPH H. HANKS.* Brigham Young University,
Provo, UT. Variation in drift patterns of endangered larval June suckers in the lower Provo River.
The population of June suckers (Chasmistes liorus), which is endemic to
Utah lake and the lower Provo River, has experienced a dramatic decline
in young-of-year recruitment for the past forty years. The Utah lake population of the Utah sucker (Catostomous ardens), a similar yet more widespread species, is also experiencing this problem. To better understand the
fate of newly spawned June suckers, the Utah Division of Wildlife Resources has been collecting data on drift patterns of sucker larvae in the
lower Provo River since 1997. We described patterns of larval drift for
June and Utah suckers in the Provo River and compared patterns between
species. Of the six years in which data were collected, the drift pattern of
Utah sucker larvae in several of those years exhibited a strong trend toward
the deepest, swiftest parts of the depth/flow rate gradient. In those same
years, June sucker larvae were fairly evenly distributed across the gradient,
with a slight trend toward shallower, slower-moving waters. Although these
temporal patterns of drift did not vary within years, they did vary widely
among years. Understanding these patterns of larval drift will help in conservation of this endangered species.
BELL, CHRISTOPHER J.1,* and JACQUES A. GAUTHIER.2 1 The University of Texas at Austin, Austin, TX; 2 Yale University, New Haven, CT.
North American Quaternary Squamata: Re-evaluation of the stability
hypothesis.
The last four decades saw a dramatic increase in knowledge of Quaternary
squamate reptiles in North America. Initial research centered on primary
descriptive accounts of faunas, but these were recently supplemented by
synthetic regional summaries, and efforts to place the reptiles in a broader
context of faunal dynamics in the Pleistocene and Holocene. The picture
that emerged from these studies suggests that squamate reptiles were taxonomically and geographically stable throughout much of the Quaternary.
In stark contrast to the mammals and birds, few extinctions and no significant geographic range adjustments are reported for reptiles at the end of
the Pleistocene. Phylogenetic systematics has yet to play an important role
in Quaternary vertebrate paleontology, but it presents a potentially fruitful
alternative philosophical and methodological approach. Our recent re-evaluation of the stability hypothesis was based on this approach and leads to
several important conclusions: 1. This hypothesis suffers from circularity
because many fossil identifications ultimately are based on the modern
geographic distribution of taxa; 2. Due to their incomplete nature and our
current understanding of morphological synapomorphies for squamates,
species-level taxonomic resolution is not possible for the majority of Quaternary fossils; 3. Geographic and taxonomic stability is demonstrable at
more-inclusive taxonomic levels, but is not particularly informative with
respect to questions typically asked by Quaternary paleontologists; 4. To
answer species-level questions, we must seek synapomorphies in the isolated skeletal elements commonly preserved in Quaternary deposits. Utilization of phylogenetic systematics in Quaternary studies will require an
effort to seek synapomorphies in materials typically recovered from Quaternary deposits, acceptance of the taxonomic limitations of the data with
which Quaternary paleontologists work, and a willingness to frame questions in a phylogenetic context.

BELL, DANIELA L. and LAURA F. GALLOWAY. University of Virginia,

Charlottesville, VA, dlbell@virginia.edu. Differentiation in patterns of
phenotypic plasticity among populations of Geranium carolinianum.
Phenotypic plasticity, the change in trait expression in response to the environment, may enhance reproduction and survival in a changing environment. Plasticity may be favored under variable environments while nonplastic responses may be favored under spatially uniform environments.
Geranium carolinianum, an herbaceous annual plant, occurs in wood margins that are spatially variable for light and in old fields that are less variable for light. In a field common garden study with two light environments,
I examined if wood margin populations demonstrate greater plasticity for
morphological traits than old field populations. Three wood margin populations were compared with three old field populations. Surprisingly, wood
margin plants were significantly less plastic for petiole length, internode
length, and total plant height than were old field plants. Plants from wood
margins had significantly lower fitness in low light relative to high light.
In contrast, old field plants had constant fitness across both light environments. Selection analyses revealed significant directional selection on petiole length and selection differed significantly between the two light environments. These patterns suggest divergence for plasticity between wood
margin and old field populations. Petiole length may be a shade-avoidance
trait: the elongation of petioles may allow plants to escape shading by
neighbors. Old field populations of G. carolinianum may experience substantial shading by neighboring herbaceous plants. Wood margin sites, in
contrast, are characterized by shade from trees and other large woody
plants. Responding to shade from trees by elongating petioles may be costly: fitness may be reduced because resources have been used to alter trait
expression but the shade may not be avoided. Thus, wood margin plants
may experience selection against plasticity while in old field sites selection
favors plasticity.
BELOTE, TRAVIS* and JAKE WELTZIN. University of Tennessee,
Knoxville, TN, USA. Forest understory community responses to elevated CO2.
Rising levels of atmospheric carbon dioxide (CO2) are predicted to change
patterns and processes of native ecosystems in the near future. We are
examining species composition and ANPP of an understory plant community in ambient and elevated CO2 treatments in an ongoing, free-air CO2
enrichment (FACE) facility on the Oak Ridge National Environmental Research Park, Tennessee. Five 25-m diameter plots have received either ambient (5 control) or elevated (5 537 ppm) CO2 since 1998. The understory
plant community in these plots is dominated by several invasive plant species including Lonicera japonica and Microstegium vimineum. Results from
the 2001 growing season indicate total understory ANPP did not differ, but
species responses lead to changes in community composition and diversity
between ambient and elevated CO2. In contrast, ANPP was greater in elevated than ambient CO2 plots in 2002, but community composition or
diversity did not differ. Differences in community patterns of production
and composition were determined by the response of M. vimineum, which
was likely influenced by annual differences in precipitation. Specifically,
soil moisture was 20% less during the 2002 understory sampling time.
These results suggest that ecosystem and community responses to atmospheric CO2 may vary depending on species responses to other environmental factors.
BENAVIDES, MELISSA A.,* MEGAN J. OSBORNE, DOMINIQUE
ALO and THOMAS F. TURNER. Department of Biology, MSC03 2020,
1 University of New Mexico, Albuquerque, NM, USA. Temporal genetic
variability in pelagic eggs in the endangered Rio Grande silvery minnow.
The Rio Grande silvery minnow (Hybognathus amarus) exhibits a rare life
history feature for a freshwater fish in that it spawns pelagic eggs and
larvae. Consequently, much of the annual production of offspring is moved
downstream with river currents after spawning. This life history may suggest that juveniles and adults historically migrated upstream in attempt to
return to their natal sites. The Rio Grande is now highly fragmented by
dams and other diversion structures that impede upstream movement of
fish. We have undertaken a long term study of temporal and spatial genetic

variation in silvery minnow, and have shown that genetic effective size Ne
is three orders of magnitude smaller than census size. The cause of low
Ne/N appears to be associated with high variance in reproductive success
among individuals, exacerbated by annual loss of reproductive output to
entrainment through diversion dams. Loss of production would exert maximal effect on variance in reproductive success if genetic diversity of eggs
is structured spatiotemporally, such that groups of genetically related eggs
are subject to mortality that varies in space and time. We tested whether
genetic diversity of eggs collected at a single location from natural spawning varied among temporally-spaced samples. Preliminary genetic screening of eggs using mtDNA gene markers, SSCP, and nucleotide sequencing
suggests strong temporal variability among egg samples, implicating high
variance in reproductive success as an important factor for lowering genetic
diversity in wild populations of silvery minnow.
BENBOW, M. ERIC,1,* SCOTT L. ROLLINS2 and RICHARD W. MERRITT.1 1 Department of Entomology, Michigan State University, East Lansing, MI, USA; 2 Department of Zoology, Michigan State University, East
Lansing, MI, USA. Road salt effects on community structure and function in small wetland mesocosms.
Few studies have addressed the effects of road salt on the structure and
function of wetlands. The objectives of this study were to assess road salt
impact on primary production, Typha decomposition, and eukaryotic species composition in wetland enclosures. Mesocosms were treated with 0,
500, 2500, or 10,000 mg l-1 of road salt during the summer of 2002. Community structure changes were tested by assessing road salt effects on algal
communities after 9 d of exposure, and on invertebrate colonization of
Typha reed packs. Functional changes were measured by primary production and Typha reed decomposition (with and without invertebrates excluded). Primary production was impacted at 10,000 mg l-1, but significant
differences were not detected at lower salt concentrations. At 10,000 mg l1
, increased Typha mortality and phytoplankton production were observed.
After 9 days of exposure to elevated salt levels, only minor differences in
periphytic diatom species composition were evident at 10,000 mg l-1. Decomposition was measured with and without macroinvertebrates using of
cattail reed packs inside fine mesh screen. We found no significant effect
of macroinvertebrate exclusion on decomposition, but did find a significant
negative effect of road salt at 10,000 mg l-1 of road salt. Macroinvertebrate
colonization of reed packs was reduced by an order of magnitude at the
same concentration. Diptera made up the highest percentage of taxa in all
enclosures; however, this taxon was lowest at 10,000 mg l-1 road salt where
Gastropoda made up nearly an equal percentage. High concentrations of
road salt reduced macroinvertebrates, but these heterotrophs did not play a
role in reed decomposition. Thus, road salt appears to reduce reed decomposition either by affecting microbial communities directly, indirectly
through cascade effects associated with macroinvertebrate reductions, or
through alternative carbon availability resulting from salt effects on primary producers.
BENDA, NICOLE D.1,2,* 1 ndbenda@ncsu.edu, Raleigh, NC; 2 North Carolina State University, Raleigh, NC. Oviposition preference of a specialist moth Heliothis subflexa.
Locating appropriate host plants is an essential part of survival for herbivorous insects. This is especially important for specialist insects, whose
range of acceptable food plants is very narrow. There are ten times more
specialist than generalist insect species. Physiological, ecological, and environmental factors may constrain or give advantage to either range of host
plant use. Understanding the genetics and mechanics of host-finding can
be useful in explaining how specialization develops. Heliothis virescens
[Lepidoptera: Noctuidae] is an extreme generalist, feeding on plants in over
fourteen families. A close relative, H. subflexa, feeds solely on fruit of
Physalis species [Solanaceae]. Viable hybrids and backcrosses of these two
species can be produced. These offspring can then be analyzed using AFLP
markers and bioassays to identify QTL that affect oviposition preference.
The first step in this investigation is to identify the cues used for hostfinding by H. subflexa. The species of Physalis most preferred by H. subflexa for oviposition was unknown. A common garden experiment was
conducted using six species of Physalis as well as two outgroup species
(cotton and tomato). Oviposition on the different species was monitored

Abstracts

27

throughout the summer by inspecting the plants regularly for eggs. Phenology of the plants, evidence of attack by H. subflexa and by other herbivores was also recorded to examine the effects of these factors on H.
subflexa oviposition. The H. subflexa population was also monitored using
pheromone traps. P. pubescens had the highest number of eggs, followed
by P. heterophylla, P. angulata, the wild variety of P. philadelphica, and
P. cordata. The data collected on P. pubescens was analyzed further to
examine how phenology and herbivory affected oviposition preference. Extracts of the most preferred species were made and await bioassays. Future
goals of this research will be discussed
BENJAMINS, MAIJA E.1,* and DONALD J. LEOPOLD.2 1 SUNY- College of Environmental Science and Forestry, mebenjam@mailbox.syr.edu,
Syracuse, NY; 2 SUNY-College of Environmental Science and Forestry,
dendro@mailbox.syr.edu, Syracuse, NY. Habitat management recommendations for the endangered Karner blue butterfly based on its oviposition preferences.
Oviposition preferences of the Karner blue butterfly, Lycaeides melissa
samuelis Nabokov (Lepidoptera: Lycaenidae), were studied in the pine barrens of eastern New York. L. m. samuelis oviposite on blue lupine, Lupinus
perennis, the obligate food source of the larvae. In the eastern New York
pine barrens, L. perennis is often interspersed with Quercus ilicifolia and
Pinus rigida. Most conservation management techniques have focused on
restoring the eastern NY pine barrens to an early successional stage by
eliminating the woody plant cover. It has been demonstrated elsewhere that
larval growth of L. m. samuelis was significantly faster on L. perennis
grown in moderate shade than in full sun. By reaching pupation more
quickly, larvae avoid senescing L. perennis during the second brood, as
well as decreased rates of predation and parasitism. Do ovipositing L. m.
samuelis select for L. perennis in moderate shade to increase the survival
of their offspring? Does area of the L. perennis play a role in this selection?
Larvae and ovipositing L. m. samuelis were exposed to various sized L.
perennis in full sun and two levels of shade (30% and 50%). Results for
two growing seasons show that ovipositing females lay eggs more often
on L. perennis in moderate shade, independent of host plant density and
proximity to the center of the patch. Plant area and stem length also are
significant factors in selection of a host plant, with small and moderate
sized L. perennis having the highest number of eggs and larva. Plant characteristics selected as most favorable oviposition sites were used with a L.
perennis population data set collected in response to various management
techniques on 25 Right-of-ways in eastern NY. Habitat management methods for this study that produce similar sized L. perennis and maintain habitat heterogeneity inherent to the natural pine barrens ecosystem will be
discussed.
BENNETT, ASHLEY B.,1,* CLIFFORD S. SADOF2 and LAWRENCE M.
HANKS.3 1 University of Illinois, Urbana-Champaign, Il; 2 Purdue University, West Lafayette, In; 3 Univerisity of Illinois, Urbana-Chamapign, Il.
Manipulating floral density in ornamental landscapes to encourage
natural enemies of herbivorous insects.
A principle tenet of conservation biological control is that some natural
enemies feed on floral resources and so can be encouraged by providing
them flowers. Our study focuses on the effect that floral density has upon
population regulation of the plant feeding pest pine needle scale. We planted four species of perennial flowering plants (white clover, goldenrod, euphorbia, and coreopsis) around pine trees that were infested with the herbivores. Three densities of flowers were used: no flowers, a low density of
flowers, and a high density of flowers. Rates of predation and parasitism
were significantly greater in study plots having higher densities of flowers,
enhancing biological control of the pest.
BENNETT, ELENA M.,1,* STEPHEN R. CARPENTER,1 GARRY D. PETERSON,1 GRAEME S. CUMMING,2 MONIKA ZUREK3 and PRABHU
PINGALI.3 1 University of Wisconsin, Madison, WI; 2 University of Florida, Gainsville, FL; 3 Food and Agriculture Organization (FAO), Rome,
Italy. The role of ecology in global scenario development.
Human well-being depends on consistent provision of ecosystem services.
Yet ecosystems and the services they provide are changing, often in ways

28

Abstracts

that we cannot anticipate. The ubiquity of change in ecosystems explains

the frequency of surprise in ecosystem management. How can we manage
for surprises and uncertainties when we cannot predict them? One approach
is to make decisions that are robust to a number of different futures. Scenarios, sets of stories about the future, have been used in the business and
planning communities for decades as an alternative to single future planning methods. Scenarios have been used extensivly to examine global environmental futures. Global scenarios have successfully focused the worlds
attention on key global environmental issues; involved experts and stakeholders in discussions about the environment; and have been used to examine the prospects for world development. To date, however, most global
environmental scenarios have treated ecosystem dynamics as the result of
large scale drivers, and have not considered ecosystem feedbacks to largerscale processes. To the contrary, recent studies suggest that cross scale
ecological feedbacks and emergent properties of interacting subsystems can
and do serve as major drivers of global processes. Global scenarios could
greatly benefit from the input of ecologists to incorporate more realistic
ecosystem dynamics. Similarly, ecology could benefit from involvement in
scenario planning. Unlike many technical models, scenarios, easily understood as stories, can also be used for communication and outreach that
builds public appreciation of ecological science and the ecological dilemmas that society faces.
BENNETT, T. M. BULL.* South Dakota School of Mines and Technology,
Rapid City, SD, USA. Tracking buffalo from space: An analysis of
ecosystem change using remote sensed data.
The purpose of this paper is to report the landscape scale ecological changes that have occurred on a bison ranch in the Powder River Basin of Eastcentral Wyoming. Prior to westward expansion, the Powder River Basin of
Wyoming supported millions of bison (Bison bison bison) and were considered primary hunting grounds for many Plains Tribes. With the near
extinction of bison and subsequent introduction of domestic cattle and agriculture, and more recently mining and coalbed methane production, this
area has undergone extensive ecological change over the last century. Surplus bison from the Yellowstone Park herd were reintroduced onto a 60
thousand acre ranch (the Durham Ranch) in 1959 and populations have
flourished under various management strategies. The Holistic Resource
Management (HRM) strategy was introduced onto the ranch in 1985 and
has continued to present. The objectives for this study were to quantify
long-term short grass system response to bison grazing under a HRM regime using information derived from satellite based multispectral radiometry. Specifically, data on surface albedo and normalized difference vegetation index (NDVI) were gathered from 1985 to 2002 on a semi-arid shrub
steppe grassland system grazed by bison under the HRM strategy. The
results indicate that ecological conditions have changed under this management strategy, however, drought within these years may have influenced
the findings. Further studies that combine ground-based data with satellite
data will provide a clearer picture of ecological impacts across multiple
spatial scales.
BENNING, TRACY L.1,* and R. F. HUGHES.2 1 University of San Francisco, San Francisco, CA, USA; 2 Institute of Pacific Islands Forestry,
USDA Forest Service, Hilo, HI, USA. Quantifying the distribution and
spread of an invasive alien tree, Falcataria moluccana, across Hawaiian
landscapes.
Spread of invasive species is frequently cited as a major threat to the integrity of native ecosystems. In the Hawaiian Islands in particular, invasion
of alien tree species into wet montane tropical rainforests has become a
major management concern. Depending on the characteristics of the specific invader, these species have the potential to change ecosystem processes and alter successional trajectories of native vegetation. As a result,
we have initiated an investigation to document the distribution, rate of
spread and impact on canopy structure of the nitrogen fixing alien tree,
Falcataria moluccana (Albizia) into lowland forest habitats on the leeward
side of the Island of Hawaii. We acquired a time series of high-resolution
aerial photography collected in 1965, 1977, and 1993 for which we classified vegetation cover, land use and other physical attributes relevant to
ecosystem level processes (i.e. soil type, disturbance history, mean annual
precipitation) in a GIS environment. Classification of vegetation was con-

ducted using textural analyses. An initial accuracy assessment of this classification indicates an 87% success rate for identification of Falcataria
cover. Rate of spread and parameters associated with canopy growth were
measured for each time period and a change detection analysis was performed to document changes in extent of cover and growth of Falcataria
stands. Preliminary results indicate increases in Falcataria canopy cover
from 33 to 95% depending on substrate age and disturbance history. Results
obtained from this exercise will be coupled with process level data to determine and compare the factors that might constrain or enhance the spread
of Falcataria across the landscape.
BENNINGTON, CINDY.* Stetson University, DeLand, FL. Sex allocation
and stylar movement in Passiflora incarnata.
Andromonoecious plant species produce both hermaphrodite and male
flowers on individual plants. Male flowers presumably represent a resource
savings, and the frequency of their production is thus expected to increase
with resource limitation. Passionflower (Passiflora incarnata), an andromonoecious perennial vine, produces two floral morphs that are effectively
male. The first morph is similar to male flowers in other andromonoecious
species in that the pistil is atrophied. In the other male floral morph, however, the pistil is similar in size to that of hermaphrodite flowers, but the
styles are held erect where they are unlikely to come into contact with
pollinators. Hermaphrodite flowers deflex their styles to a position where
contact with pollen-carrying pollinators is likely. Other flowers partially
deflex their styles. In a field study in two Central Florida passionflower
populations during the summer of 2002, I categorized 226 flowers into four
classes: atrophied pistil, erect-style, intermediate, and hermaphrodite (i.e.,
those whose styles were held parallel to the flower petals). I asked whether
dry weight allocation to gynoecium, androecium and perianth differed
among the four floral morphs. In addition, I compared pollen deposition
on stigmas of open-pollinated flowers and ovule number among morphs.
As expected, the gynoecium weight of flowers with atrophied pistils was
9x less than that of hermaphrodite flowers. In addition, for all measured
traits except ovule number there was a predictable increase in trait value
with increased style movement. Thus, those flowers with "intermediate"
style position tended to be larger and receive more pollen than male flowers
and smaller and receive less pollen than hermaphrodite flowers (p , 0.05
in all cases). Ovule number, which differed significantly between the two
populations (F1, 74 5 36.65, p,0.001) did not differ among gender categories. Continued investigations will determine whether the degree of style
movement is related to the timing and/or the severity of resource limitation.

temporal window of prey availability, and the predictability of their respective food resources.
BENSON, GARY L.1,* and GORDIE H. REEVES.2 1 USDA Forest Service, Pacific Northwest Research Station, Portland, Oregon, USA; 2 USDA
Forest Service, Pacific Northwest Research Station, Corvallis, Oregon,
USA. A dynamic landscape approach to maintain and restore freshwater habitat for U.S. Pacific Northwest anadromous salmonids.
Populations of all Pacific salmon species (Oncorhynchus spp.) in the Pacific
Northwest (PNW) of the United States are in need of special management
consideration because of low or declining numbers. A major factor responsible for these declines is the loss of freshwater habitat quantity and
quality. To preserve and recover these populations, both long- and shortterm processes that create and maintain freshwater habitat must be protected and restored. Streams in the PNW are highly dynamic in space and
time, and lack of consideration of their dynamic nature has limited the
effectiveness of habitat management and recovery programs. Riverine-riparian ecosystems used by Pacific salmon are periodically subjected to
large disturbance events. These ecosystems then move through a series of
recovery states over time periods extending from decades to centuries.
Large natural disturbance events such as wildfires, floods, and debris flows
may result in the extirpation of local populations, followed by re-colonization from surrounding areas as more favorable conditions develop. Every
stream in a watershed cannot be expected to have good habitat and abundant fish populations at all times. Rather, natural landscapes consist of a
mosaic of patches in various successional stages, with conditions that are
favorable or less favorable to Pacific salmon. Spatial and temporal configuration of successional stages will vary widely within river basins and
across eco-regions. Over time, less productive streams can become more
productive and streams with more favorable conditions can transition to
less favorable conditions. Life-history attributes of Pacific salmon suggest
that they are adapted to a dynamic environmment. The long-term cycles
of aquatic ecosystems havent been adequately addressed in current management schemes. Management regimes should be based on maintaining
historical disturbance patterns and drainage systems to the extent possible,
in order to provide a landscape context similar to the conditions in which
salmonid populations evolved.

BENOIT-BIRD, KELLY J.* Hawaii Institute of Marine Biology, University

of Hawaii at Manoa, Kaneohe, HI. Sound-scattering layers and the foraging ecology of pelagic marine mammals.

BERG, ED,1,* DAVID HENRY2 and ANDREW DEVOLDER.3 1 Kenai

National Wildlife Refuge, Soldotna, AK, USA; 2 Kluane National Park,
Haines Junction, YT, Canada; 3 US Forest Service, Seward, AK, USA. A
250-year record of spruce bark beetle outbreak history on the Kenai
Peninsula, Alaska and Kluane National Park, Yukon Territory.

The foraging ecology of two pelagic predators, spinner dolphins in Hawaii

and dusky dolphins in New Zealand, was examined with respect to the
characteristics of the sound-scattering layers that serve as their prey. Active
sonar surveys were used to simultaneously assess the distribution of dolphins and the density and distribution of their prey. Scattering layers in
both Hawaii and New Zealand underwent a diel vertical migration. As a
result of these migration patterns, the potential prey of dusky dolphins was
within their depth range for longer each night than the prey of spinner
dolphins. The vertical distribution of both dolphin species closely matched
the diel vertical migrations of their prey over time. The mean depth of both
dolphin species was within a few meters of the depth at which the scattering
layers were most dense. The Hawaii scattering layer was much more heterogeneously distributed over horizontal space and reached much higher
densities than the New Zealand layer. Groups of spinner dolphins were
observed to actively aggregate patches of the layer while dusky dolphins
were not, perhaps because of the relative homogeneity of the scattering
layer in New Zealand. Spinner dolphins were found to exhibit very fixed
foraging patterns; they were nearly always found in pairs that were part of
very coordinated groups of up to 22 animals. Dusky dolphin group size
ranged from 1 to 5 animals in each coordinated group. The size of these
groups varied as a function of time of night, layer depth, and variance in
the layers density. I hypothesize that the differences in behavior observed
between the two species can be accounted for by the differences in their
degree of foraging specialization, their risk of immediate starvation, the

Both the Kenai Peninsula and southwestern Yukon Territory have experienced extensive outbreaks of the spruce bark beetle (Dendroctonus rufipennis Kirby) during the 1990s. When bark beetles thin a forest canopy,
surviving trees grow more rapidly for 60 to 80 years until the canopy recloses. We used dendrochronology to reconstruct bark beetle outbreak history at 23 Kenai Peninsula sites and at 4 sites near Haines Junction in the
Yukon Territory. On the Kenai we found tree-ring evidence of regional
canopy-thinning events during the 1810s to 1820s, 1870s to 1880s, 1910s,
and 1970s, which we attribute to spruce bark beetle outbreaks. Every Kenai
stand showed evidence of at least one and as many as five thinning events,
whereas only one Yukon stand showed evidence of thinning. For that Yukon stand we have historical documentation of a local 1940s outbreak, as
well as standing beetle-scarred trees with death dates from 1934-42. We
conclude that regional spruce bark beetle outbreaks have occurred on the
Kenai at intervals of 20 to 60 years over the last 250 years, and that no
regional outbreaks occurred in the Kluane area during this time period,
prior the 1990s. The Kenai outbreaks of the 1970s and 1990s followed runs
of unusually warm summers (1967-69, 1987-97, respectively). The Kluane
outbreak of the late 1990s followed the warm summers of 1989-95. Treering reconstructions of summer temperatures on the Kenai indicate that the
warm summers of the 1990s are the longest run of warm summers in the
last 350 years. Fire history studies on the Kenai have found no evidence
of fire following beetle-kill over the last 250 years, prior to the 1990s.

Abstracts

29

BERGER, MICHELLE L.* and GEOFFREY G. PARKER. Smithsonian

Environmental Research Center, Edgewater, MD. A multivariate analysis
of structural development in Tulip Poplar Association forests.
We combined measurements of the vertical distribution of leaves in the
canopy with more common stem measurements in a multivariate analysis
of structural development in Tulip Poplar association forests. Eighteen variables were measured or derived for a 240 year chronosequence of 46 forest
plots in the vicinity of the Smithsonian Environmental Research Center in
Edgewater, MD. With a factor analysis, the original variables were reduced
to three independent factors that together accounted for .82% of the variability in the dataset. Variables that change early in development, when
height growth is greatest and before crowns begin to differentiate, loaded
strongly onto the first factor. This factor was positively correlated (r2 5
0.68, p,0.0001) with age, particularly for forests younger than 70 years.
Variables that change as the canopy differentiates and the understory begins
to grow loaded onto the second factor. This factor was negatively correlated
(r2 5 -0.36, p50.014) with age. Only Leaf Area Index and total leaf biomass loaded strongly onto the third factor, which was not significantly
correlated with stand age. We conclude that: 1. Common measurements of
stem height and diameter used in many studies of forest development do
not capture all of the structural changes that occur, particularly in older
forests. 2. The only leaf variables usually measured, LAI and leaf biomass,
do not change much during development in this forest type. Additionally,
the changes in the vertical distribution of leaves within the canopy can
have clear implications for how a forest functions as it ages.
BERGNER, BELLE,1,* JILL JOHNSTONE2 and KATHLEEN K. TRESEDER.1 1 University of Pennsylvania, Philadelphia, PA; 2 University of
Alaska-Fairbanks, Fairbanks, AK. Experimental warming increases soil
CO2 flux but not microbial abundance in a recently burned Alaskan
boreal forest.
Boreal forest soils comprise a large portion of the terrestrial global carbon
pool and are currently a net carbon sink, but this carbon storage capacity
may change in the future due to global warming. In addition, boreal forest
fires have increased in frequency and amount burned over the past 50 years.
We tested the hypotheses that (1) fire-affected boreal forest soils will have
greater CO2 flux rates in a global warming scenario and (2) increased CO2
flux rates are correlated with an increase in microbial abundance. We tested
this hypothesis by experimentally raising the average daily ambient temperature of a recently burned black spruce forest in central Alaska during
the growing season by 1 oC (4 oC mid-day maximum) using open-top
greenhouse chambers. Three years after the fire, soil CO2 flux was significantly higher in warmed plots than controls for all measurement dates
(greenhouse mean: 103.1 mg C m-2 h-1 +/- 7.1 (S.E.); control mean: 82.8
mg C m-2 h-1 +/- 6.3 (p , .05); repeated measures ANOVA). Microbial
abundance did not increase in the warmed plots as we expected. BIOLOG
ecoplateTM analysis indicates that bacterial diversity and bacterial abundance were not higher in warmed plots. Similarly, using an ergosterol extraction technique, we found that fungal abundance was not affected by the
warmer temperatures. Higher soil respiration rates in the warmed plots may
be explained by higher metabolic activity due to the effect of temperature
on enzymatic reaction rates. Our results suggest that in a warmer climate,
fire-disturbed boreal forests may store less carbon than their present carbon
storage. Furthermore, the interaction between global warming and fire may
result in a positive feedback to atmospheric CO2, and therefore, a positive
feedback to global warming.
BERISH, CORY.* US EPA Region 4, Atlanta. Southeastern Ecological
Framework - a tool for managing ecosystem integrity.
The southeastern United States has large areas of unique ecological character which help sustain a healthy environment. Ecosystem fragmentation
by urban sprawl is becoming the major problem in protecting of environmental processes that protect human health. Threats to ecological function
and conflicts in resource protection need to be identified and prioritized in
order to safeguard the functionality of large ecosystem processes that provide environmental services and protect human health. The Southeastern
Ecological Framework (SEF) identifies the most important ecological areas
in the southeastern US and the best ways to connect them for sustaining a

30

Abstracts

functional and healthy ecosystem. The SEF was developed by EPA Region
4 in cooperation with the University of Florida and several State natural
resource management agencies. The SEF has been reviewed and found to
be useful by federal agencies in the Southeastern Natural Resource Leaders
Group including USFS, USFWS, Federal Highways Administration, Department of Defense, USGS, and the Natural Resource Conservation Service as well as county planners and private watershed conservation groups
for planning greenways and riparian habitat protection. Additionally, the
SEF received favorable reviews from EPA ORD as well as the Science
Advisory Board who suggested that it would be valuable to perform a
similar analysis at the national level. Long term effects of global climate
change are a topic for intense discussion. In order for the ecosystem to
maintain its ability to respond to changes in global climate, large scale
connectivity and preservation of biological hot spots is imperative. The
SEF provides some protection for preserving the ability of the ecosystem
to respond to those changes.
BERKOWITZ, ALAN R.,1,* J. MORGAN GROVE,2 KAREN E.H.
STEELE,3 WAYNE C. ZIPPERER,4 RICHARD V. POUYAT,5 CHARLES
H. NILON,6 GEORGE MIDDENDORF7 and KATALIN SZLAVECZ.8 1 Institute of Ecosystem Studies, berkowitza@ecostudies.org, Millbrook, NY;
2
USDA Forest Service, Burlington, VT; 3 Carver Center for Arts and Technology, Towson, MD; 4 USDA Forest Service, Syracuse, NY; 5 USDA Forest Service, Baltimore, MD; 6 University of Missouri-Columbia, Columbia,
MO; 7 Howard University, Washington, DC; 8 Johns Hopkins University,
Baltimore, MD. Teacher and student investigations of urban ecosystems with the Baltimore Ecosystem Study.
The Baltimore Ecosystem Study (BES) School/Community Research Partnership Program connected BES scientists, teachers, students and community groups in order to engage students in on-going studies of the local
environment. Emphasis was on integrating biological, physical and social
dimensions in spatial and historical contexts, using an inquiry approach.
We: 1) conducted professional development programs in urban ecosystem
education with small groups of middle and high school teachers; 2) provided teachers with on-going support from small grants, scientists, peers
and staff; 3) collaborated directly with teachers and their students in research; 4) linked school-based work with community resources; and 5)
worked with a small number of students as interns. Program teachers had
tremendous interest and enthusiasm, and impressive expertise in inquirybased teaching, but reported that they did not teach outdoors frequently,
nor did they teach very much about urban ecosystems. Concerns about
having enough time to do field research and about fitting urban ecosystems
into their curriculum were rated most significant, while concerns about their
teaching skills were rated less important, in part reflecting the exceptional
nature of the participating teachers. Teachers reported increased understanding of the local environment as a result of their participation, and felt more
able to teach about urban ecosystems. Noteworthy achievements included:
1) a student-generated, comprehensive analysis of land use history of the
citys reservoir system, with participating students showing gains in standardized test scores as well; 2) a multi-grade, multi-subject study of a
whole schools grounds; 3) establishment of a summer soil research program for female high school students; 4) development of an earthworm
sampling protocol; and 5) establishment of permanent soil and stream sampling stations at a school. Key factors leading to and limiting our success,
insights about our initial hypotheses, and future directions will be discussed.
BERKSON, JIM,1,* AUTUMN-LYNN HARRISON1,2 and MICHELLE
DAVIS.1 1 Department of Fisheries and Wildlife Sciences, Virginia Tech,
Blacksburg, VA; 2 Society for Conservation Biology, Arlington, VA. Will
telling the students the truth about environmental management scare
them away?
Although the science of ecology should be objective, the process of environmental management in many cases is not. Fresh graduates, armed almost
solely with the science and theory of ecology are ill prepared to deal with
the reality of environmental management in practice. While many universities have tackled the subjective aspects of this field through the addition
of courses in economics, human dimensions, and law, there remains little
focus on their integration. Too often, our curricula focus on the role science

could ideally play in this process; neglecting the roles science actually does
play when integrated with politics, economics, and sociology. In response
to this omission, we created a capstone course to integrate relevant subjects
and issues into a realistic representation of the process of environmental
management. We have employed case studies, guest speakers, and lively
debates of issues not discussed within the traditional ecology or environmental science curriculum. These issues challenge students preconceived
notions. Inevitably, some students are disheartened and others, inspired
when they discover the reality of environmental decision-making. They are
not, however, turned away from the field. We have four years of data to
suggest that graduates of this course stay within the field of environmental
science, despite the discouraging reality. Moreover, most change their desired role within the process leading them towards more satisfying and
effective environmental careers. Graduates from this capstone course are
armed not only with science, but also with realistic expectations.

forts. We outline a new initiative, the National Riverine Restoration Science

Synthesis (NRRSS), that involves a large interdisciplinary team of scientists and engineers working in partnership with the river conservation organization, American Rivers. Participants are assembling a data set that
spans multiple ecoregions and many types of restoration activities performed by diverse groups with various stakeholders. We are addressing:
what kinds of restoration activities, at what scale, and by what means have
taken place; how goals were set and success measured; the extent to which
scientific criteria were used; the extent to which adaptive management was
an explicit component; and the extent to which scientists formed partnerships with restoration practitioners in order to use restoration projects as
opportunities for scientific experimentation. The goal of the project is to
facilitate the linkage between the practice of ecological restoration and the
science of restoration ecology as well as establish standards for data gathering to scientifically assess restoration methods and success.

NDEZ and TERESA VALVERDE.

BERNAL, ROCIO,* LAURA HERNA
Departamento de Ecolog
a y Recursos Naturales, Facultad de Ciencias,
Universidad Nacional Autonoma de Mexico (UNAM), Mexico, D.F., Mexico. Establishment patterns in a metapopulation of the epiphyte Tillandsia recurvata in a Mexican desert shrubland.

BERRY, JOE.* The Balance Between Respiration and Photosynthesis.

Tillandsia recurvata (Bromeliaceae) is an epiphyte that colonizes the canopies of shrubs and trees in the semidesert areas of Central Mexico. Each
colonized tree may be conceived as a local population within a metapopulation. Little is known regarding the factors that determine patch selection
in plant metapopulations. Here we aimed to describe patch occupancy, as
well as microhabitat preferences in this epiphyte as a preliminary step to
understand its metapopulation dynamics. We surveyed a 1ha plot in the
Tehuacan Valley and recorded the position, size and species identity of
every tree and shrub (N51159), as well as the number of Tillandsia recurvata plants established in its canopy. We also carried out germination
experiments, and observations on seedling survival and growth in different
microhabitats within the canopy of three tree species. The results show that
T. recurvata occupy certain tree species with a higher frequency than expected by chance. These preferred host species were Prosopis laevigata,
Acacia bilimekii and Cercidium praecox, which were colonized with a frequency of 65-73%. Additionally, large trees were colonized more frequently than relatively smaller ones, and colonized trees had colonized nearest
neighbors with a higher frequency than expected by chance. Seed germination was equivalent in the three host species tested (Prosopis laevigata,
Acacia bilimekii and Cercidium praecox). However, seeds germinated better when located in the upper canopy (5.15%) than at medium heights
(2.26%). Seedling survivorship after one year was higher on Acacia bilimekii (80.6%), followed by Cercidium praecox (65.4%) and Prosopis laevigata (45.9%). Seedling growth rate was highest in the outer-upper canopy
of A. bilimekii. This study has allowed us to identify the colonizable patches for Tillandsia recurvata within this metapopulation, which is a key issue
for the understanding of metapopulation dynamics in many plant species.
BERNHARDT, EMILY,1,3 MARGARET PALMER1,3 and DAVE ALLAN.2,3 1 Departments of Biology and Entomology, University of Maryland, College Park, MD; 2 School of Natural Resources, University of
Michigan, Ann Arbor, MI; 3 National Center for Ecological Analysis and
Synthesis, Santa Barbara, CA. Improving the science of river restoration: the National River Restoration Science Synthesis.
Streams and rivers have experienced some of the most dramatic humaninduced alterations of any ecosystem types. Species extinctions and declining water quality and quantity are of such urgent concern that restoration
efforts are now a major focus. We argue that successful restoration of
running-water ecosystems is best accomplished by a shift from a strict
engineering focus to efforts directed by interdisciplinary teams of engineers, ecologists, and geomorphologists. Stream structure and function
from a flow, sediment, or biological perspective are inextricably linked to
channel dynamics and sediment movement and geomorphologists have
unique contributions to make. Ecologists provide critical input relative to
the interplay between on-site ecological processes and the hydrologic and
geomorphic context in which these processes are grounded. Interdisciplinary restoration efforts have enormous potential; however, restoration has
suffered from lack of evaluation and lack of synthetic studies of past ef-

We now observe that terrestrial ecosystems are sequestering some of the

CO2 entering the atmosphere from fossil fuel combustion and land use
conversion. Over thousands of years prior to the industrial revolution, the
CO2 concentration of the atmosphere remained quite stable at about 280
ppm. This implies that the rates of uptake and release of CO2 must have
been in steady-state balance. This presentation will consider the extent to
which the current sink can be attributed to feedback responses that maintained steady-state balance between photosynthesis and respiration in preindustrial times. The analysis will draw on measurements of photosynthesis
and respiration at the ecosystem scale conducted at Biosphere 2, and modeling. Contradictory evidence from other approaches will be discussed.
BERTNESS, MARK D.* and BRIAN R. SILLIMAN.* Department of
Ecology and Evolutionary Biology, Brown University, Providence, Rhode
Island, USA. Anthropogenic Modification of New England Salt Marsh
Landscapes.
In spite of the important role that salt marshes play in the ecology of many
shoreline ecosystems, as many as 50-70% of New England salt marshes
have already been lost to development. We present data that suggests that
the remaining salt marshes in southern New England are currently being
rapidly degraded by shoreline development. Shoreline development, operationally defined as the removal of the woody vegetation bordering marshes, increases nutrient input to marshes and lowers soil salinities as a consequence of increased runoff that is not intercepted and processed by the
woody vegetation bordering marshes. On the seaward border of these
marshes, by releasing plants from nutrient competition, nitrogen eutrophication stimulated by local shoreline development is shifting the competitive
balance among marsh plants, leading to the displacement of natural high
marsh plants by low marsh cordgrass. On the terrestrial border of these
same marshes, by removing the woody vegetation buffer between terrestrial
and salt marsh communities, shoreline development is precipitating the
invasion of the common reed, Phragmites via nitrogen eutrophication and
reduced salinities. As a consequence of these human impacts, traditional
salt marsh plant communities and the plants and animals that are dependent
on these habitats are being displaced by monocultures of weedy species.
BETHERS, SUZANNE* and DAVID R. BOWLING. Dept of Biology,
University of Utah, Salt Lake City, UT. Evapotranspiration of a semiarid Utah grassland during the summer monsoon.
Water is the most limiting resource for plant growth on the Colorado Plateau. The timing of precipitation is bimodal, with the majority of moisture
falling in the winter as rain or snow or during monsoon rains in the summer.
The degree to which different plant species will make use of short-duration
moisture inputs is a function of several factors, including photosynthetic
physiology, rooting depth, phenology, nutrient availability, soil crust activity, and competition with other plants. We used the eddy covariance technique to examine the physiological response of a semi-arid grassland in
southeast Utah to summer precipitation events associated with the North
American monsoon. Our study years were part of a multi-year drought that
began in 2000. Precipitation was 79% of the long-term average in 2001
and 54% in 2002. Rainfall amounts during the monsoon periods (days 180-

Abstracts

31

250) of 2001 and 2002 were 61 and 27 mm, respectively, which amounted
to 38 and 24 % of total annual precipitation (30 and 13 % of mean annual
precipitation). Soil moisture content at 10 cm depth during the monsoon
ranged 3-10 % (2001) and 3-6% (2002). Latent heat fluxes increased following rain events, but peaked below 200 W m-2 and only persisted for a
few days. The mean latent heat fluxes (day and night) during the monsoons
each year were 20.7 (2001) and 11.1 W m-2 (2002). We expected an increase in plant activity by C4 grasses during this period, but found little
evidence for a transpiration contribution to latent heat flux, based on analyses of latent heat flux versus vapor pressure deficit during day and night
periods.
BEVER, JAMES D.,1,* PEGGY A. SCHULTZ,1 LIDIA C. YOSHIDA,2
BALA CHAUDHARY,2 KEITH VOGELSANG1 and BANU SUBRAMANIAM.3 1 Indiana University, Department of Biology, Bloomington, Indiana, USA; 2 University of CaliforniaIrvine, Irvine, California, USA;
3
University of Massachusetts, Amherst, Massachusetts, USA. Degradation of mycorrhizal mutualisms and the success of naturalized plants.
Species characteristics and habitat properties can interact to contribute to
successful plant invasions and, in some locations, dominance by naturalized
plant species. We suggest that the dominance of naturalized plant species
in Southern California is facilitated by the degradation of the mycorrhizal
mutualism. Both through examination of floras and experimental tests, we
find that naturalized plant species are generally less dependent on arbuscular mycorrhizal (AM) fungi than native plant species. As a result, naturalized plant species would be predicted to have higher success in disturbed
habitats. We also find that naturalized plant species are poorer hosts for
AM fungi. Therefore, the initial establishment and dominance of an area
by naturalized plant species can inhibit the reestablishment of effective
mycorrhizal mutualists, thereby inhibiting the reestablishment of native
plant species. We find support for this hypothesis of reinforcement in tests
of soil community feedback. Native plant species generally grow better
with soil communities derived from native dominated sites than from sites
dominated by naturalized plant species. Moreover, when the soil community derived from native dominated sites was trained on either native or
naturalized plant species, native plant species generally grow better with
soil communities previously trained with native plants.
BEVIS, KEVIN P.* and ROBERT W. THACKER. University of Alabama
at Birmingham, Birmingham, AL. Herbivore-mediated competition between cyanobacteria and green algae: Impacts on cyanobacterial toxin
production.
Although the filamentous, mat-forming cyanobacterium Lyngbya wollei is
known to produce saxitoxin, a paralytic shellfish poison, experimental analyses of the potential role of saxitoxin in freshwater ecosystems are limited.
Lake Guntersville, AL contains persistent mats of L. wollei that occupy
many shoreline areas, and in most cases are found with sympatric green
algae and high densities of herbivorous snails (Pleurocera annuliferum).
Competition between L. wollei and a common sympatric green alga, Rhizoclonium hieroglyphicum, was studied in a response surface design in the
presence and absence of P. annuliferum. Both L. wollei and R. hieroglyphicum show strong intraspecific competition at high densities. Although
L. wollei growth was not significantly affected by R. hieroglyphicum density, L. wollei exhibited a density-dependent facilitative effect on R. hieroglyphicum growth in the absence of herbivory. The presence of snails had
no effect on R. hieroglyphicum growth, but increased L. wollei growth.
Saxitoxin concentrations were highest in L. wollei grown at high density.
In the presence of herbivores, saxitoxin concentrations decreased with increasing R. hieroglyphicum density. These results suggest that saxitoxin
plays a limited role in interspecific competition and may be produced primarily in response to intraspecific competition. Since the presence of palatable green algae reduces herbivory on this cyanobacterium, L. wollei may
decrease saxitoxin production and gain an associational defense when both
herbivores and alternative foods are present.
BHASKAR, RADIKA,1,* ALFONSO VALIENTE-BANUET2 and DAVID
D. ACKERLY.1 1 Stanford University, Stanford, CA; 2 Instituto de Ecologa,
Universidad Nacional Auto`noma de Mexico, D.F., Mexico, Mexico. Water
deficits and hydraulic conductance: Effects of water availability vs.
evaporative demand.
The influence of seasonal water deficit on plant water relations has been
studied extensively in mediterranean-type climates, which are characterized

32

Abstracts

by hot, dry summers and cool, wet winters. Comparisons with non-mediterranean climates provide an ideal study system to examine the consequences of water availability in the soil vs. atmospheric demand (high
vapor pressure deficits) on plant water relations. In central Mexico evergreen shrub communities closely related to California chaparral experience
a winter dry season followed by a warm summer with higher insolation
and monsoonal rains. Thus the period of highest evaporative demand coincides with the period of highest water availability, whereas the opposite
is true under mediterranean-type climates. We compared whole plant hydraulic conductance, leaf gas exchange, and water use efficiency of 13-15
dominant evergreen shrubs in two sites, one in the state of Puebla, Mexico
and the other in Santa Barbara, California. The sites had similar average
annual rainfall and potential evapotranspiration. Within both sites, species
that experienced greater water deficits, as determined by minimum seasonal
water potentials, had lower hydraulic conductance and light-saturated photosynthetic rates. Between sites, species in California had higher hydraulic
conductance than species in Mexico, relative to minimum water potentials.
This result may be explained by two climatic factors that distinguish the
sites. The first is the concurrence of high atmospheric demand at the time
of extreme soil drought in California; the second is the occurrence of
freeze-thaw events during the onset of soil drought in Mexico. Both factors
may select for higher hydraulic conductance in California relative to Mexico. The contribution of evolutionary divergence and phenotypic plasticity
to these patterns observed in the field will be tested in future studies under
common environmental conditions.
BIERWAGEN, BRITTA G.* University of California, Santa Barbara,
britta@bren.ucsb.edu, Santa Barbara, CA. Changes in habitat connectivity resulting from different patterns of urban growth.
The conversion of habitat to other land-uses is one of the major drivers of
global change. Many studies document the negative impacts of habitat loss
and fragmentation on species persistence and community composition;
however, the functional effects on ecological processes such as dispersal
are less clear. This research develops a theoretical framework for understanding the impacts of urban growth on habitat connectivity over time. I
evaluated the impacts of land-use change on butterflies with different dispersal abilities across a range of initial habitat configurations. I addressed
the problem by applying a cellular automata-based land-use change model
to simulate different urban growth patterns, varying relative emphasis on
road-mediated, urban edge-mediated, or leap-frog type development. I generated initial landscape configurations using a neutral landscape model and
varied the amount of habitat and its aggregation systematically. Image analysis techniques were used to calculate landscape metrics with a variable
window size corresponding to different butterfly dispersal distances. This
approach effectively scaled the view of each urbanizing landscape according to dispersal capabilities of different butterfly classes. Results suggest
that urban growth spreading from one compact area has the highest habitat
connectivity at first; however, as the urban area spreads, connectivity drops
nearly exponentially for the shortest distance dispersers. Initial configurations with many small urban areas distributed across the landscape begin
with up to 60% lower habitat connectivity compared to landscapes with a
few urban centers. The growth of the many small urban areas results in a
slower rate of decline in habitat connectivity. This work underscores the
importance of understanding land-use changes as dynamic drivers of ecological systems.
BIGELOW, SETH W.1,2,* and CHARLES D. CANHAM.1 1 Institute of Ecosystem Studies, Millbrook, NY; 2 USDA-FS Sierra Nevada Research Center, Davis, CA. Tree-soil relationships in a northern hardwood forest
are reinforced by seedling survival patterns.
Tree communities in northern hardwood forests are organized along gradients of calcium and co-varying soil factors. Because clear relationships
already exist at the sapling stage, we tested how survival at the seedling
stage is affected by base cation availability. Seedlings of six species were
transplanted into plots in a southern New England forest that had either
been fertilized with calcium sulfate or magnesium sulfate or left as unfertilized controls. After two years there were no effects of fertilization on
survival. In a companion experiment soil chemistry was measured in randomly located plots of naturally established seedlings, and maximum like-

lihood estimation was used to model two-year survival as a function of

soil chemistry. Significant increases in predictive ability were gained by
inclusion of soil chemistry. At higher levels of soluble aluminum, sugar
maple survival declined and beech survival improved, which is consistent
with the positions of these species along the soil gradient. Eastern hemlock
had increased survival at higher soil pH, which is not consistent with its
position on the soil gradient. We conclude that survival of sugar maple and
beech seedlings in response to soil chemistry plays a key role in establishing the soil relationships characteristic of these species as adults; nevertheless aluminum, rather than calcium, is the factor that the species respond
to most strongly.
BIGGERS, ANDREW D.* and JUREK KOLASA. McMaster University,
Hamilton, Ontario, Canada. Community disassembly What causes divergence?
Communities assembled under the same environmental conditions often
follow divergent trajectories. Some have postulated that such divergence is
due to minute differences in initial conditions; but the question appears to
be unresolved. However, in natural systems, differences in environmental
conditions strongly influence the community assembly process. In particular, variability in physical conditions has often been suggested as a prominent force in structuring communities. The crucial question emerges then
as to what extent community divergence can be attributed to differences in
habitat variability. One possible approach to address this question is to
tackle it in reverse. Thus, our approach has been to create maximum communities and then follow up on their disassembly as a function of time and
local habitat variability. Specifically, we hypothesize that, while communities diverge over time, this divergence will be greater among habitats
with greater variability. We test this hypothesis with experimental data from
20 tropical coastal rock pools at Discovery Bay, Jamaica. First, the contents
of each rock pool were removed and combined together (in a tank) to form
maximum assemblages (max S). Next the emptied pools were thoroughly
cleaned of propagules and the mixed maximum assemblages returned to
each pool bringing them to their original volumes. We recorded temperature, salinity, pH, conductivity, and chlorophyll as well as the invertebrate
communities for over 2 years at varying intervals. Physical measurements
were used to calculate pool variability. Our results indicate that (a) pool
richness and abundance declined over time, (b) these declines were associated with the degree of community divergence in each pool, and (c) that
in the later stages of community disassembly, the physical variation of the
pools interacted with the time that had elapsed since the beginning of the
experiment to determine the magnitude of divergence.
BIGLER, WENDY. Arizona State University, Tempe, AZ. Historical ethnographies as a source of geomorphic knowledge: The Akimel Oodham
and the Gila River.
A collection of thirty-two interviews transcribed in 1914 provides important insights into flood events and environmental change from the 1830s
to the early 20th century. Akimel Oodham (Pima) elders provided detailed
information about the Gila River and their irrigation practices as part of a
federal water rights investigation. Elders describe the timing and impact of
floods on their villages and the ecological changes to their valley following
upstream water diversions by settlers. Ethnohistorical analysis provides
valuable insights into human/environment interactions. Historical maps
augment the interviews by providing a spatial context for their observations. These observations have important implications for understanding
the Hohokam (a prehistoric irrigation culture presumably ancestral to the
Akimel Oodham) and providing a rich historical basis for understanding
extreme environmental change.
BILLOCK, ARLENE G.1 and JAMES B. GRACE.2 1 Johnson Controls
World Service, National Wetlands Research Center, Lafayette, Louisiana,
USA; 2 USGS, National Wetlands Research Center, Lafayette, Louisiana,
USA. Baccharis halimifolia: Seed germination response after exposure
to heat, smoke, and charred wood extract.
Currently, much effort is being made to reduce the abundance of Baccharis
halimifolia using prescribed fire. While exposure to fire often top kills the
shrub, the effects of fire on B. halimifolia seed germination have not been

determined. The objective of this study was to determine the individual

and combined effects of heat shock, smoke, and charred wood extract on
seed germination by this species in order to evaluate the potential effects
that fire may have. Seeds were tested for germination after exposure to
heat shock and smoke with different duration times, and two different concentrations of charred wood extract. Germination was evaluated for a period of 48 days in a greenhouse. We found that germination of presoaked
seeds (soaked in DI water for 24hours before treatments) was significantly
reduced by exposure to 105oC or 120oC for all durations and by exposure
to smoke for all durations, whereas dry seed (i.e., seed not presoaked) was
only reduced by exposure to 120oC and 20% charred wood extract. In a
combination treatment, dry seeds were exposed to 105oC, 10% charred
wood extract and exposure to smoke for 5, 10, and 15 minutes. In these
combined treatments, germination was reduced to less than 1%, a dramatically greater reduction than caused by the individual components in isolation. Overall, the main findings of this study were (1) none of the treatments caused a stimulation of germination, (2) presoaked seeds were impacted more negatively by heat, smoke, and wood extract than dry seeds,
and (3) the combination of treatments had a much greater effect on germination than the individual components. These findings suggest that fire
can be expected to reduce postburn germination by B. halimifolia and that
preburn soil moisture conditions may influence the degree of this effect.
BILYEU, DANIELLE M.,1,* FRANCIS J. SINGER,1,2 N. THOMPSON
HOBBS1 and DAVID J. COOPER.3 1 Natural Resource Ecology Laboratory, Colorado State University, Fort Collins, CO, USA; 2 Earth Resources,
Colorado State University, Fort Collins, CO; 3 Biological Resources Division, USGS, Fort Collins, CO. Willow persistence in Yellowstone National Park: Interacting effects of hydrology and herbivory.
Willows (Salix spp.) in the ungulate winter range of Yellowstone National
Park (YNP) have undergone a well-documented decline in height, distribution, and recruitment over at least the past 70 years. The decline has
coincided with both an increase in ungulate populations and a decrease in
water tables, due to declines in beaver populations and/or a trend towards
a drier, warmer climate. We constructed an experiment with two treatments,
exclosing (to prevent herbivory by ungulates) and artificial damming (to
raise water tables), in order to examine the effects of herbivory and water
table on the growth of S. geyeriana, S. boothii, and S. bebbiana (n54).
The first year of post-treatment data suggests that damming had a positive
effect on current annual growth for S. geyeriana, although exclosing had
a negative effect for both S. geyeriana and S. bebbiana. However, seed
production was much higher inside of exclosures, suggesting that browsed
plants allocate more resources to growth and less to reproduction relative
to unbrowsed plants. Furthermore, increased water availability may also
stimulate growth in browsed plants.
BINCKLEY, CHRISTOPHER A.* and WILLIAM J. RESETARITS. Old
Dominion University, Norfolk, VA, U.S.A. Habitat selection of aquatic
organisms across gradients of forest canopy coverage.
Many species segregate amongst aquatic habitats that differ in resource
abundance. Two mechanisms, differing primarily in assumed colonization
strategies used by species to select habitats, could explain these field patterns. The first mechanism emphasizes differential mortality within habitats
after a random colonization of all habitat types. A second mechanism emphasizes nonrandom colonization of habitats (e.g. habitat selection) based
on habitat specific fitness consequences. We tested the strength of this second mechanism by conducting two experiments that quantified the abundance and species richness of colonizing aquatic beetles. In the first experiment we established 24 experimental ponds with 12 placed 3m into the
forest (closed canopy) and 12 placed 3m out from the forest edge (open
canopy). Beetles were collected weekly for 6 weeks and mean abundance
and species richness were both significantly higher in open canopy ponds.
In the second experiment we established 24 experimental ponds as in experiment 1, but utilized a split-plot design with canopy as the whole plot
factor and nutrient addition (0, 30, or 60 g) as the sub-plot factor. For both
response variables there was no significant canopy by nutrient interaction
or significant nutrient effect as beetles again strongly avoided closed canopy ponds. Behavioral avoidance of particular habitats resulted in a greater
degree of spatial co-occurrence than would occur if ponds were chosen

Abstracts

33

randomly, suggesting that habitat selection is one primary mechanism

structuring aquatic communities.
BINFORD, MICHAEL W.,1,* GREGORY STARR,1 HENRY GHOLZ,1,2
GRENVILLE BARNES,1 LEVENT GENC,1 SCOT E. SMITH1 and ALLISON FLEMING.1 1 University of Florida, Gainesville, FL, USA; 2 US
National Science Foundation, Arlington, VA. Decadal-scale dynamics of
land ownership and carbon storage in the southeastern lower coastal
plain region of the U.S.
Forests of the Southeastern U.S. Coastal Plain are owned and managed by
a wide variety of landowners and leaseholders, and their activities may
influence ecosystem processes. We study carbon exchange and storage over
25 years at the landscape level by integrating on-the-ground measurements
of biomass accumulation and net ecosystem exchange (NEE), ecosystem
modeling, and satellite remote sensing methods to create several independent estimates of C dynamics within four 15 x 15 km study areas in north
Florida. One of the study areas has been the site of long-term ecosystem
measurements, and provides data for calibrating the models. The landscapelevel, spatially explicit estimates are then combined with a time series of
parcel-level ownership classes (commercial, forest products industry, private, government, mining companies, other) to examine how ownership
influences C dynamics. We show an annual C uptake by ecosystem processes at the landscape level, but tree harvests, some land conversion or
mining, and occasional fires often result in net annual C losses from each
of our study areas. Much of the harvest C loss is removed for manufacturing forest products (e.g. paper or lumber), so contribution to atmospheric
CO2 is unknown. Large-extent fires generated major C losses to the atmosphere several times in the time period. Land ownership class does not
significantly affect C dynamics because owners usually lease land to others
for various purposes including plantations, and the management practices
of individual leaseholders determine the patterns of forest growth and harvest, and consequently C dynamics.
BIRD, CHRISTOPHER E.* and CELIA M. SMITH. University of Hawaii
at Manoa, Honolulu, HI. Effects of grazers on algal composition on a
wave-exposed rocky shore in Hawaii.
The rocky shores of Hawaii are characterized by a distinct horizontal zonation in gross algal functional form along wave exposure gradients. Waveexposed shores are dominated by crustose algae, with scattered patches of
erect algae, while more protected shores are dominated by erect algae. The
densities of the shingle urchin, Colobocentrotus atratus, and the limpet,
Cellana sandwicensis, are also highly correlated with wave exposure, occurring at densities exceeding 60/m2 on exposed shores. We investigated
the contribution of these two grazers to the observed distribution of algal
form along the shoreline. Six blocks of four 0.25m2 fences were installed
on the wave exposed sections of Manana Islet, Oahu. The absence and
presence of C. atratus and C. sandwicensis at high natural densities (60/
m2) were manipulated factorially. The percent cover and composition of
algae were determined prior to the addition of grazers in April 2002 and
at the conclusion of the experiment in November 2002. Five out of six of
the grazer removal fences were successful at excluding grazers. Of these
five fenced areas, four experienced at least a 35% decrease in the cover of
coralline crusts and were dominated by a combination of erect algae and
fleshy crusts, while the fifth was dominated solely by fleshy crusts. Both
C. atratus and C. sandwicensis were able to maintain areas originally free
of erect algae, and both reduced the percent cover of fleshy crusts by 100%.
Only C. atratus was able to clear crustose coralline algae, reducing cover
by up to 60%. Based on these results, we conclude that both C. atratus
and C. sandwicensis contribute significantly to the low cover of erect algae
on exposed rocky shores. We hypothesize that asymmetrical competition
for food is occurring between these two grazers, C. atratus being the stronger competitor, which could be important in the management of the C.
sandwicensis fishery.
BIRD, SUZANNE,1 DAN BINKLEY1 and RANDY SENOCK.2 1 Colorado
State University, Fort Collins, CO; 2 University of Hawaii at Hilo, Hilo,
HI. Tree competition: Species and soil fertility in mixed stands of Eucalyptus saligna and Facaltaria mollucana.
The growth of forests has been examined at the scales of hectares and
landscapes for more than a century, but quantitative information is sparse

34

Abstracts

on the growth interactions of trees sharing a neighborhood (at a scale of

10 to 100 m2). An understanding of the immediate competitive neighborhood of trees (the number, size, distance, and identity of neighbors), and
how these vary with soil fertility, may lead to better predictions of standlevel growth and structural characteristics. An individual tree, neighborhood analysis was used to asses the relative importance of inter- and intraspecific competition and facilitation for tree growth along a fertility gradient. We conducted our research in a 20-year replacement series of Eucalyptus saligna and nitrogen-fixing Facaltaria mollucana in Hawaii.
Eucalyptus and Facaltaria showed a synergistic growth response in mixtures. Increasing mortality and size differentiation in recent years suggests
that competition has intensified between neighboring individuals. We used
diameter measurements from Age 18 and 20 years to calculate annual
above-ground net biomass increment for individual trees, and mapped all
stems. Mortality in the 2 year period studied was highest in Facaltaria,
regardless of the species mix, while Eucalyptus have their greatest mortality
in pure stands. The growth of both Eucalyptus and Facaltaria declined
with increasing biomass of neighborhood trees within a 2 meter radius.
Neighborhood biomass explained more of the variation in growth than the
number of neighbors, and the impact of neighbor biomass on Eucalyptus
growth was smaller when the neighbors were Facaltaria. The nutrient status of the soil varied by a factor of more than 2 among plots, and had a
significant effect on these competitive interactions.
BISHOP, BLAIR B.,* PAUL P. MOU and ANNE E. HERSHEY. University
of North Carolina, Greensboro, tel. 336-256-1074, email: bbbishop@
uncg.edu, Greensboro, NC. Retention and mitigation of anthropogenic
nitrogen and phosphorus in headwater urban riparian systems.
Urban areas in headwater regions have the potential to strongly influence
water quality throughout the entire river basin. However, few studies have
investigated the importance of urban riparian zones in retaining and mitigating inputs of anthropogenic nitrogen and phosphorus. We conducted
seasonal experimental applications using 25 tons of nutrient solution (18
mg/L NO3-, 10 mg/L NH4+, and 9 mg/L PO43-) in surface water to forested
and grassy buffer zones in the City of Greensboro, NC, which forms the
headwaters of the Cape Fear River basin. Preliminary results suggest that
riparian soils in both vegetation types have a strong buffer capacity in
retaining 95% of experimentally applied NH4+ and PO43-, and 80% of NO3. Buffer capacity for NO3-, NH4+, and PO43-, varied based on season (p,
0.0001) winter,spring indicating different dominant biogeochemical processes. Forested riparian buffers had a higher infiltration and retention capacity for inputs of anthropogenic nutrients, reducing mean concentrations
from 15 mg/L NO3-, 7 mg/L NH4+, 11 mg/L PO43- in upslope concentrations to 1.6 mg/L NO3-, 0.08 mg/L NH4+, and 0.15 mg/L PO43- near stream
bank. A strong pattern was found between the retention of NH4+ within the
upper portions of the soil horizon (p , 0.05), while NO3- retention was
strongly related to season and riparian width (p, 0.05), suggesting that
hydrologic conditions and site-specific soil properties control nutrient retention. Longitudinal (down slope) changes in surface and subsurface nutrient concentrations across urban riparian buffers represent the net effects
of input, infiltration, and buffer capacity, and create a viable sink for anthropogenic nitrogen and phosphorus. Changes in common storm water
management practices from piping discharge to spreading discharge along
vegetated riparian zones could greatly improve water quality.
BJORNDAL, KAREN A.,1,* ALAN B. BOLTEN,2 THOMAS DELLINGER,3 CLAUDIA DELGADO3 and HELEN R. MARTINS.4
1
kab@zoology.ufl.edu, University of Florida, Gainesville, FL; 2 University
of Florida, Gainesville, FL; 3 Universidade da Madeira, Funchal, Portugal;
4
Universidade dos Azores, Horta, Portugal. Compensatory growth in
oceanic loggerhead sea turtles, Caretta caretta: response to a stochastic
environment.
Compensatory growth is accelerated growth that may occur when an organism that has grown at a reduced rate as a result of sub-optimal environmental conditions is exposed to better conditions. Compensatory growth
is considered an adaptation to variable environments. Although documented thoroughly under captive conditions, compensatory growth has rarely
been studied in wild populations. In their first years of life, oceanic-stage
loggerhead sea turtles have relatively little control over their geographic

position or movements and thus have an extremely stochastic lifestyle with

great variation in food availability and temperature. This environmental
variation results in variable growth rates. We evaluate somatic growth functions of oceanic-stage loggerheads from the eastern Atlantic based on skeletochronology that allowed us to assign age to each individual. We demonstrate that oceanic-stage loggerheads exhibit compensatory growth based
on three analytical approaches: changes in coefficients of variation in sizeat-age, generalized additive model regression analyses of somatic growth,
and linear regression of age-specific growth rates. As a result of compensatory growth, variation in size-at-age in these juvenile loggerheads is substantially reduced. Thus, size is a better predictor of age than expected
based on variation in growth rates. Compensatory growth decreases with
age, apparently as loggerheads gain greater control over their movements.
BLACK, BRYAN A.* and MARC D. ABRAMS. Penn State University,
University Park, PA. Applicability of boundary-line release citeria: a
survey of species.
Identifying releases from suppression represents one of the most fundamental dendroecological procedures for quantifying forest disturbance histories. Previous methods have scaled releases using such broad predictors
as species, crown class, age, or diameter, and possibly over-generalize the
ability of a tree to respond to a disturbance event. In recent studies, we
show that prior growth is a more fundamental predictor of release response.
Maximum release response sharply declines with increasing levels of prior
growth, and is quantified as a negative exponential boundary line. We have
used this prior-growth boundary line to develop new release criteria that
standardize each release with respect to maximum growth potential. To
date, the relationship between prior growth and maximum release response
has been quantified for Tsuga canadensis, Quercus prinus, and Pinus strobus. In this study, we describe prior-growth boundary lines for eight additional tree species indigenous to wide range of ecotypes in North America. These species include Picea glauca, Picea mariana, Pinus echinata,
Pinus ponderosa, Pseudotsuga menzesii, Quercus alba, Quercus macrocarpa, and Quercus stellata. Configuration of the prior-growth boundary
line is unique to each species, and reflects adaptations to understory conditions. For example, species with high levels of understory tolerance have
relatively high levels of slow growth and are better adapted to respond to
disturbance at these slow growth rates. Overall, the diversity of species that
exhibit a clear prior-growth boundary line underscores the broad applicability of this approach to determining forest disturbance histories.
BLACK, R. ALAN1,* and R. BRANDON PRATT.1,2 1 School of Biological
Sciences, Washington State University, Pullman, Washington, USA; 2 Natural Science Division, Pepperdine University, Malibu, California, USA.
Cavitation repair under tension: Is it consistent with the cohesion-tension theory?
We examined diurnal patterns of cavitation in two genotypes of black cottonwood (Populus trichocarpa) displaying divergent stomatal behavior under water stress: one genotype incompletely closing stomata (ISC genotype)
while the second closes its stomata (SC genotype). To test for cavitation
we made diurnal measurements of xylem specific hydraulic conductivity
(Ks), and ultrasonic acoustic emissions (UAE) under well-watered conditions in a greenhouse. The Ks of stems for the ISC genotype grown for one
year were 47% lower at 13:00 hours compared to predawn. Predawn xylem
water potentials (Cx) were -0.20+0.02 MPa with no evidence of positive
root pressure overnight. After two years of growth Ks displayed the same
diurnal pattern but only when predawn Cxs were . -0.20+0.02 MPa. In
contrast, the SC genotype displayed no diurnal difference in Ks and Cx at
predawn was ,-0.20 MPa. Ultrasonic acoustic emissions measured in situ
were not different between the genotypes. For field grown plants, Kss were
not different between predawn, midday, and dusk measurements and predawn Cxs were , -0.30 MPa for both genotypes. Experimentally raising
Cx of excised stems increased Ks by about 15% for both genotypes. Vulnerability to cavitation of lateral stems was not different between the genotypes (Cx at 50% loss of hydraulic conductivity (C50) was -0.85+0.1
and -0.95+0.1 MPa for the ISC and SC genotypes respectively; p.0.05).
We conclude that the decline in Ks diurnally displayed by the ISC genotype
cannot be explained by differential vulnerability to cavitation, hydrogel
behavior of pit membranes, or positive root pressure. Instead, the ISC ge-

notype refilled embolized vessels when overnight Cx was , -0.20 MPa

while the SC genotype did not refill vessels because its Cx was , -0.20.
MPa. Our data are consistent with the Yang and Tyree (1992) model of
cavitation repair and the cohesion-tension theory of water transport.
BLAIR, LAUREN P.* and MATTHEW D. MORAN. Hendrix College,
Conway, AR. Primary forest in Arkansas post oak savannas.
The amount of primary forest (i.e. old-growth) has declined dramatically
in the United States during the past three centuries. However, remnants of
this habitat type remain, even in areas with substantial human populations.
These primary forests may exist on sites with unfavorable environmental
conditions, exhibit slow growth rates, and therefore not be evident. We
hypothesized that primary forest could be found on post oak savannas that
exist on south-facing slopes in scattered areas throughout central Arkansas.
We analyzed the demography of trees in two field sites to determine the
age structure of the different tree species. Both sites were dominated by a
few species, especially post oak (Quercus stellata), black hickory (Carya
texana), and winged elm (Ulmus alata). According to the demography traits
of these three species, site #1 had a major disturbance even approximately
50 years ago and therefore does not represent primary forest. However,
demography traits from site #2 indicated a much older forest with the
majority post oaks between 175-225 years old, which is pre-European settlement for this portion of Arkansas. Therefore, this site does represent a
primary forest and may shed light on the community structure of undisturbed forests in Arkansas. We suggest that further investigation will reveal
other primary forests in this geographic area.
BLAIR, ROBERT B.1,* and JOSEPH A. REALE.2 1 Dept of Fisheries,
Wildlife, and Conservation Biology, University of Minnesota, St. Paul,
MN, USA; 2 City of Boulder, Open Space and Mountain Parks, Boulder,
CO, USA. Life history attributes of bird communities along an urban
gradient: Does nesting success influence species composition?
The increase in urbanization in North America has raised concerns regarding impacts on avian populations. In this study, we assessed the life history
attributes of birds along an urban gradient in southwestern Ohio. We found
that the number of species that use a multiple-brood breeding strategy increases with urbanization. Further, birds identified as high-nesting species
reached peak levels at the most natural sites and decrease in number with
the urbanization. In contrast, low-nesting species exhibited the reverse
trend. These findings suggested that nesting success may drive the distribution of avian species along an urban gradient. To investigate this, we
measured the nesting success of American robins and Northern cardinals
and analyzed the changes along the gradient. We found that nesting failure
was not significantly correlated with location on the gradient, but was correlated to the height of the nest, which decreased significantly from the
most natural to the most urban sites. Further, the density of adult birds did
not predict nesting failure. These findings suggest that nesting site is a
critical resource in urban environments that regulates bird community composition.
BLAUSTEIN, ANDREW R.,1,* BARBARA A. HAN,1 BELDEN K. LISA,5
JOSEPH M. KIESECKER,6 DOUG CHIVERS,3 ADOLFO MARCO,4
ELIZABETH FASY1 and LEE B. KATS.2 1 Oregon State University, Corvallis, OR, USA; 2 Pepperdine University, Malibu, CA, USA; 3 University
of Saskatchewan, Saskatoon, Saskatchewan, Canada; 4 Donana Biological
Station, Sevilla, Sevilla, Spain; 5 University of Washington, Seattle, WA,
USA; 6 Pennsylvania State University, University Park, PA, USA. Are amphibians exposed to harmful levels of ultraviolet-B radiation?
As part of an overall biodiversity crisis, populations of numerous species
are in decline. This includes populations of amphibians from a wide variety
of habitats and locations around the world. Various factors appear to be
contributing to amphibian population declines. These include habitat destruction, pathogens, introduced non-native species, contaminants, and
global environmental changes. The diversity of locations where amphibian
populations have declined has lead to studies examining the role of atmospheric factors such as increasing ultraviolet-B (UV-B) radiation. Both
laboratory and field experiments have shown that UV-B harms many amphibian species. However, the effects of UV-B radiation on amphibians

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35

differ between species and are often dependent upon life history stage.
Even short-term exposure to low levels of UV-B radiation can kill the
embryos of some species, whereas the embryos of other species are resistant to long-term exposure and higher doses of UV-B radiation. Although
the embryos of some species may not be noticeably affected by UV-B, they
may accrue sublethal effects that appear in later life stages. The effects of
UV-B on amphibians are influenced by ecological parameters, synergistic
effects with other agents, amphibian behavior and amphibian defenses
against UV-B. Recent surveys of hundreds of ponds and lakes in the Pacific
Northwest suggest that many amphibians are subjected to harmful UV-B
doses and corroborate the results of experiments that illustrate these effects
in the field.
BLISS, KRISTIN M.,1,* ROBERT H. JONES,2 RACHEL C. WAGNER2
and PAUL P. MOU.3 1 Randolph-Macon Womans College, Biology Department, Lynchburg, VA, USA; 2 Virginia Tech, Department of Biology,
Blacksburg, VA, USA; 3 Department of Biology, University of North Carolina at Greensboro, Greensboro, NC, USA. Root foraging for patchy
nutrients: Morphological, physiological and demographic plasticity in
four species.
In response to soil nutrient heterogeneity, plant root systems exhibit morphological, physiological, or demographic plasticity. We predicted that species adapted to different positions within a successional sequence would
vary in their foraging ability. We tested this prediction using two pot experiments, each with two herb and two tree species that co-occur in the
southeastern USA but differ in successional status. In the first experiment,
we created heterogeneity by concentrating slow-release fertilizer in one half
of the pot; homogeneity was created in additional pots by distributing the
same amount of fertilizer evenly. Nutrient heterogeneity led to an increase
in root mass in all four species (P , 0.001), however all species responded
similarly with respect to morphological plasticity. Plants were dosed with
a low concentration of 15N labeled ammonium nitrate 48 hours prior to
harvest. Uptake rates per unit of root mass were greater for roots conditioned to low nutrient levels (P , 0.009), but species tended to differ in
the degree of this response (P , 0.080), indicating variable levels of physiological plasticity. In a second experiment, seedlings were grown in rhizotron boxes with high and low fertility halves (19:1 nutrient ratio). Over
21 weeks, biweekly root tracings were used to estimate production, mortality and turnover. In all species, the high fertility half had significantly
greater mortality (P 5 0.033) and turnover (P 5 0.006). Significant differences among species were detected for each demographic variable; however, species did not differ in the degree of demographic plasticity as measured by the relative difference between high and low nutrient patches.
Plants exposed to fertile soil patches show within-root system plasticity in
morphology, physiology, and demography. Contrary to our prediction, degree of plasticity was not strongly variable across the four tested species.
BLOOD, ELIZABETH R.1,* and JAMES HOOK.2 1 Joseph Jones Ecological Research Center, Newton, GA, USA; 2 University of Georgia, Tifton,
GA, USA. Science, culture, and the need for local knowledge in understanding process and implementation.
Ecological knowledge has a central role in resolving the most critical and
complex societal challenge we face this century: freshwater sustainability.
Formulation of adaptive co-management strategies that foster both the vitality and sustainability of intrinsically linked human and ecological systems typifies the complexity, challenges, and opportunities that scientists
and society face in achieving sustainable freshwater resources. Regional
differences in freshwater systems, human systems, their interdependence,
their dynamics, drivers of change, and resource use trajectories challenges
traditional management approaches. Synthesis of ecological, economic, and
social knowledge is essential to defining ecological bounds for human use
and defining strategies that assures ecological integrity is maintained.
Emerging cross-disciplinary theory, science-based policy efforts, and intellectual syntheses are frameworks and foundations for incorporating ecological knowledge into the sociopolitical processes. Equally important is
designing new social and institutional partnerships and processes that effectively translate and integrate scientific knowledge into societal beliefs
and norms and conversely local knowledge and cultural practices into ecological understanding. Through communication, adaptation, and transfor-

36

Abstracts

mation of scientific knowledge and professional judgment, citizens can

grasp the issues, construct a sustainable vision and regional knowledge,
define appropriate co-management prescriptions, implement them, and assess their effectiveness. A grassroots regional leadership effort will be described that has evolved a participatory democratic process that is building
knowledge, infrastructure, resources, support, trust, consensus, negotiated
solutions, and a collective regional vision to sustaining fresh water resources.
BLUM, MICHAEL J.,1,2,* JASON MCLACHLAN2 and COLIN J. SAUNDERS.2 1 Molecular Ecology Research Branch, National Exposure Research Laboratory, US EPA, Cincinnati, OH; 2 Duke University, Dept of
Biology, Durham, NC. Adaptation of coastal marsh species to global
climate change.
Recent work has shown that plants exhibit heritable responses to changes
in atmospheric CO2. Given that atmospheric CO2 has steadily increased
since the industrial revolution, it is possible that plants are under selection
for tolerance to high CO2 concentrations. Like migration, adaptation may
buffer populations and limit the threat of extinction due to climate change.
We have begun studies on two eastern North American coastal marsh plant
species (Scirpus olneyi, Spartina patens) to determine whether these coastal
plants may evolve in response to increased CO2 concentrations. Studies
involving both species are now underway to determine the genetic component of phenotypic responses to changes in atmospheric CO2. Common
garden experiments determining responses to CO2, salinity and temperature
changes, alongside comparison of genotype frequencies across time will
likely indicate whether specific genotypes are favored under conditions of
increased CO2. Complementary work is being done to assess the genetic
variability of extant populations at microsatellite loci, alongside research
that involves the recovery and analysis of DNA from macrofossils deposited in coastal sediments since the industrial revolution. Our preliminary
studies show that DNA can be recovered from 100-200 year old S. olneyi
seeds, and greenhouse studies have shown that these seeds are viable after
such long periods of dormancy. Data on adaptive potential stand to improve
current physiology-based models and may provide more accurate predictions of how coastal species respond to global climate change.
BLUMENTHAL, DANA M.,1,2,* NICHOLAS R. JORDAN1 and ELIZABETH L. SVENSON.1 1 University of Minnesota, St. Paul, Minnesota;
2
USDA-ARS Rangeland Resources Research Unit, Fort Collins, CO. Tallgrass prairie restoration controls weeds.
Weed control in non-cropland often precludes management for biological
diversity. It may be possible, however, to use native late-successional plant
communities to exclude weeds, thereby managing simultaneously for diversity and weed control. We studied a limited form of this generalization,
considering the invasibility of a particular native community, tallgrass prairie, by a class of invader, early-successional weeds. We measured invasion
of extant and added weeds within randomized, 6-year-old plots of restored
prairie and unrestored, old field vegetation. This experimental approach
enabled us to separate the effects of late successional vegetation per se
from factors that commonly covary with late successional vegetation, such
as propagule pressure and time since disturbance. Restoration reduced extant weed biomass by 94%, extant weed stem number by 76%, and stem
numbers of four individual extant weed species, relative to no restoration.
Similarly, restoration reduced added weed biomass by 85% and reduced
biomass of 4 individual added weed species, each by greater than 92%. To
examine potential mechanisms underlying weed responses, we treated subplots with burning (to reduce establishment limitation), and N addition (to
reduce competition). The role of establishment limitation in reducing weed
invasion is suggested by increased litter mass and reduced weed density
with restoration, and by stronger weed responses to burning in restored
than unrestored plots. The role of competition in reducing weed invasion
is suggested by increased biomass of extant vegetation, decreased light
levels and decreased weed size with restoration.
BOBICH, EDWARD G.,* KATIE A. GRIEVE, GREG A. BARRON-GAFFORD and RAMESH MURTHY. Biosphere 2 Center, Columbia University,
Oracle, AZ, USA. Sap flux responses to drought and humidity for cottonwoods under current and increased CO2concentrations.
Sap flux in stems of woody plants depends on available soil water, xylem
conductivity, and leaf water status, which in turn depends on stomatal con-

trol of water loss. Predicted increases in atmospheric CO2concentrations

([CO2]) and changes in rainfall patterns associated with global climate
change can influence sap flux and hence the physiological success of certain
species. In the present study, four-year-old eastern cottonwoods (Populus
deltoides Bartr.) growing in a sealable glasshouse structure under current
(400 ppm) and doubled (800 ppm) ambient [CO2] were investigated with
respect to their sap flux responses (measured using Granier-type probes) to
drought combined with periodic changes in vapor pressure deficit (VPD).
Total daily sap flux decreased an average of 80% after 5 weeks of drought
for plants under both elevated and ambient [CO2]; the decreases are likely
due to increased cavitation of vessels as the drought progressed. Decreases
in total daily sap flux occurred primarily during periods of high VPD (3.1
kPa); sap flux was relatively unchanged during periods of low VPD (1.2
kPa). Decreases in sap flux were responsible for over 90% of the decreases
in hydraulic conductivity (calculated using sap flux, midday leaf water
potentials, and soil water potentials) at midday for plants under both [CO2].
For the first 4 weeks, total daily sap flux was greater for plants in doubled
versus current [CO2], reflecting the greater total leaf area and stomatal
frequency for plants growing under the doubled [CO2]. In addition, wood
of plants under the doubled [CO2] was 13% less dense than wood of plants
under the current [CO2], possibly indicating a greater conducting area per
unit wood area. The results suggest that P. deltoides will take up and
transport more water, allowing for greater photosynthesis, during short periods of drought in future atmospheric [CO2].
BOECKLEN, WILLIAM J.* New Mexico State University, Las Cruces,
NM. Leaf fluctuating asymmetry and its role in plant-herbivore interactions in the Quercus undulata species complex.
Fluctuating asymmetry (FA) is a measure of developmental instability, and
is characterized by a random pattern of deviations from perfect symmetry
in bilaterally or radially symmetric traits. High levels of FA has been associated with environmental stresses, such as competition, pollution, ultraviolet radiation, temperature, and soil moisture and nutrients; and with genetic stresses, such as inbreeding, intraspecific hybridization, and interspecific hybridization. The degree to which FA is heritable and the fitness
consequences of FA have generated much recent controversy. While it is
generally excepted that FA may play a role in sexual selection, its role in
natural selection is more equivocal. Whether FA is an important source of
plant heterogeneity and important in plant-herbivore interactions remains
an open question. I examine leaf fluctuating asymmetry in the Quercus
grisea x Q. gambelii species complex and assess its role in plant-herbivore
interactions. Preliminary analysis indicates that FA of leaves may be heritable and may have fitness consequences. I have measured leaf FA for
seedlings generated by artificial crosses and transplanted to a common garden. There was a significant parent-offspring regression in levels of FA.
For Q. gambelii, there was a significant inverse relationship between FA
and acorn loads at one site (p , 0.04), but not in the other two sites
examined. In addition, leaf FA appears to be a significant source of plant
heterogeneity to herbivores, as there was a significant positive relationship
(p , 0.007) between average leaf asymmetry in Q. gambelii and densities
of the leaf-miner, Pyllonorycter sp. In Quercus grisea, leaf FA is significantly associated with ovipositional preferences of two species of cynipid
gall wasps, although gall formation does not appear by itself to influence
leaf FA.
BOELMAN, NATALIE T.,* MARC STIEGLITZ and KEVIN L. GRIFFIN.
Lamont-Doherty Earth Observatory, Columbia University, Palisades, NY.
Global scale NDVI of arctic tundra landscapes: What does it really tell
us?
In order to accurately interpret global scale NDVI datasets, it is crucial that
we better understand how the high spatial heterogeneity of arctic tundra
vegetation translates into a single, large-scale NDVI pixel value acquired
from a satellite. We have established relationships between photosynthetically active aboveground biomass to NDVI in two of the three dominant
tundra vegetation communities (wet sedge, r250.84 and moist tussock,
r250.6) at Imnavait Creek, Alaska (688 379 N, 1498 329 W). These data
demonstrate that the relationship between NDVI and aboveground biomass
is not constant between vegetation communities and therefore the interpretation of global scale NDVI values may not be straightforward. We also

compared data sampled from various altitudes (1 m, 4 m, 15 m, 50 m,100

m, 1.7 km and 705 km), and therefore different field of view (FOV) sizes.
A handheld spectroradiometer fitted with a 208 field of view (FOV) restrictor was used to acquire NDVI values from the 1 m to 1.7 km sampling
distances using extension poles on the ground or mounted on a helicopter.
We have also included NDVI values for Imnavait Creek derived from the
MODIS and Landsat sensors orbiting the Earth at 705 km. These data show
that NDVI values vary among the FOVs sampled resulting from the dominant scales of "patchiness" inherent to arctic tundra landscapes.
BOERNER, RALPH E.,1,* THOMAS A. WALDROP,2 JENNIFER A.
BRINKMAN1 and MAC A. CALLAHAM.2 1 Ohio State University, Columbus, OH, USA; 2 U.S.D.A. Forest Service Southern Research Station,
Clemson, SC, USA. Structural and functional restoration effects on soil
microbial activity in two contrasting ecosystems.
As part of a national-scale evaluation of the consequences of structural and
functional restoration treatments in ecosystems that historically had frequent fire, we determined the effects of reintroduction of dormant season
fire (functional restoration) and thinning from below (structural restoration)
on soil organic matter characteristics and microbial activity in South Carolina pine-oak and Ohio oak-hickory ecosystems. In South Carolina the
three blocks in which treatments were imposed varied in age and size
distribution whereas in Ohio they varied in parent material and site fertility.
In the pine-oak ecosystem, fire resulted in reduced acid phosphatase and
chitinase activity in the youngest block, but not in the older two. Thinning
resulted in reduced chitinase activity in the oldest block. Phenol oxidase
activity was reduced by both burning and thinning in the oldest block.
Multiple regression indicated that posttreatment enzyme activity in this
pine-dominated ecosystem was most closely linked to pretreatment soil
carbon content and posttreatment soil C:N ratio. In the oak-hickory ecosystem, acid phosphatase activity was reduced by burning in all three
blocks, whereas chitinase activity was unaffected. Phenol oxidase activity
was increased by fire in the most fertile block. Thinning reduced acid
phosphatase activity, but only in the least fertile block. Multiple regression
indicated that posttreatment enzyme activity in the hardwood-dominated
ecosystem was most closely linked to posttreatment soil carbon content,
and to a lesser extent posttreatment soil C:N ratio, soil moisture, and soil
texture. These preliminary results suggest that the consequences of restoration treatments on soil microbial activity may vary within and among
ecosystem types in relation to their effects on the quality and quantity of
soil organic matter.
BOHANAN, ROBERT E.1,2,* and MARIANNE E. KRASNY.3 1 Center for
Biology Education, University of Wisconsin-Madison, Madison, WI, US;
2
Center for Limnology, North Temperate Lakes Long-term Ecological Research, University of Wisconsin-Madison, Madison, WI, US; 3 Graduate
Studies, Department of Natural Resources, Cornell University, Ithaca, NY,
US. Integrating K-12 Education with Scientific Research and Teaching.
Our goal is to integrate K-12 outreach with scientific research and the
training of undergraduate and graduate students. The challenge is to develop and understand models for effectively integrating K-12 outreach and
the hope is that the interactions will benefit all involved: K-12 students,
their teachers, and university faculty and their graduate and undergraduate
students. We describe the development, implementation, and assessment of
several models including: K-12 teacher professional development, involvement of Graduate Teaching Fellows in K-12 Education (GK-12), development of instructional materials for K-12 classrooms, informal science
activities, and initiatives aimed at attracting diverse students to science.
Underlying teaching and learning goals emphasize inquiry, updating content, and improving students understanding of the nature of scientific research. Using case studies focusing on urban ecology, biocomplexity, and
community dynamics we have developed models that are robust in nature
and include promising elements for replication and adaptation to other sites.
Over the past 3 years, more than 50 graduate teaching fellows have participated in K-12 environmental sciences projects at 20 middle and high
schools in New York and Wisconsin. In interviews, graduate teaching fellows reported that they learn more about teaching than they would through
teaching assistantships. Similarly interviews with K-12 science teachers

Abstracts

37

and curriculum coordinators, found that the graduate teaching fellows provided unique resources for connecting content and student research. This
model of graduate teaching fellows and K-12 teachers working together
has helped middle and high school students learn how to ask meaningful
scientific questions and how to develop rigorous student inquiry projects
modeled after ecological research. Our results suggest that programs such
as GK-12 can create synergistic and mutually beneficial effects between
universities (e.g. effectively leveraging faculty involvement in outreach,
improving graduate training) and K-12 schools (e.g. improving student and
teacher understanding of the nature and practice of science).
BOLKER, BENJAMIN M.,1,* DOUGLAS J. LEVEY,1 JOSHUA J.
TEWKSBURY,1 SARAH SARGENT,2 TOSHINORI OKUYAMA1 and
NATHANIEL SEAVY.1 1 University of Florida, Gainesville, FL; 2 Allegheny College, Meadville, PA. Data-driven models of bluebird movement
in and near corridors.
The function and design of habitat corridors in ecological landscapes to
mitigate the effects of habitat fragmentation continues to excite interest.
Previous results from a landscape-scale, replicated corridor experiment in
South Carolina showed that open corridors of second growth in a matrix
of forest facilitate colonization of connected habitat patches by bird-dispersed fruits (yaupon holly, Ilex vomitoria) relative to unconnected patches;
they fail to show significant evidence for a "drift-fence" effect where corridors increase colonization of patches by intercepting animals moving
through matrix habitat. We used follow data from individual Eastern Bluebirds (Sialia sialis) to parameterize a statistical model of bird movement
in order to probe the mechanisms underlying the observed effects of corridors on seed dispersal. Our best-fit model included random movement
within patches and corridors, a tendency for birds in the matrix to follow
the nearest edge, and correlated directions of subsequent moves in the
matrix. We simulated the movement of individuals, starting from the center
of a patch and following for the typical gut passage time (45 minutes), to
obtain average proportions of simulated seeds deposited in each patch type.
The model suggests that our previous supposed mechanism for corridor
functionsthat birds prefer to move within corridorsis wrong. Instead,
birds move within the matrix but use the corridor edge as a movement cue.
This edge-based mechanism also suggests that drift-fence effects should
occur, in contrast to previous results. In fact, our results provide evidence
for a small drift-fence effect, and suggest that the failure to find it in the
experimental results results from a lack of power.
BOLTON, GARY H.1,2,* 1 The University of Arizona, Tucson, AZ; 2 University of Wyoming, Laramie, WY. Age/size structures of subtropical
montane tree species influenced by differing resource harvest practices
in Nepal.
Age structures and size class distributions were developed for three common tree species in a subtropical montane forest in west-central Nepal. The
200 ha forest study area lies between 2300 and 2800 m, near the village
of Chimkhola in Myagdi District. Villagers selectively harvest fuelwood,
shelter poles, and tree-leaf fodder for livestock throughout the forest area.
The tree species selected for study represent different resource types with
differing harvest practices. Symplocos ramocissima is used for poles in
temporary livestock-herding shelters, which are constructed in abundance
as herds are kept in various field and forest areas away from the village.
Machilus duthiei is lopped for livestock fodder, which is required in abundance for herds used to fertilize agricultural fields prior to planting. Lindera
pullchermia is not harvested. Species specific harvest practices are reflected
in differing patterns of age structure and size class distribution. Preliminary
results suggest that selective harvesting of forest products by indigenous
peoples may be causing changes in forest composition and structure in this
area of Nepal.
BONNET, VERONIQUE H.,1,2,* ANNA W. SCHOETTLE1 and WAYNE
D. SHEPPERD.1 1 USDA - Rocky Mountain Research Station, Fort Collins,
CO; 2 MATCOM, Fort Collins, CO. Post-fire establishment and regeneration of Pinus ponderosa in relation to environmental conditions and
competitive interactions.
Ponderosa pine (Pinus ponderosa) benefits from fire partly due to changes
in chemical properties of soil and to the reduction of canopy and understory

38

Abstracts

coverage. In 2000, the Jasper Fire burned 34,000 ha of ponderosa pine

forests in the Black Hills of South Dakota, creating a mosaic of environmental conditions. We characterized the spatial establishment of ponderosa
pine seedlings two years after the fire in relation to environmental conditions and influences of competitive interactions. We quantified ponderosa
pine establishment and survival, the corresponding environmental conditions, and the floristic patterns along 22 transects set up from unburned to
the center of burned areas. To supplement this study, we tested the effects
of inter- and intra-specific competition on ponderosa pine seedlings in a
greenhouse. The establishment of ponderosa pine seedlings after fire was
correlated to understory plant composition, canopy, and soil conditions.
The number of established seedlings also decreased with distance from the
unburned areas. The survival rate was correlated to understory plant composition and varied depending on the number of seedlings established.
Growth rates depended on soil conditions, competitor plant species, and
intra-specific interactions. These results demonstrated that ponderosa pine
seedling regeneration after fire is not only dependant upon dispersal distance and environmental conditions, but is also linked to inter and intracompetitive pressure.
BOOTH, ROBERT K.* and STEPHEN T. JACKSON. University of Wyoming, Laramie, WY. Influence of late Holocene moisture variability on
forest dynamics in the western Great Lakes region.
High-resolution records of past moisture variability have recently been obtained from peatlands in the western Great Lakes region using testate amoebae as proxies for past surface-moisture conditions. We used these records
together with pollen and plant macrofossil data to determine how late Holocene population dynamics of mesic tree species (Tsuga canadensis, Fagus grandifolia, Betula alleghaniensis) were influenced by climate variability in the region. In Upper Michigan, populations of mesic tree species
underwent expansions during times of increasing moisture, particularly
when these moisture increases were preceded by drought. For example,
regional populations of B. alleghaniensis and local populations of T. canadensis expanded 4000 years BP during a rapid increase in moisture that
was directly preceded by a high-magnitude, yet relatively short-duration
(100-200 years) drought. High-resolution pollen and charcoal analysis
spanning the drought interval indicates regional wildland fires and abundant
Pteridium ferns, suggesting that the expansion of B. alleghaniensis and T.
canadensis was caused by the combined effect of drought-induced disturbance and the subsequent rapid increase in moisture. Our results suggest a
new explanation for the decline of F. grandifolia populations in southeastern Michigan and southern Ontario during the last 1000 years, which
has been variously attributed to climatic cooling or anthropogenic disturbance. Our peatland proxy-climate record indicates that drought and an
associated increase in fire frequency and/or intensity led to replacement of
F. grandifolia by Pinus populations. Comparisons between records of paleoclimate and vegetation history show that major late Holocene vegetation
changes in the western Great Lakes region were strongly influenced by
moisture extremes, and demonstrate the power of independent paleoclimate
records in evaluating records of vegetational change.
BORER, CATHERINE H.,1,* PAUL G. SCHABERG,2 DONALD H. DEHAYES1 and GARY J. HAWLEY.1 1 The University of Vermont, Burlington, VT; 2 USDA Forest Service, South Burlington, VT. Differential biochemical partitioning and physiological availability of calcium and aluminum in red spruce foliage.
Calcium (Ca) is an essential macronutrient in plants, and is an important
component of many cellular structures and physiological processes. Aluminum (Al) can obstruct Ca uptake, and may compete for biochemical Ca
binding sites, thus inhibiting many Ca-dependent processes. Ca to Al ratios
in the soil have been described as important indicators of both site-specific
forest health and plant damage that can result from soil acidification. By
means of sequential extractions of foliage from mature red spruce (Picea
rubens Sarg.) trees growing under ambient environmental conditions, we
found that Ca preferentially accrues to labile and physiologically active
forms at low total foliar Ca concentrations. At higher levels of total Ca,
substantial quantities of low-solubility Ca also accumulate in the foliage.
This inducible chemical sequestration of Ca occurs only within the documented foliar Ca sufficiency range. A similar pattern of differential chem-

ical sequestration was found for Al. Our results suggest that foliar Al sequestration may provide for passive Al tolerance as a "side-effect" of Ca
partitioning. Accretion of sufficient physiologically available foliar Ca, in
conjunction with Al sequestration, was associated with greater tree growth,
elevated accretion of other nutrients, enhanced cold hardiness, and reduced
winter injury. In contrast, total foliar Ca and Al content, as well as foliar
Ca to Al ratios were not closely related to cold hardiness or winter injury.
These data demonstrate that not all chemical forms of foliar Ca and Al are
equally physiologically relevant, and underscore the importance of assessing relevant components in biogeochemical research studies.
BORER, ELIZABETH,1,* ERIC SEABLOOM,2 JONATHAN SHURIN,2
KURT ANDERSON,1 CAROL BLANCHETTE,1 BERNARDO BROITMAN,1 SCOTT COOPER1 and BENJAMIN HALPERN.1 1 Department of
Ecology, Evolution, and Marine Biology, University of California, Santa
Barbara, CA; 2 National Center for Ecological Analysis and Synthesis, University of California, Santa Barbara, CA. Within- and among-system differences in the strength of trophic cascades: Methodology or biology?
A recent meta-analysis examining data across ecosystem types shows that
trophic cascades, indirect positive effects of predators on plant biomass,
are stronger in benthic marine and stream systems than they are in terrestrial or pelagic systems. This finding begs further investigation into the
causes of this pattern. Using the same studies and meta-analysis techniques,
we examined over 20 biological and methodological factors that have been
proposed to explain variation in the strength of trophic cascades among
studies within an ecosystem and among ecosystem types. After accounting
for inherent biological differences among systems, we found that cascades
are stronger when plant generation times are longer, and that predation on
invertebrate herbivores is associated with stronger cascades than is predation on vertebrate herbivores. A few extremely efficient herbivores (e.g.
sea urchins) are consistently associated with very strong trophic cascades.
In addition, cascading effects tended to decline as the spatial and temporal
scales of the studies increased. One strictly methodological factor was important: predators that are mobile with respect to an experimental unit produced weaker indirect effects on plants than did enclosed predators. Among
ecosystem types, percent nitrogen in plants had the strongest effect size,
suggesting that cascades are stronger in systems with a smaller stoichiometric imbalance between plants and herbivores.
BOROWICZ, VICTORIA A.* Illinois State University, Normal, IL, USA.
Mycorrhizal fungi and plant resistance to herbivory: Does mode of
herbivore feeding matter?
In addition to facilitating phosphorus uptake, mycorrhizal fungi have been
hypothesized to alter relations between plants and their enemies, including
herbivores. Early experiments with caterpillars suggested that mycorrhizae
reduce herbivore performance but subsequent experiments with aphids
found the opposite trend. Based on these studies investigators speculated
that mycorrhizae increase resistance to chewing herbivores but reduce resistance to insects that feed by sucking. Using data from published studies
I conducted a meta-analysis to test the hypothesis that effects of mycorrhizal fungi depend on herbivore mode of feeding. A literature search produced only nine papers that presented data in a form amenable to metaanalysis, yielding a total of 27 separate experiments. Two experiments involved ectomycorrhizal fungi and oaks, and the remaining involved arbuscular mycorrhizal fungi and herbaceous plants. Herbivores included aphids
("suckers" - 8 experiments), and root or leaf chewers (13 experiments), leaf
scrapers (4 experiments), and stem gallers (2 experiments). Due to small
sample sizes of two categories and low within group variance when pooled,
all non-sucker studies were combined in one group and compared to suckers. The effect of mycorrhizal fungi on insect mass or growth rate was
large and positive for suckers; aphids grew larger or faster when they fed
on mycorrhizal hosts. Mycorrhizal fungi did not significantly affect nonsucker growth. Nine experiments with non-suckers also included data on
survival. Mycorrhizal fungi had a moderately large and significant negative
effect on herbivore survival; insects exhibited lower survival on well-colonized plants. Studies thus far provide equivocal support for the hypothesis
that mycorrhizal fungi alter plant resistance and that the direction of the
effect is associated with mode of feeding. The trends were not robust and
are likely to be challenged as more data become available.

BORRETT, STUART R.* Institute of Ecology, University of Georgia, Athens, GA. Development of environ indirect effects in ecological flow
networks: Initial lessons from the Neuse River Estuary.
Indirect interactions are a critical component of ecological environments
and are one source of biocomplexity. Results from environ analysis suggest
that indirect flow intensities tend to dominate direct flow intensities in
conservative transaction networks. One criticism of network environ analysis, however, is that it sums interactions over pathways of infinite length,
thus implicitly assuming infinite time. In this paper, I investigate the development of environ indirect effects to determine the validity of this assumption and the robustness of the conclusion that indirect flow intensities
tend to dominate direct in ecological networks. The analysis presented is
restricted to the output oriented throughflow case of environ analysis. I
used two threshold values to measure development of indirect flow intensities in the extended pathway network. The first, kI.D, is the pathway
length at which indirect flow intensities surpass direct. The second, k95, is
the pathway length at which 95% of the integral (direct + indirect) throughflow is achieved. These thresholds were measured in a seven compartment
nitrogen cycling model developed for the Neuse River Estuary, and parameterized for sixteen seasons from 1985-1989 (Christian, R.R., Thomas, C.R.
2000. Neuse River Estuary Modeling and Monitoring Project Stage 1: Network Analysis for Evaluating the Consequences of Nitrogen Loading. Water Resources Research Institute of the University of North Carolina, Report
325-F). In all sixteen networks kI.D53, and the mean k95 was 224 (6149
SD). Variation in k95 correlates with both total system throughflow (R2 5
0.72) and the degree of cycling (Finn cycling index, R2 5 0.92). These
results indicate that very short durations of system stasis may be sufficient
for indirect flow intensities to dominate direct, but that longer durations
are required for full development.
BORSUK, MARK E.,1,* PETER REICHERT1 and PATRICIA HOLM.2
1
Systems Analysis, Integrated Assessment and Modelling (SIAM), Swiss
Federal Institute of Environmental Science and Technology (EAWAG), Dubendorf, CH, Switzerland; 2 Fischnetz Project Leader, Swiss Federal Institute of Environmental Science and Technology (EAWAG), Dubendorf, CH,
Switzerland. A probability network of brown trout population changes
in Swiss midland rivers.
Catches of brown trout have decreased about 50% in many rivers and
streams in Switzerland in the past 15 years. Additionally, the health status
of numerous brown trout populations has been assessed to be impaired. To
evaluate the causes for these phenomena, a probability network model is
being developed. A probability network is the combination of a graphical
depiction of the relationships among the most important variables in the
system with a probabilistic quantification of these relationships. The graphical model explicitly represents cause-and-effect assumptions between variables that may be obscured under other modeling approaches. In this case,
the variables selected are those describing anthropogenic impacts on habitat, water quality, hydrology, temperature, stocking, and catch, as well as
those describing population characteristics, including growth, condition,
survival, reproduction, and density at various life stages. The probabilistic
relationships are being established using a combination of a dynamic fish
population model, experimental investigations, field data, published reports,
and elicited expert opinion. The quantified network will be used to assess
the historical causal importance of anthropogenic changes, as well as predict the effect of proposed management actions. Analyses are being carried
out for individual streams using site-specific information as evidence to
update less specific prior beliefs.
BOSSART, JANICE.* The College of New Jersey, Ewing, NJ. Species
assembly rules and nestedness of butterfly communities in long-protected relict forests of Ghana.
Indigenous sacred forest groves represent some of the earliest forms of
conservation in the world because of centuries-old restrictions that prohibit
human access. Sacred groves were originally once part of continuous forest
cover, but now exist as isolated relicts of climax forest embedded in a
converted, agropastoral landscape. As such, they represent a remarkable
opportunity in which to ultimately test theoretical predictions about community assembly theory and to explore, in situ, impacts of long-term pro-

Abstracts

39

tection and subsequent complete isolation on species persistence and extinction in human-dominated landscapes. Here I summarize results of a
year-long survey of fruit-feeding butterfly communities in relict sacred forests and forest reserves in the moist semi deciduous forest zone of Ghana.
Nearly 7000 individuals were collected during the survey, representing
more than 80 species. Species composition and evenness were highly variable among sites. Common species at one site were rare or nonexistent at
other sites. Nearly one-third of species collected were trapped at only a
single site. A null-model analysis is used to evaluate the hypothesis that
species composition in these long protected, relict forest islands is a random
subset of the regional species pool of forest dwelling, fruit-feeding butterflies.
BOSWELL, GANTT. Tulane University, New Orleans, LA. Changes in
leaf morphology and biomass allocation reflect soil nitrogen availability
in a carnivorous plant.
Sarracenia rosea, a recently described Gulf Coastal species of carnivorous
pitcher plant, occurs mainly in the western Florida panhandle and adjacent
areas of Georgia, Alabama, and eastern Mississippi. The leaves (pitchers)
serve as both photosynthetic organs and prey-capture devices, and plants
procure nutrients from both soil and prey. We hypothesized that increased
soil nitrogen-availability would be reflected by 1) changes in leaf morphology to maximize surface area (i.e. decreased structural allocation to
carnivory) and 2) decreased biomass allocation to roots. We estimated potential N-availability in two S. rosea habitats characterized by different
soils using buried-bag incubations to assess net N-mineralization rates. We
then measured leaf wing width (a larger wing results in greater leaf surface
area) and percent biomass allocated to roots in plants from the two sites.
The mean annual rates of N-mineralization were significantly different at
the two locations. Leaves of plants from the site with the higher rate of Nmineralization had significantly larger wings and allocated significantly less
biomass to roots, compared to plants from the lower N-mineralization site.
These results suggest that S. rosea responds to differences in available soil
nitrogen by altering leaf morphology and patterns of biomass allocation to
roots. These responses by the plant may be indicative of a decreased dependence on prey-derived nitrogen when growing in soils with higher levels of available nitrogen.
BOUDELL, JERE A.* and JULIET C. STROMBERG. Arizona State University, Tempe, Arizona. Potential for herbaceous community restoration of an impounded southwestern riparian ecosystem of North America via seed bank manipulation.
With the construction of Waddell dam in 1927, the Agua Fria River was
impounded, which impeded flow to downstream reaches. Other than occasional flood releases and dam leakage, the river bed is dry. Our seed
bank investigations from a nearby free-flowing, river revealed the importance of seed banks to riparian vegetation dynamics and to the maintenance
of riparian biodiversity. We initiated seed bank studies of the degraded
riparian ecosystem of the Agua Fria in 1998 to assess site potential for
natural revegetation. 180 soil samples were collected from four dominant
patch types. Seeds within soil samples were allowed to germinate in a
growth chamber, programmed to mimic site conditions, over a two-year
period. Species found in seed banks were placed into categories based on
life-history traits, growth form, moisture tolerance and origin. Wetland ruderals dominated both Tamarix and Tamarix-Salix forest seed banks. Native wetland perennial species, such as Mimulus guttatus, were found in
deeper soil layers of both of these forest seed banks. A few species (e.g.,
Polypogon monspeliensis) heavily dominated the seed banks of these channel-side forests, mirroring the loss of above-ground diversity; this pattern
may reflect the loss of the disturbance processes that typically allow for
high biodiversity in riparian ecosystems. The seed banks in the shrubland
patch types had many upland individuals and thus are beginning to mirror
the community shift towards xeric vegetation. However, some hydromesic
riparian graminoids and forbs (e.g., Cyperus oderatus) were present in
deeper soils. The presence of seed bank species with a range of life history
strategies and moisture tolerances suggests to us that, if the flow regime
was restored, a herbaceous riparian community would establish. The loss
of riparian biodiversity in the seed bank, and shift towards upland species,
suggest that reclamation may be a more probable outcome than full restoration.

40

Abstracts

BOULTON, APRIL M.1,* and KEITH D. AMBERMAN.2 1 Biology Dept.,

Villanova University, Villanova, PA, USA; 2 Veterinary Hospital, University of Pennsylvania, Philadelphia, PA, USA. How Ant Nests Increase
Soil Biota Species Richness: A Mechanistic Field Experiment.
A few studies have shown that ants increase soil biota diversity and soil
nutrient concentrations, but none have examined the mechanism behind
this "ant effect." In this study, we employed a 2 X 2 complete factorial
design (water X food) in the field to mimic the effects of harvester ant
nests (Messor andrei). We hypothesized that, in the absence of ants, moisture and food additions would interact to produce conditions found in ant
nests. Our results indicated that the addition of food to the soil (regardless
of water addition) best mimicked the conditions found inside M. andrei
nests. Our food-treated and ant-nest soils resembled each other with higher
numbers of bacteria, nematodes, miscellaneous eukaryotes, and microarthropods compared to the other soil treatments with relatively depauperate
soil biota. Microbial species richness was also highest in ant and foodtreated samples. Moreover, the effect that ants can have on the soil occurs
quickly-in a period of just two months based on our results. Because ants
are widespread and are the most abundant eusocial insect with many longlived species, they could substantially influence soil and belowground foodwebs in a number of ecosystems and may have important restoration/conservation implications for soil communities.
BOURG, NORMAN A.,1,2,* WILLIAM J. MCSHEA,1 DOUGLAS E.
GILL2 and KEVIN KOY.1,3 1 Smithsonian Institution Conservation and Research Center, Front Royal, VA; 2 University of Maryland - College Park,
College Park, MD; 3 American Museum of Natural History, New York,
NY. CART and GIS-based predictive habitat modeling for Turkeybeard (Xerophyllum asphodeloides), a rare fire-adapted Appalachian
forest herb.
We used a classification and regression tree (CART) model in combination
with GIS to predict suitable habitat and potential new population occurrences for Turkeybeard (Xerophyllum asphodeloides), a rare lily associated
with southern Appalachian pine-oak forests. Turkeybeard is endangered or
rare in portions of its range, listed as a sensitive species on National Forest
lands, and is included in the Center for Plant Conservations National Collection of Endangered Plants. We gathered evidence from field experiments
and long-term population monitoring that supports the hypothesis that turkeybeard is a fire-adapted species, dependent on disturbance by fire for
population maintenance. Based upon these findings, we compiled fire occurrence records for three districts of the George Washington National Forest and Shenandoah National Park and then performed kriging interpolation
on a subset of these data to construct a fire likelihood GIS layer. GIS data
layers describing forest and soil types, elevation, slope, aspect, and planar
and profile landform indices were also compiled for the study area. Randomly located 90 meter diameter sample points were placed within 25
mapped turkeybeard populations as well as across the entire study area,
and data values from each GIS layer were then extracted for each point
and input into CART modeling software. The resulting cross-validated classification tree contained five predicted presence classes, with elevation,
forest type, fire likelihood and slope being the main explanatory variables.
This tree classified 74% of the known turkeybeard population areas, and
90% of known absence areas, correctly. Additionally, eight new occupied
patches were located during ground-truthing of the predicted habitat map
produced from the model. This modeling effort highlights the amenability
of combining CART analysis methods with GIS to define suitable habitat
for rare species over large landscape-scale areas. Our findings also have
important implications for the study of metapopulations and disturbance
regimes in Appalachian forests.
BOVA, ANTHONY S.* and MATTHEW B. DICKINSON. USDA Forest
Service, Northeastern Research Station, 359 Main Road, Delaware, OH,
USA. Making sense of fire temperatures: A thermocouple heat budget
correlates temperatures and flame heat flux.
The ecological and forestry literature abounds with papers in which forest
fires are characterized by "fire temperatures" measured with thermocouples
or temperature sensitive paint applied to metal tags or other objects. The
temperatures indicated by such measurement devices are a function of heat

transfer resulting from the devices thermophysical properties, along with

flame temperature, velocity and emissivity. Unfortunately, these temperatures, by themselves, do not provide information about fundamental attributes of flames (e.g., heat fluxes) that are responsible for the heat transfer
that causes ecological effects (e.g., conduction into tree stems, convection
into tree crowns). There seem to be two ways around this problem: (1)
designing the measurement device so that it mimics the heat transfer properties of the organism (or one of its component parts) or (2) translating
temperatures into fundamental flame attributes by describing the measurement devices heat budget and thermophysical properties. Here, we use a
heat budget to translate in-fire cylindrical thermocouple probe temperatures
and their rates of change into heat fluxes at the surface of tree stems. Net
heat gain (q"net) by the thermocouple probe is the result of heat gains by
convection (q"convection) and net gains by radiation (q"net radiation):
q"net 5 (Thermal Constants) * [q"convection + q"net radiation] Net heat
gain by radiation is the balance of heat gain from the flame and heat losses
by probe re-radiation. Thermal constants describe the probes volumetric
heat capacity. Flame heat fluxes calculated from the thermocouple probe
heat budget were closely correlated with flame heat fluxes measured at
stem surfaces in experimental burns (R2 0.80). In turn, flame heat fluxes
were closely related to the depth of tissue kill (R2 5 0.78).
BOWERS, RACHEL N.* and PETER H. WYCKOFF.* University of Minnesota, Morris, MN. Climate sensitivity of Quercus macrocarpa along
the prairie-forest border in west-central Minnesota.
Quercus macrocarpa (bur oak) is the dominant tree in forest patches of
western Minnesota. Sensitivity of this species to climate may dictate any
movement of the prairie-forest border as western Minnesota warms. We
used tree rings to examine growth patterns for Quercus macrocarpa growing at three locations on a 150 mile transect running along the prairie-forest
border in west-central Minnesota. Results indicate a strong sensitivity to
growing season drought (r . 0.5). Mean growth declines of 60% were seen
in the droughts of the 1930s, and a 40% growth decline accompanied a
drought in 1988. Estimates of the relationship between growth and mortality suggest that growth declines of this magnitude subject trees to greatly
increased mortality risk. IPCC predictions show substantial warming of the
region in coming years, possibly accompanied by declined growing season
rainfall. Our results suggest that these changes may precipitate a decline in
the dominance of Quercus macrocarpa in west-central Minnesota.
BOWERS, RICHARD W.,1,* MATTHEW J. BARRY2 and FERENC A.
DE SZALAY.3 1 Edwards-Pitman Environmental, Inc., Smyrna, Georgia;
2
Arizona State University, Tempe, Arizona; 3 Kent State University, Kent,
Ohio. Effects of herbivory, sediment disturbance, and hydrology on
plant recruitment in a Lake Erie coastal wetland.
In 2000, we examined plant recruitment on exposed mudflats in a Lake
Erie coastal marsh during two years of below average water levels. Mudflat
vegetation was sampled in shallow, medium and deep water transects and
compared to germinable seeds found in sediments from 0-15 cm below the
surface. Using 1m2 quadrats, we examined the effects of sediment disturbance and herbivory on plant recruitment in 2000. Vegetation was then
sampled in 2001 to assess how well the disturbance/herbivory experiment
predicted plant communities that became established on the mudflats. Eighteen species were found on mudflats and 19 species were reared from the
seedbanks in 2000. The shallow transect had the highest number of plants,
but the deep transects had the highest seed abundance and richnesss. Herbivory decreased both plant cover and richness, and sediment disturbance
decreased taxa richness; however responses varied among taxa. Our results
indicate that abundant seedbanks exist in this wetland, but plant recruitment
will be restricted to shallow areas due to turbidity. Communities in 2001
were more diverse (40 species) than predicted from vegetation in 2000.
This was probably because propagules were introduced from nearby habitats. Communities in 2001 were most similar to ungrazed and undisturbed
quadrats in 2000.
BOWLES, CHRISTY M.* and SANDY J. ANDELMAN.* National Center
for Ecological Analysis and Synthesis, 735 State Street Suite 300, Santa
Barbara, CA, 93101. Invasion and composition in North American
grasslands: Is the presence of non-native invasive species related to
plant species diversity and productivity?
Invasive non-native species have been of concern to many grasslands conservation efforts. Non-native invasive species potentially may affect native

species richness and composition, but these relationships are not well-understood. I used data from six North American grasslands (2-10 years) to
examine the relationships between distribution and occurrence of non-native invasive species, composition, native and total richness, disturbance
(grazing and burning) and aboveground annual net primary productivity
(ANPP). I examined these relationships at multiple scales, ranging from 1
m2 to 1000 m2, using a series of t-tests to examine the relationships between
non-native species presence and richness and principle component ordinations to examine the relative influences of each component. The principle
component analyses revealed a negative association between non-native
invasive presence and components of native richness. The identity of the
non-native invasive species is also important. At each site, a small number
of non-native invaders show the strongest negative relationships with native
species composition and native richness. Disturbance and ANPP are weakly
correlated with non-native invasive presence and native richness. This
study has implications for understanding the role of species composition
in grassland communities and for the restoration of grasslands impacted by
invasive non-natives.
BOWLING, ANNA M.,* MICHAEL J. VANNI, WILLIAM H. RENWICK
and ALAN D. CHRISTIAN. Miami University, Oxford, OH, United States.
The effects of land use and precipitation on nutrient and sediment
export from three Ohio watersheds.
Land use, precipitation, soil and landscape characteristics can drive variation in the quantity and quality of nutrients and sediments that are exported
from a given watershed. Although numerous studies have shown that nutrient export rates are higher in agricultural watersheds than in forested
watersheds, export rates are highly variable due to variation in precipitation,
physical factors, and sampling regimes. This variation can obscure trends
in land use effects, and highlights a need for comparative studies with
consistent and intensive sampling. Our research focused on how land use
and precipitation affect the amounts of nutrients and sediments exported in
streams from three Ohio watersheds. Our study sites included Acton Lake
watershed (88% agriculture, 9% forest), Pleasant Hill Lake watershed (51%
agriculture, 45% forest), and Burr Oak Lake watershed (14% agriculture,
81% forest). We used a storm-intensive sampling program to quantify particulate, as well as inorganic and organic dissolved forms of nitrogen (N)
and phosphorus (P), and suspended solids. We found a strong and consistent effect of land use on nutrient and sediment export. Thus, the concentration of NO3-N was highest in Acton Lake watershed (0.99-11.98 mgN/
L), followed by Pleasant Hill (0.81-2.29 mgN/L), and then Burr Oak (0.010.53 mgN/L). Most of the dissolved N exported was in the from of NO3N for the agricultural watershed (Acton Lake), whereas most of the N
exported in the forested watershed (Burr Oak) was in the form of dissolved
organic N. Concentrations of soluble reactive P were also highest in Acton
Lake watershed (4.06-132.37 ugP/L), followed by Pleasant Hill (3.82-40.08
ugP/L) and then Burr Oak (2.55-9.57 ugP/L). Most nutrient loading (dissolved and particulates) occurred during storm events. Particulates ( particulate P and suspended solids) exhibited the greatest variation in response
to precipitation.
BOWLING, DAVID R.1,* and DAN YAKIR.2 1 Dept of Biology, University
of Utah, Salt Lake City, Utah; 2 Dept of Environmental Sciences and Energy Research, Weizmann Institute of Science, Rehovot, Israel. Interpreting ecosystem respiration fluxes using stable isotopes.
Respiration by terrestrial ecosystems is one of the most important fluxes
in the global carbon cycle. Stable isotopes have been an integral component
of carbon cycle studies for several decades, and both, 13C and 18O in CO2
have been used to assess the magnitude of respiration fluxes at the global
scale. During the last decade, application of stable isotopes to ecosystem
carbon cycle studies gained popularity and progress was made in several
areas of biosphere-atmosphere exchange research. In this talk, we will review the application of stable isotope studies to analyses of respiration,
focusing primarily on measurements conducted at the ecosystem scale. We
will discuss linkages that have been observed between mean annual precipitation, atmospheric vapor pressure deficit, and soil moisture availability
with the carbon isotope ratio of total ecosystem respiration (d13CR). We will
also describe results from continental-scale networks of stable isotopes in
ecosystems (BASIN, Carboeuroflux-Stable Isotopes Network) showing var-

Abstracts

41

iation in d13CR that follows general trends with seasonal, meteorological

and geographical parameters. We will explore the potential for using dynamics of d13CR to improve estimates of ecosystem discrimination D13CE
for regional and global-scale models of carbon source/sinks in the land
biosphere.
BOWMAN CUTWAY, HEATHER. Rutgers University, New Brunswick,
NJ. Invasibility and dispersal in urban wetlands.
In urban fragments, the incidence of invasive exotic species is considered
common. Urbanized areas are exposed to many factors that are known to
increase invasibility including high perimeter to area ratio, anthropogenic
influence, and disturbance. This study evaluates the effect of land use on
the invasibility of forested wetlands in the highly urbanized Arthur Kill
watershed in northeastern New Jersey. The diversity of land use within the
urban landscape is usually ignored, causing diverse urban fragments to be
lumped together improperly. Through a vegetation survey of 17 forested
wetlands of different sizes and surrounding land, I determined that wetlands
surrounded by residential land use had a significantly higher richness and
density of exotic species than wetlands surrounded by industrial land use.
Large residential sites had a surprisingly high number of invaded plots
(46.768.2%). However, all industrial sites had a relatively low percentage
of invaded plots (12.366.2%) when compared with small residential sites
(52.6621.9%). Riverine sites proved to be the most invaded with
84.0618.9% of the plots having at least one exotic species present. Differences in invasibility of these urban forested wetland are likely the result
of seed dispersal dynamics. I conducted a series of experiments that examine several components of seed dispersal including a seed trap study
that measures current seed flux, a seed bank survey which examines past
seed flux and a seed addition experiment that examines germination after
dispersal. The information garnered from these experiments will help to
build on our current level of understanding regarding the function surrounding land use in determining seed flux and site invasibility.
BOWMAN, WILLIAM P.,1 MARGARET M. BARBOUR,2 DAVID T. TISSUE,3 MATTHEW H. TURNBULL,4 DAVID WHITEHEAD2 and KEVIN
L. GRIFFIN.1 1 Columbia University, New York City, New York, USA;
2
Landcare Research, Lincoln, New Zealand; 3 Texas Tech University, Lubbock, Texas, USA; 4 University of Canterbury, Christchurch, New Zealand.
Influences of sap flow and respiratory activity on CO2 efflux from
woody stems in a New Zealand podocarp forest.
Measurements of CO2 efflux from woody stems and branches were collected for Dacrydium cupressinum, the dominant species in a podocarpbroadleaf forest in south Westland, New Zealand. An automated, multichambered system was utilized to make continuous measurements on the
stems of nine trees for up to 160 hours and on canopy branches for 24
hours. Temperature-normalized CO2 efflux rates from stems ranged from
0.51mmol m-2 s-1 to 2.43 mmol m-2 s-1, while respiration rates from branches
were generally larger, between 0.72 mmol m-2 s-1 to 6.8 mmol m-2 s-1. CO2
efflux rates were largely predictable from sapwood temperature; however,
efflux rates were lower than predicted from temperature alone during periods of maximum sapflow. This indicates that CO2 may be transported in
the xylem transpirational stream from respiring tissues in woody stems to
the canopy. The measurements of CO2 efflux were also analyzed with respect to wood density, cambial and sapwood respiratory activity, and tissue
nitrogen content. The results of this study highlight the importance of considering CO2 efflux from woody stems as a function of both the activity
of the respiring tissues within stems and the barriers to diffusion for CO2
that exist between the source tissues and the atmosphere.
BOYCE, RICHARD L.* Northern Kentucky University, Highland Heights,
KY. Choosing the best similarity index when performing fuzzy set ordination on abundance data.
Fuzzy set ordination (FSO) requires similarity indices that return values
between 0 and 1. Many indices that meet this requirement are available,
but their suitability for FSO with abundance data has not been tested. I
selected 12 indices that meet the above criterion; some are extended versions of presence/absence indices, following Tamas et al. (2001), and some
were modified to generate similarities between 1 and 0. Simulated plant

42

Abstracts

community data sets with variation along one gradient were generated with
COMPAS. Factors that were varied included beta diversity, species response shape, data standardization and qualitative and quantitative noise.
Sampling occurred along a regular sampling grid. Indices were evaluated
by performing rank correlations (Spearman r) between the apparent ecological positions generated by FSO and actual gradient positions; the abilities of the best-performing indices to minimize the curl-over effect, which
is similar to the horseshoe effect and affects samples near gradient endpoints, were also compared. Most indices performed well at low levels of
beta diversity (R,1). The Yule and Baroni-Urbani & Buser coefficients
also performed well at intermediate and high beta diversity levels, followed
by the Horn coefficient. With skewed species responses and noisy data, the
Yule and Baroni-Urbani & Buser coefficients were clearly the best choices.
A step-across algorithm improved correlations and reduced curl-over with
these two indices at R.1. Data standardization had little effect on any
index. Preliminary results suggest that these conclusions also hold for data
sets with two gradients. Thus, I recommend that one of the two best-performing indices be used when performing FSO on data with species abundances. A step-across algorithm should also be employed when analyzing
data sets with high beta diversities.
BOYD, ROBERT S. and EDWARD JHEE. Auburn University, Auburn,
AL. Elemental defense of Streptanthus species against herbivores: A
comparison of accumulated and hyperaccumulated levels of nickel.
Plants containing more than 1000 mg Ni/kg are considered nickel hyperaccumulators whereas those with lesser yet still elevated levels are nickel
accumulators. Hyperaccumulated nickel may defend plants from herbivores, but the defensive role of lesser levels of metals is unknown. We
raised five species of Streptanthus (Brassicaceae) native to serpentine soils,
one of which is a hyperaccumulator (S. polygaloides) and the other four
are accumulators, on Ni-amended and unamended greenhouse soils to create plants of differing Ni concentration. Slugs were fed plant material in
no-choice tests over a 50-day period and slug survival and mass changes
were recorded. All slugs fed high-Ni leaves of the hyperaccumulator species died within 21 days. Slugs fed high-Ni leaves of other plant species
did not significantly differ in survival or mass change from those fed lowNi leaves of those species. Choice tests using slugs offered both high- and
low-Ni S. polygloides plants showed a strong preference for low-Ni plants.
We conclude that hyperaccumulated Ni can defend S. polygaloides from
herbivory via both toxicity and deterrence, but these defensive effects do
not extend to Streptanthus species that simply accumulate Ni.
BOYER, ALISON G.* and MATTHEW D. MORAN. Hendrix College,
Conway, AR. Fire timing in a relict grassland: Effects on the plant
community.
We investigated the effects of a spring fire on the plant community in a
relict grassland and compared this experiment to one utilizing fall burning.
The field site was a naturally occurring grassland dominated by the perennial grass little bluestem (Schizachyrium scoparius) which co-occurs with
numerous other grasses and forbs. One-half of the plots underwent a prescribed burn on 1 April while other plots remained unburned. Plots were
sampled three times post-manipulation to determine seasonal changes in
the plant community. There was no effect of burning on overall plant biomass. However, forb biomass was weakly elevated during the summer in
burned plots, although this effect disappeared by the end of the season.
Grass biomass was not affected by burning. These results are in contrast
to a previous experiment that utilized a fall burn. In that experiment, forb
biomass was enhanced and grass biomass was reduced throughout the following growing season. This shows that the effects of fire are dependent
upon the season of burning which has implications for the management of
these relict grasslands.
BOYER, KATHARYN E.,1,* JOHN F. BRUNO1 and J. EMMETT DUFFY.2
1
University of North Carolina at Chapel Hill, Dept of Marine Sciences,
Chapel Hill, NC, USA; 2 College of William and Mary, Virginia Institute
of Marine Science, Gloucester Point, VA, USA. Biodiversity and ecosystem functioning in plant-grazer systems: experimental tests in a marine
benthic community.
Despite intense interest in the relationship between species diversity and
ecosystem functioning, the majority of research has focused at the base of

terrestrial food webs, and to a lesser degree, on aquatic microbial microcosms. Whether and how biodiversity might be related to ecosystem functional processes at higher trophic levels and in other ecosystem types have
received little attention. We conducted a series of experiments to explore
the relative effects of marine macroalgal and grazer identity and diversity
on net primary production in North Carolina hard substratum communities.
As in many previous grassland studies, we found a diverse assemblage of
algae to be more productive than the average monoculture, but not more
productive than all component species. However, overyielding was due not
only to the inclusion of some highly productive species but also to greater
performance of some species in mixture than in monoculture. High algal
diversity reduced consumption by a diverse assemblage of herbivores relative to consumption of the most palatable monocultures, perhaps reflecting
associational benefits of proximity to chemically defended species. In manipulations of herbivore diversity, net algal biomass production was two
times lower in the presence of a mixed grazer assemblage than in any of
the grazer monocultures, suggesting dietary complementarity among herbivores. In a factorial manipulation of algal and grazer diversity, we calculated variance components and found the magnitude of effects to be
greatest for the herbivore diversity factor. Although producer diversity enhanced net primary production, diversity at higher trophic levels may have
an equivalent or greater influence on the structure and function of this, and
perhaps other, ecosystems.
BRACHO, ROSVEL,1,* THOMAS POWELL,2 C. ROSS HINKLE3 and
BERT DRAKE. 1 National Research Council, Washington, DC, U.S.A;
2
Smithsonian Environmental Research Center, Edgewater, MD, U.S.A;
3
Dynamac Coprporation, Kennedy Space Center, FL, U.S.A. Comparing
evapotranspiration in Florida pine Flatwoods and Scrub-oak ecosystem
at the Merritt Island National Wildlife Refuge.
Florida pine Flatwoods and Scrub-oak are forest ecosystems with different
structures. At the Merritt Island National Wildlife Refuge we assessed seasonal and annual evapotranspiration (Et) in both forests using the eddy
covariance technique. Annual Et was 16% lower in the Scrub-oak ecosystem. The components of the energy budget (Et, sensible heat flux and soil
heat flux) were tightly related to the independent measurement of net radiation (Rn). Et represented 40% and 35% of Rn for the pine Flatwoods
and the Scrub-oak, respectively. Bulk canopy conductance (gc) was a
strong function of vapor pressure deficit (VPD) during the dry season (November-May) and the average of maximum gc was highly related to Rn
for the wet period.
BRADFORD, JOHN B.,1,2,* WILLIAM K. LAUENROTH,1,2,3 INGRID C.
BURKE1,2,3 and JOSE M. PARUELO.4 1 Graduate Degree Program in Ecology, Colostate State University, Fort Collins, CO, USA; 2 Department of
Forest, Rangeland and Watershed Stewardship, Colorado State University,
Fort Collins, CO, USA; 3 Natural Resource Ecology Laboratory, Colorado
a, UnivState University, Fort Collins, CO, USA; 4 Facultad de Agronom
ersidad de Buenos Aires, Buenos Aires, B.S.A.S., ARGENTINA. Controls
over large-scale ecosystem processes: The role of land use.
Identifying the conditions and mechanisms that control ecosystem processes is a central goal of ecosystem ecology. Ideas have ranged from single
limiting-resource theories to colimitation by nutrients and climate, to simulation models with edaphic, climatic and competitive controls. Although
some studies have begun to consider the influence of land use practices,
especially cultivation, few studies have quantified the impact of cultivation
at large scales relative to other known controls over ecosystem processes.
We utilized a 9-year record of productivity, phenology, climate, weather,
soil conditions and cultivation in the U.S. Great Plains to quantify the
controls over spatial and temporal vegetation patterns and estimate the process sensitivity to specific driving variables. We considered climate, soil
conditions and long-term average cultivation as spatial controls and weather
and interannual cultivation variations as temporal controls. We found that,
within this region, productivity variation is primarily spatial, whereas phenology variation is more evenly split between spatial and temporal components. Our models explained more of the variation in productivity than
phenology and more of the spatial than the temporal patterns. Our results
indicate that although climate is the most important spatial variable, cultivation explains a substantial fraction of the residual variability. Soil con-

ditions contributed very little to our spatial models. Weather and cultivation
deviation both made modest contributions to the temporal models. These
results suggest that the controls over phenology and temporal processes are
not well understood. Our sensitivity analysis indicates that productivity is
more sensitive to climate than weather and is very sensitive to cultivation
intensity. In addition to identifying potential knowledge gaps, these results
provide insight into the probable long- and short-term ecosystem response
to changes in climate, weather, and cultivation.
BRADLEY, KATE L.,* JOHANNES M.H. KNOPS and RHAE DRIJBER.
University of Nebraska, Lincoln, NE. Feedbacks between nitrogen availability, plants, and soil microorganisms.
Experiments involving nitrogen (N) fertilization have mainly focused on
plant responses, clearly demonstrating that increasing N availability increases primary productivity and may also lead to decreases in plant species
diversity. However, these studies have largely ignored potential feedbacks
that may occur between N availability, plants, and soil microorganisms,
even though soil microorganisms are considered the key regulators of decomposition and nutrient supply. In this study, we examined the in situ
effects of a one-time nitrogen pulse on the soil microbial communities
associated with the warm season grass, Schizachyrium scoparium. Nitrogen
was added in early June at the rate of either 0 gN/m2 (N 5 6), 16 gN/m2
(N 5 6), or 80 gN/m2 (N 5 6). Soil CO2 flux, soil microbial biomass, soil
community composition, and soil available N (ammonia + nitrate) were
quantified four times throughout the growing season (before fertilization
and 2, 4, and 8 weeks after fertilization). Soil CO2 flux was significantly
higher in the 16 gN/m2 treatment than in the control or 80 gN/m2 treatments
throughout the growing season. More interestingly, N fertilization significantly decreased microbial biomass throughout the entire growing season.
Control plots had significantly higher amounts of microbial biomass at the
three sampling periods following application of the fertilization treatment
(adjusted means: 97, 115, 133 nmol lipid/g soil) compared with both the
16 gN/m2 (adjusted means: 92, 102, 118 nmol lipid/g soil) and 80 gN/m2
(adjusted means: 90, 100,106 nmol lipid/g soil) treatments. These results
support the hypothesis that under situations of N limitation plants can stimulate soil microorganisms, through rhizodeposition, to actively decompose
SOM, and therefore increase N mineralization. When their N limitation is
alleviated through anthropogenic sources, plants no longer need to maintain
as large of a soil microbial community to meet their N needs. It is likely
that they reduce both rhizodeposition and their root biomass and invest
their carbon resources to aboveground tissues.
BRADSHAW, G. A.* Oregon State University, Corvallis, Oregon, USA.
Complexity, Ethics, and the Postmodern Dilemma.
By emphasizing relationships, complexity theory helps bridge conceptual
and disciplinary schisms. Unlike most other scientific theories, however,
complexity not only includes insights and description of the observed world
such as an ecosystem or socio-ecological landscape, but also brings the
observer, science and scientists, within the domain of study. This conceptual re-orientation shifts from a model of science and scientists wherein
society and the problems they address are seen as separate, to a model of
participation where fundamental epistemological assumptions and purpose
of scientific enquiry become topical. In this view, complexity re-configures
the epistemological and ethical framework in which science is engaged and
compels discussion concerning the implications for scientific institutions,
concepts, and subject matter of science itself.
BRAGG, DON C.,1,* ERIC HEITZMAN2 and JAMES M. GULDIN.3
1
USDA Forest Service, Southern Research Station, Monticello, AR, USA;
2
University of Arkansas-Monticello, Monticello, AR, USA; 3 USDA Forest
Service, Southern Research Station, Hot Springs, AR, USA. Age class
distribution of a virgin shortleaf pine stand.
A recently uncovered set of shortleaf pine (Pinus echinata Mill.) increment
cores from the Lake Winona Research Natural Area (LWRNA) in the
Ouachita National Forest of Arkansas was assessed for age and increment.
According to the core sample, this uneven-aged virgin stand contains shortleaf pine that date back to the mid-1700s, although most individual trees
are less than 130 years old. Several prominent spikes in decadel represen-

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43

tation suggest major recruitment events occurred in the 1870s to 1890s and
again in the 1920s, possibly arising from disturbances related to the exploitation and settlement of the Ouachita Mountains. Canopy recruitment
declined dramatically from 1930 to 1990 (with very few shortleaf pine
originating after 1950), corresponding to the implementation of effective
fire control in this region. Since shortleaf pine responds well to frequent
burning and most hardwoods do not, the loss of fire from the LWRNA has
contributed to the virtual cessation of pine establishment and the development of a prominent oak, hickory, maple, and gum midstory. Additionally, small-scale lightning, wind, insect, and disease events continue to
claim old overstory shortleaf without much potential for their replacement.
The LWRNA, created to preserve a rare old-growth shortleaf pine remnant,
stands to gradually lose its pine component and become a hardwood-dominated forest if major natural disturbances like fire continue to be excluded.
BRAKE, ROBIN A.* and RUTH ANN KERN. Biology Department, California State University, Fresno, Fresno, CA, USA. An investigation into
Pinus sabiniana range discontinuity in the Kings, Kaweah and Tule
watersheds, California, USA.
Blue oak woodlands are Californias most common hardwood forest, covering nearly three million acres. These woodlands are endangered due to
increasing blue oak senescence, as well as environmental stresses related
to pollution, suburban expansion, rural building, firewood harvesting, and
agricultural/ranching enterprises. Pinus sabiniana is a dominant endemic
species of the valley and foothill woodlands surrounding the San Joaquin
Valley in California. P. sabiniana, or gray pine, is inexplicably missing in
a fifty-five mile segment on the southeastern portion of its potential range.
This study hypothesized that P. sabiniana does not grow in the gap due to
an environmental gradient established by slope and aspect compared to
those areas that do support the species. Field sampling was conducted in
foothill woodland research sites situated north, south, and within the range
gap at 15 locales in Fresno, Tulare, and Kern counties. Data was collected
for 900 point samples during the spring/summer season 2002 and 2003.
All tree and shrub species within each point sample were identified and
recorded along with the environmental variables of slope, aspect, potential
radiation, topographic position, slope configuration, elevation, and the nature and degree of disturbance. Data will be examined using Canonical
Correspondence Analysis (CCA) and analysis of covariance to describe the
relationship between P. sabiniana distribution and physical site factors.
This study is significant in providing much needed research into a historically under-studied community. Because the gray pine is a major associate
in the blue oak foothill woodland community, solving the P. sabiniana
range discontinuity mystery will add one more piece of information required to sustain the community in a scientifically appropriate manner.
BRAMBLE, DENNIS M.1 and RAYMOND M. TURNER.2 1 University of
Utah, bramble@bioscience.utah.edu, Salt Lake City, UT; 2 U.S.G.S. - Desert Research Laboratory, Tucson, AZ. Documenting long-term persistence of the western harvester ant (Pogonomyrmex occidentalis).
The nests of the western harvester ant, Pogonomermyx occidentalis, are
conspicuous features on the landscape of the western United States. These
structures, consisting of a pyramidal mound and a surrounding cleared
zone, appear to be remarkably long-lived as well. Using repeat photographic and dendrochronologic techniques, we have obtained evidence for minimum site persistence of P. occidentalis ranging from many decades to
over a century at two environmentally contrasting study sites in Utah. At
a lower elevation (1920m), desert-scrub locality in west central Utah (Millard Co.), all anthills discernable in an historic 1901 photograph were still
occupied by ants in 2001. Periodic repeat photography and direct monitoring indicate that most of these ant sites have been continuously occupied
since at least 1969. Repeat aerial photography and big sage (Artemesia
tridentata) dendrochonology were used to document ant persistence at a
cooler, wetter, and higher elevation (2210m) pinyon-juniper site in south
central Utah (Garfield Co.). At this locality, dozens of individual sites inhabited by harvester ants in 1960 still possess active colonies at this time.
Growth ring series from sage shrubs bordering the cleared zones (together
with other historic data) strongly suggest that many of these ant-dominated
sites predate Anglo settlement and livestock grazing in the area. If so, such
sites have been occupied by harvester ants more or less continuously for

44

Abstracts

at least 120 years. Although targeted recolonization of abandoned sites is

likely a factor in the long-term persistence of P. occidentalis, we so far
have little direct evidence of this. Our observations indicate that P. occidentalis is probably the most enduring, site-specific animal in the American
West. The extraordinary persistence of this ant is all the more impressive
given the dramatic, anthropogenically induced changes in vegetation that
have occurred at both study sites.
BRASSIL, CHAD E.* University of Toronto, brassil@zoo.utoronto.ca, Toronto, ON, Canada. The similar effect of intrinsic versus extrinsic cycles
on mean population densities.
The long-term mean density of a population across time is a basic measure
that is widely used to summarize populations with variable densities. I
demonstrate the similar affect of cycling on the mean density, whether the
cycling is due to extrinsic or intrinsic noise. Intrinsic cycles are those cycles
driven by density dependence or population interactions such as predation.
Extrinsic cycles are those driven by random variation in the environment,
reflected as variation in a model parameter. The analyses of intrinsic and
extrinsic cycles have largely developed along independent lines, and by
many measurements this separation is entirely justified. However, with regards to the long-term mean density, a basic and widely utilized indicator
of a fluctuating population, the affect of intrinsic and extrinsic cycles can
largely be summarized by an examination of density variance. In simple
models such as a discrete logistic equation, or the Ricker equation, a similar
expression describes the effect of both extrinsic and intrinsic variation on
the mean density. In most standard population models, intrinsic or extrinsic
variation will decrease the mean density compared to the equilibrium. Depending on the form of non-linearity, variation could lead to an increase
in the mean density, as in some formulations of the Beverton-Holt equation.
This analysis is extended to two and three species systems, demonstrating
how variation interacts with processes such as competition and predation.
The similarity of extrinsic and intrinsic variation is based in part on the
assumption that external environmental variation is independent among
generations. This assumption is examined by looking at the effect of correlated environmental variation.
BRAY, SARAH R. University of Florida, sbray@botany.ufl.edu, Gainesvillle, FL, 32611-8526. Microbial community function and composition
over a range of geographical locations, plant communities and invasive
plant species.
Invasion of natural systems by non-native plant species has been shown to
alter nutrient cycling, nutrient regimes and native plant biodiversity. These
effects of invasive species may be due to unique qualities of the invader
or the dominance of that species in the community. The majority of this
information, however, has come from case studies of individual species,
limiting our ability to make conclusions about invasive species effects
across systems. For a standardized assessment of the effects of invasive
species on microbial communities across a geographical range, I examined
soil samples of communities heavily impacted by five non-native plant
species in Florida and Texas: Sapium sebiferum, Meleleuca quinquenerva,
Schinus terebenthifolius, Ardisia crenata, and Ruellia tweediana. Soil samples were taken from both non-invaded and invaded areas for each species.
Microbial community function was determined by Biolog and composition
was analyzed by extraction of microbial phospholipid fatty acids (PLFA).
Carbon substrate utilization patterns from Biolog as analyzed by principle
components analysis showed little difference between invaded and noninvaded areas within and across species. While the metabolic abilities of
the microbial communities were not altered by plant invasion, the identities
of the microbes as determined by PLFA did change. These results suggest
that while invasion may alter the species of microbes present in the soil,
the function of the community remains intact.
BREEN, AMANDA N.* and JAMES H. RICHARDS. University of California, Davis, Davis, CA. Water and nutrient limitations affect seed
number, size, and quality: Implications for establishment of two desert
shrubs.
Plants have limited resources that they must allocate to growth, survival,
and reproduction. It is theorized that larger seeds will have at least an initial

advantage over smaller seeds and subsequent seedlings will be better able
to establish under stressful environmental conditions. I tested the effects of
fertilizer and nutrient additions on seed production in Sarcobatus vermiculatus and Atriplex parryi by measuring seed number per length of branch,
seed weight, seed viability, seed quality, and initial seedling length. The
results are for two field seasons. Additionally, I looked at the effects of
treatment on seeds in one subsequent generation in a controlled environment experiment. I found that both the number of seeds produced
(p50.036) and the weight per seed (p50.017) increased with additions of
fertilizer over that of water alone. Seed width increased with treatment
(p50.001). Seed viability was mixed for S. vermiculatus but increased
under treatment for A. parryi (p50.041). Seedling size at three days was
highest in fertilizer treatments for both species (p50.034). Seed quality
analyses support these results. In both species, seed nitrogen content was
higher in treated plants (p50.02). In a growth chamber experiment, I found
that large seeds were able to withstand 200mM NaCl while smaller seeds
could not. These results indicate that larger seeds contain more nitrogen,
have higher percent viability, and produce significantly larger seedlings,
enabling them to more successfully establish under stress.
BRESHEARS, DAVID D.,1,* GEOFFREY B. WEST,1 JAMES H.
BROWN,2 CRAIG D. ALLEN3 and BRIAN J. ENQUIST.4 1 Los Alamos
National Laboratory, Los Alamos, NM; 2 University of New Mexico, Albuquerque, NM; 3 US Geological Survey, Midcontinent Ecological Science
Center, Los Alamos, NM; 4 University of Arizona, Tucson, AZ. Biophysical scaling relationships for tree structure and stand patterns: Extensions to semiarid ecosystems.
Recently developed theory on biophysical scaling relationships quantitatively predicts how many biological attributes of size and energy are related. The theory has been applied from molecular through biosphere
scales. There are four critical hypotheses relative to the theory: (1) The
flow of biological energy at all scales is limited by a hierarchical resource
distribution network. (2) A constant amount of energy is required to produce unit biomass at all size scales ("energy equivalence"). (3) Metabolic
power scales as a fractional power function of mass at all levels of biological organization. (4) Variations from predicted behavior result from nicherelated adaptations driven by evolution (species-dependent variation) or
dynamic responses to recent perturbations (environmentally dependent variation). Here we test several aspects of theory using data on plant community structure in a semiarid pinyon-juniper woodland. Our findings indicate that for both of the co-dominant woody species, Pinus edulis and
Juniperus monosperma, large trees exhibit a trend similar to the predicted
scaling relationship that the number of tress of a given size is proportional
to the inverse diameter squared. These relationships did not differ between
species. Small trees, however, violate the predicted relationships, thereby
providing insight into ongoing ecosystem processes. Both species exhibit
scaling trends within individuals relating basal diameter and height that are
similar to those predicted by theory. These results extend the predicted
scaling relationships, which have thus far been tested primarily in tropical
species to semiarid ecosystems with a more patchy structure. Further, the
results highlight how deviations from the theory can provide a diagnostic
for patterns and processes in biology.
BREWER, CAROL.* Division of Biological Sciences, University of Montana, Missoula, MT, USA. Integrating Undergraduate Research Experiences From Matriculation through Graduation.
There are natural connections between teaching, research, and learning that
can engage students in the thrill of scientific discovery. I will illustrate how
a research-rich environment can be created to provide high-quality educational experiences for undergraduate students from their first biology course
through graduation by describing a curriculum innovation program at the
University of Montana. Over the last four years at UM, we have introduced
a series of new team-taught introductory biology courses featuring a combination of lectures, active learning techniques designed for large classrooms, and small group discussions to introduce undergraduates to cutting
edge research. To date, more than half of the tenure-line faculty in Biology
have taught in this sequence. Core concepts are presented using examples
with research themes. Lectures are complemented by research-centered laboratories where students work in collaborative teams to explore research

questions they generated from the topics covered during the lectures and
follow-up discussions. Many lab investigations in the introductory biology
sequence have formed the basis for larger-scale student research projects
that have been presented at annual campus-wide research symposia. This
curriculum innovation program also has supported the development of new
student-centered, research-based approaches to teaching existing upper division courses. Today, hundreds of undergraduates participate in research
experiences as part of their biology course training every semester. One
outcome of this approach is that the number of students seeking capstone
research experiences, including independent research and participation in
faculty research, has increased substantially. Focusing on integrating research experiences from matriculation through graduation has changed the
landscape of undergraduate biology education at UM, with research programs providing important platforms and opportunities for student learning
(as well as generating new knowledge) and student and faculty research
informing and enlivening what goes on in the classroom.
BREWER, ELIZABETH A.,1,* ELDOR A. PAUL2 and SHERRI J. MORRIS.1 1 Bradley University, Peoria, IL; 2 Colorado State University, Fort
Collins, CO. Role of nitrogen and calcium in stabilization of soil carbon
in forest soils.
Carbon (C) sequestration in soils is important for soil fertility and may play
a role in short-term mitigation of elevated CO2 on global change. Unfortunately, the mechanisms for C stabilization are poorly understood. Previous studies have determined that nitrogen (N) and soil cations, especially
calcium (Ca) are essential to C storage. Our research evaluates the importance of N and Ca, especially the form of Ca, in stabilization of soil C.
Laboratory incubations were established using soil from an afforested red
pine plantation with relatively low C content. We chose this specific site
because earlier field studies have suggested that soils under pine stands
with higher Ca content have greater C sequestration. Incubated soils were
amended with Ca added at a rate consistent with adjacent high C soils and
N at twice and six times atmospheric deposition rates for this area. The Ca
and N amendments were added separately and together with and without
incorporation of litter. The addition of Ca in the form of CaCl2 decreased
the amount of CO2 lost in the first 45 days of incubation when compared
to control soils. The addition of Ca in the form of CaCl2 also resulted in
less CO2 loss than Ca in added in the form of lime and N addition treatments. Incorporation of litter increased CO2 loss in all treatments but lime
additions consistently resulted in the greatest CO2 loss. Addition of N at
twice and six times atmospheric deposition rates for this area resulted in
CO2 evolution rates comparable to unamended control soils. These results
suggest that stabilization of C through the addition of Ca is dependent on
form added and N availability. Maximizing C sequestration by increasing
stabilization may be achieved but requires greater understanding of the
mechanisms controlling C storage in soils.
BREWER, STEPHEN.* University of Mississippi, jbrewer@olemiss.edu,
University, Mississippi. Resource Competition and Fire-Regulated Nutrient Demand in Plants of Longleaf Pine Savannas.
Longleaf pine savannas are fire-dependent ecosystems typically characterized by relatively nutrient-poor soils. It is generally agreed that repeated
fires reduce competition and maintain plant species diversity in these systems. One documented effect of fire is the reduction of aboveground competition through the destruction of aboveground parts of plants and the
combustion of associated litter. In addition to fire, low soil fertility is
thought to maintain species diversity in pine savannas. Two alternative
hypotheses are offered to explain species coexistence in nutrient-poor soils.
1) Plants avoid competition for nutrients with one another by occupying
different niches at fine spatial scales. 2) Slow growth and low nutrient
demand limit competition. Recent investigations of competition between
carnivorous pitcher plants and their neighbors have shown that, in years
without fire, depriving pitcher plants of prey does not reduce their competitive ability. Apparently, low soil nutrient demand and slow growth obviate intense competition between pitcher plants and their neighbors. However, nutrient demand (measured as investment in carnivory) is a plastic
trait, which increases significantly following fires. Because most fires in
pine savannas cause little mortality and tend not to damage belowground
parts of plants, increases in nutrient demand following fire could lead to

Abstracts

45

increased belowground competition for nutrients. Hence, complementary

nutrient use by carnivorous and non-carnivorous plants after fires could
contribute to coexistence. I argue that studies that attempt to test alternative
hypotheses about competitive mechanisms in longleaf pine savannas should
consider the effect of fire-regulated plasticity in nutrient demand.
BREZINSKI, BRIAN,* KRISTIN DEGROOT* and EVAN WEIHER. University of Wisconsin - Eau Claire, Eau Claire, WI. Patch- and landscapelevel effects on bird diversity in an oak savanna - bottomland hardwood forest mosaic.
Local vegetation structure, habitat geometry, and the quality of the surrounding landscape can affect bird diversity, but rarely have all of these
factors been simultaneously addressed. Furthermore, we know of no cases
where structural equation modeling has been used to assess the independent
effects of these factors. We mapped the main vegetation units (habitat
patches) in a 1200 ha mosaic of bottomland hardwood forest, oak forest,
oak savanna, prairie, and shrub thicket in western Wisconsin and conducted
four replicate breeding bird counts at 90 sampling points (from late May
to early July). Vegetation data included measurements of cover, structural
complexity, composition, and diversity. We used GIS to determine patch
geometry (area, area-perimeter ratio, distance to edge), and surrounding
landscape context (surrounding habitat diversity at five scales). In general,
standardized richness (species richness divided by the log of the number
of detections) increased with vegetation cover and evenness, with bird
abundance, and with landscape diversity. We used structural equation modeling to determine the independent effects of four latent variables on standardized richness: landscape diversity, patch geometry, local vegetation,
and bird abundance. The final structural equation model explained 33% of
the variation in standardized richness. Vegetation and bird abundance independently had twice the effect of landscape factors, while habitat geometry was not significant. Landscape diversity and evenness within a 200
m radius (about 12.6 ha) had the strongest effects on bird richness, and
this suggests a possible critical scale for birds in this system.
BRIGHT, KERRY L., CAROL A. BREWER,* CAROL SNETSINGER
and ALISON E.H. PERKINS. University of Montana, Missoula, MT, USA.
Fatal Flower frailties: Addressing misconceptions about evolutionary
theory found in nature films.
Science and nature films often are used in instruction because they engage
students through appealing visual and sound effects. Educators must use
these media with caution, however, because they often are produced to meet
entertainment rather than educational goals. How can conceptually flawed,
but beautifully produced, media be used in the classroom without promoting misconceptions? We present a method that uses flaws in educational
videos to promote conceptual change in students. In this study, we used a
visually stimulating, but conceptually flawed, video to help college students
confront their conceptions about evolutionary theory. After watching a video about co-evolution between orchids and their pollinators, students were
asked to do two things: predict how Darwin would have explained this
phenomenon; and examine prepared explanations, some of which contained
evolutionary misconceptions. Initially, a significant proportion of students
proposed Lamarckian explanations, and, when evaluating the prepared explanations, students considered teleological (need- or goal-based) explanations acceptable. After class discussions, students re-examined the prepared explanations, and this time many students identified the inconsistencies with evolutionary theory. Ultimately, students were asked to apply
their understanding on an exam by evaluating the supposition that bat
wings evolved because shrew-like ancestors needed to fly to catch food.
Indeed, most students now recognized the flaws in this explanation of bat
wing evolution. At the end of this unit, students were surveyed to explore
what classroom experiences influenced any changes they had in understanding evolutionary concepts. Discussions in lecture were reported the most
influential, followed by critiquing the prepared answers.
BRINKMAN, ROBERT B.,* WILLODEAN BURTON and STEVE HAMILTON. Center for Field Biology, Austin Peay State University, Clarksville,
Tennessee, United States. Evaluation of Crayfish as a Keystone Species.
ABSTRACT A keystone species is critical to a community and has the
ability to affect more than one trophic level. Crayfish are ecologically im-

46

Abstracts

portant to small stream ecosystems and have been described as keystone

species. They occupy numerous trophic levels and are instrumental in making coarse particulate organic material (CPOM) into fine particulate organic
material (FPOM) and available for other macroinvertebrates in streams.
Two enclosure/exclosure experiments were conducted in 2002 to test the
role of Orconectes placidus as a keystone species. An experiment to determine the impact crayfish have on processing leaf litter was started in
February and lasted 28 days. A second experiment, lasting 21 days, to
study possible impacts crayfish have on the abundance of other macroinvertebrates was started in May. Orconected placidus significantly reduced
the weights of leaf packets they were enclosed with. There were no significant differences in the abundances or richness of macroinvertebrates
between crayfish enclosure channels and exclosure channels, used as controls.
BRISBIN, I. LEHR1,* and MICHAEL STUREK.2 1 Savannah River Ecology Laboratory, Aiken, SC, USA; 2 School of Medicine, University of Missouri, Columbia, MO, USA. Invasive aliens with thrifty genes: Diabetic
feral swine on a southeastern coastal barrier island.
Over three decades of study of the feral swine of Ossabaw Island, Georgia
suggest that during the nearly 500 years that these animals have existed in
a feral state, they have developed a number of unique physiological traits
which might well confer fitness in an environment of fluctuating and unpredictable food (acorn mast crop) availability. Of particular interest are
those traits related to fat storage and utilization, which have produced a
"metabolic syndrome" phenotype, the precursor to type 2 diabetes. The
occurrence of the metabolic syndrome in these swine without the usual
sequelae of cardiovascular disease has brought these animals to the attention of the biomedical research community. Of particular interest is the
opportunity to use studies of these free-ranging swine in their island habitat,
to better understand the "thrifty gene hypothesis" for the evolutionary origin and maintenance of this now epidemic disease condition in humans.
In Ossabaw Island feral swine moreover, this condition occurs without apparent detrimental effects within the population. However these swine are
invasive aliens with a clear potential to negatively impact other island resources (e.g. sea turtle nesting beaches). This creates unique challenges for
integrating biomedical research interests with goals for the long-term management and control of these swine as components of this barrier island
ecosystem. It also suggests the importance of collecting basic biological
data from removed offenders in all programs designed to control or eliminate invasive exotics - particularly those with a long history of residence
and potential adaptation in the environments that they are impacting.
BROCKWAY, DALE G.,1,* KENNETH W. OUTCALT2 and BECKY L.
ESTES.1 1 USDA Forest Service, Auburn, AL; 2 USDA Forest Service,
Athens, GA. Restoring southern pine forest ecosystems: Plant community response to mechanical midstory reduction and prescribed fire
on sandhills at Fort Benning, Georgia.
Developed during periods of fire exclusion, dense midstory vegetation, that
reduces understory plant diversity (competitive shading) and increases the
risk of damaging wildfire (fuel ladder from ground to canopy), has impeded
restoration efforts to safely reintroduce prescribed burning in southern pine
ecosystems. Our study evaluated the effects of midstory reduction on plant
community structure, composition and diversity in forests treated by mechanical means alone and also when followed by prescribed fire during the
winter, spring or summer. Mechanical treatment caused declines in overstory and midstory tree density (1220 to 258/ha) and basal area (31 to 18
m2/ha) and a corresponding increases in mean DBH (13 to 29 cm), with
the largest reductions in Pinus taeda, Liquidambar styraciflua, Quercus
nigra and Q. hemisphaerica. Despite declines in tree species richness (9 to
4) and diversity (1.32 to 0.84), increased evenness (0.72 to 0.85) indicated
improved distribution equity among residuals. Understory tree cover declined 17% overall and cover increases of 9% were observed for shrubs &
vines (Smilax and Vitis expansion) and grasses (Chasmanthium appearance). Forb cover increases were very modest, averaging , 4%. Understory
plant diversity remained generally unchanged following treatment, with
increases noted for species richness corresponding to declines in evenness.
While not producing large initial growth increases in herbaceous plants,
prescribed fire is needed following mechanical treatment to stimulate grasses and forbs and curtail redevelopment of the woody plant midstory.

BRODERSEN, CRAIG R.,1,* MATTHEW J. GERMINO2 and WILLIAM

K. SMITH.1 1 Wake Forest University, Winston-Salem, NC; 2 Idaho State
University, Pocatello, ID. Carbon uptake limitations in Abies lasiocarpa
and Picea engelmannii across an alpine treeline ecotone.
The debate over the mechanistic factors that limit the altitudinal limit of
forest trees to specific elevations, and no higher, has been a source of
controversy for over a century. Environmental effects on both photosynthetic carbon gain and respiratory-driven growth processes have been used
to evaluate limitations at the alpine treeline. It was hypothesized here that
microsite factors associated with ecological facilitation are more important
for determing the potential for photosynthetic carbon gain than altitudinal
differences. Throughout the summer of 2002 adult trees of the codominant
Abies lasiocarpa (subalpine fir) and Picea engelmanii (Engelmann spruce)
were selected for gas exchange measurements (Li-COR 6200) of net photosynthesis, as well as microlimate, at three different altitudes that spanned
the width of the alpine treeline ecotone (3,154 m; 3,205 m; 3,345 m). High
elevation alone did not appear to be a limiting photosynthetic activity, as
the two highest sites studied showed the highest daily values of carbon
gain (0.7, 1.2, 1.9 mmol m-2s-1, respectively), as well as integrated annual
values (0.96, 2.20, and 1.79 mmol m-2s-1, respectively). Additional data also
support the idea that microsite sky and wind exposure (both day and night),
and associated biophysical factors, may be more definitive in predicting
carbon gain at treeline than the effect of altitude. Microsite differences in
sky and wind exposure may be responsible for higher carbon gain at the
middle elevation site compared to either the forest or upper ecotone sites.
Adaptive trade-offs between too much and too little sun exposure, cold
nighttime sky exposure, and wind effects reflect a complex interaction between needle temperatures, low temperature photoinhibition of photosynthesis, and water relations related to snow burial and deposition patterns.
BROOKS, KATHERINE E.* and JEROME J. HOWARD.* University of
New Orleans, kebrooks@uno.edu, New Orleans, Louisiana, USA. The effects of Chinese tallow (Sapium sebiferum) on bird populations in a
Louisiana bottomland forest.
Invasive species are seen more now than ever as a serious threat to biodiversity. In the Gulf States, the Chinese tallow tree (Sapium serbiferum)
has been steadily creating monocultures in native Louisiana bottomland
forests. Several studies on Chinese tallow have shown that this tree can
significantly alter its surrounding environment. Chinese tallow has been
shown to alter soil chemistry and hydrology, thereby affecting germination
rates of native species. Additionally, Chinese tallow seems to be resistant
to pathogens and herbivores. The germination and growth rates of Chinese
tallow tend to out-pace those of native trees, allowing rapid establishment
in old growth forests. The primary method of seed dispersal of Chinese
tallow has been shown to be through birds. For these reasons we hypothesized that a higher diversity and greater abundance of birds would be
found in native forests with few tallow trees. Likewise, a lower diversity
and smaller number of birds should be found in areas with high tallow
establishment. We conducted several point counts at the Bayou Savauge
refuge in eastern New Orleans, Louisiana. We established 12 different transects with three different levels of tallow. Through these point counts we
found a tendency towards higher levels of diversity in the low tallow sites,
with a greater abundance of birds overall. Additionally, we found lower
levels of diversity in high tallow sites. Somewhat surprisingly, we found
moderate levels of diversity and abundance in the sites with medium levels
of Chinese tallow in the understory. Canopy cover may prove to be a more
important factor for retaining high levels of biodiversity in birds than attempting to completely rid an old growth forest of an invader. This finding
may provide support for new management techniques when dealing with
an aggressive invasive species such as the Chinese tallow tree.
BROOKS, MELODY A.,* RAELYNN DEATON and EDIE MARSHMATTHEWS. University of Oklahoma, Norman, Oklahoma. Effects of
body condition on reproduction in western mosquitofish, Gambusia affinis.
Energy for current reproduction may come directly from environmental
food sources or energy fat stores. Using females that differed in body
condition, we examined maternal provisioning with external resources in a

livebearing fish by injection of radiolabeled nutrients. Gambusia affinis

females were collected from a pond in central Oklahoma, isolated in 3.7
L aquaria and randomly assigned to four food treatments in an attempt to
manipulate body condition (flake food, frozen brine shrimp, algae, and
dried fish). After 24 days females were injected with tritiated leucine to
assay mother-to-embryo transfer of nutrients (measured as embryo radioactivity in disintegrations per minute). Two hours post injection, females
were dissected and embryos removed from the ovary, dried, weighed and
prepared for liquid scintillation assay. Female condition was determined by
extraction of fat with multiple washes of petroleum ether. Body condition
differed significantly among food treatments: females fed algae or dried
fish were in poorer condition. Despite condition variation, all females
showed transfer of radiolabeled nutrients to developing embryos. Females
in poorer condition did not show decreased investment in embryos: embryo
number, embryo weight, and maternal transfer of nutrients did not vary
significantly with body condition. Because reproduction was not compromised by decreased body condition, we suggest that energy-limited females
will invest more in current than future reproduction, whereas females in
good condition may invest in both.
BROWN, BRYAN L.* Dartmouth College, Department of Biological Sciences, 6044 Gilman, Hanover, New Hampshire, USA. Quantifying sources of temporal variability in a stream macroinvertebrate community.
Natural ecosystems vary temporally due to species interactions (intrinsic
factors) and the extrinsic pressures exerted by a continually fluctuating
environment. I used observational and experimental studies coupled with
statistical techniques to quantify the influence of multiple sources of variability on stream macroinvertebrate communities in two 2nd order streams
of the Second College Grant, NH. In the summer of 2001, I performed an
observational study to examine the influence of substrate spatial heterogeneity on community temporal variability. Meso-scale spatial heterogeneity explained considerable variability in community variability (r250.58,
p50.011 with Rommes relative evenness as a heterogeneity index) and
within-stream community variability decreased over 40% with observed
increases in spatial heterogeneity. However, an experimental manipulation
of stream spatial heterogeneity in the summer of 2002 failed to reproduce
the results of the 2001 observational study, yet overall community variability was very similar in both 2001 and 2002. Causes of this discrepancy
may include 1) failure of the experiment to reproduce a gradient of spatial
heterogeneity similar to that in the observational study of 2001, 2) massive
drought in 2002, 3) sources of extrinsic variability other than spatial heterogeneity, or 4) the influence of intrinsic community variability. To further
investigate temporal variability in these stream communities, I plan to apply
several statistical techniques to the 2002 experimental data, including multivariate autoregressive models (MARs), to determine which sources of
variability significantly contribute to observed community temporal variability. Understanding how and why communities vary through time is
appealing from both an academic desire to understand the functioning of
nature, and its applicability to ecosystem restoration and/or management.
BROWN, JOHN B.,* CELIA A. EVANS, JANET A. MIHUC and RUTH
M. SMITH. Paul Smiths College, Paul Smiths, NY. Scaffolding scientific
inquiry across a Bachelors Degree in Natural Resources: Preliminary
assessment of student outcomes.
During the first year of an NSF Course Curriculum and Laboratory Improvement Grant (CCLI), investigators at Paul Smiths College developed
and implemented laboratory activities to help students construct content
knowledge and an improved understanding of the scientific process. First
year students in introductory biology and stream ecology courses (Phase
I) worked with hypothesis development and empirical data collection in
the lab and field. Students in their second year (Phase II) learned ecological
concepts through collaborative exercises using computer simulations, some
of which employed real data sets and ecological scenarios based on actual
research. Seventy percent of biology students in Phase I reported learning
something new about hypothesis development as a result of the activity.
The two most common responses were that hypothesis development was
difficult to do well, and that hypotheses are not always supported by the
scientific process. Results from the Student Assessment of Learning Gains
(SALG) instrument we administered at the end of the biology course sup-

Abstracts

47

ported that students improved their understanding of hypothesis development (3.33 of 5). Students in stream ecology improved their understanding
of stream invertebrate microhabitats (78%) as a result of the activity. Most
commonly reported concepts were that different stream insects were found
in different microhabitats, and that physical factors vary across the stream
environment. Only ten percent of students in Phase II reported that using
ecological models did not improve their understanding of concepts, even
though twenty-eight percent reported no desire to take another course in
which computer models are used. Lack of prior computer skills was a
common reason given. Preliminary data from Phase III model development
activities and an analysis of surveys given to students completing their
senior capstone research will also be presented.
BROWN, KIM J. and RITA GIULIANI.* Dept. of Environmental and Plant
Biology, 405 Porter Hall, Ohio University, Athens, OH, USA. Landscape
and treatment influences on the spatial heterogeneity in leaf area productivity in thinned and burned oak-hickory forests of SE Ohio.
Highly managed horticultural or silvicultural forest systems typically display a fair correlation between stand basal area and leaf area index (LAI),
but factors such as differences in species productivity and distribution on
the landscape, nutrient limitations, complexities in canopy architecture, and
anthropogenic management regimes make simple allometric solutions to
predicting leaf area elusive in natural forest stands. Temporal heterogeneity
in forest phenology further complicates accurate measures of seasonal LAI
production. In order to model forest carbon dynamics in recently thinned
and burned oak-hickory forests in SE Ohio, research was conducted to
assess landscape, treatment and species influences on the spatial heterogeneity of LAI. Leaf litter traps (n5120) were deployed in September 2002
at a US Forest Service FFS study which featured three replicate locations
within Vinton County, Ohio, four treatments (thin, burn, thin & burn, control), and ten plots within each treatment. Trap material was collected weekly through December and monthly thereafter. Leaf material was sorted into
oak, maple, and "other" classes and dried and weighed, and LAI was calculated using area:dry mass conversion factors. LAI values ranged from
2.0 to 7.0, with mean values of 4.0 to 4.6. Significant treatment effects
were observed with the thinning treatments imparting the largest reductions
of LAI with burning alone having minimal effects. Relationships between
LAI and the following variables will be evaluated: plot basal area, speciesgroup basal area, soil nitrogen, and soil moisture index (after Iverson et
al. 1997). The accuracy of optical LAI detection techniques will be evaluated and presented, with particular attention to canopy heterogeneity of
leaf aggregation and vertical distributions.
BROWN, NINA A.1,* and ROBERT A. RAGUSO.2 1 SUNY Stony Brook,
Stony Brook, NY; 2 University of South Carolina, Columbia, SC. Targeting pollinators and evading herbivores: Floral scent emission in two
species of Cirsium.
Floral volatile emissions attract pollinators to reproductive structures and
thus promote cross-pollination. By increasing the apparency of reproductive
parts, plants face the danger of attracting floral herbivores with the same
signals that they use to lure pollinators. In two species of thistle, Cirsium
arvense, a dioecious, exotic invasive species, and C. repandum, a native
to the southeastern United States, we have investigated the potential mechanisms by which these plants modify volatile emissions to avoid attracting
detrimental insects. Using a combination of analytical chemistry and field
research, we examined the variability in emissions in relation to the foraging patterns of both pollinators and herbivores, including phenological
preference and diel patterns. We found that in C. arvense the peak in floral
volatile emissions correlates with the peak in pollinator activity rather than
temperature. In contrast, C. repandum tracks temperature rather than the
diel pattern of pollinators, peaking in scent emissions at the hottest time
of the day. Nighttime, the coldest time of the day, is also the low point in
scent emission for both thistles. While pollination in C.arvense does not
occur at night (in the population we studied), C. repandum receives nocturnal pollination, at a level equivalent to daytime intensity. In order to
avoid attracting herbivores, scent should be minimized prior to and following reproductively receptive times. In C. arvense androecious plants produce more scent then gynoecious plants. In fact, the full blend is not represented in gynoecious inflorescences until florets begin to emerge. In both

48

Abstracts

thistle species there is a drop in scent emissions following bud break, and
both peak in scent production at full maturity. We found no potential repellent compounds prior to flowering. Following flowering, methyl salicylate increases in C. repandum suggesting a potential role as a repellent.
BROWN, REBECCA L.,1,* JASON D. FRIDLEY2 and JOHN F. BRUNO.2
The Academy of Natural Sciences, brown@acnatsci.org, Philadelphia,
PA; 2 University of North Carolina, Chapel Hill, NC. Null models and
observed patterns of native and exotic diversity: Does native richness
repel invasion?

In studies of species invasion, growing evidence from surveys of natural

habitats suggests that positive or non-significant native-exotic richness relationships at larger scales appear to give way to negative relationships at
small scales. The small-scale negative relationship has been taken as evidence that small-scale species diversity is an effective deterrent to invasion,
but this argument fails to consider expected native-exotic richness relationships under a null model of no interspecific interactions with different sample sizes. With simple simulations, we show that small sample scales produce negative relationships between native richness and exotic richness
(and invasion frequency) in the absence of native-exotic interactions. Furthermore, when samples are allowed to intrinsically vary in total species
richness (due to different rates of propagule supply or resource availability,
for example), positive relationships between native and exotic richness are
unavoidable except at the smallest scales. Departures from this null model
can be assessed with observational data by a permutation test that randomly
permutes the "native" versus "exotic" identities of each species. Our use
of this technique on plant compositional data from riparian and coastal
strandline communities reveals that observed patterns of native and exotic
richness are remarkably similar to the expected relationship generated by
a no-interaction null model, suggesting that native species richness plays
little role in regulating species invasion.
BROWNE, DAVID R.* and JOSEPH B. RASMUSSEN. Department of
Biology, McGill University, 1205 Dr. Penfield Ave., Montreal, Quebec,
Canda. Changes in ontogenetic diet shifts of brook charr in response
to the presence of yellow perch: A possible example of compensationdepensation.
Fish community composition can have a large impact on fish population
structure and dynamics. Brook charr show highly flexible life history traits
that may vary in response to habitat gradients, resource levels, and predation. In this study we use gut content data and stable isotope analysis to
examine the interaction between brook charr and yellow perch. We determined the trophic position and resource use of brook charr in a set of four
lakes containing cyprinids and white suckers either with or without yellow
perch present. Carbon isotopic signatures of brook charr tissue indicated
brook charr shift from reliance on a mixture of benthic and pelagic resources to a mixture of benthic and profundal resources when perch are
present in the lake. Gut content analyses were consistent with isotopic
results and indicated a high degree of piscivory and a reduced contribution
of zooplankton and large zoobenthos to brook charr diet when yellow perch
were present in the lake. Brook charr trophic position increased with fork
length to a maximum of 3.9 in lakes without yellow perch. When yellow
perch were present in the system, the trophic position of brook charr decreased with fork length from small to intermediate size classes then increased with length to a maximum of 4.3 in large size classes. Analysis of
brook charr condition factor across size classes in lakes containing perch
indicates a bottleneck in late juvenile growth in contrast to non-perch lakes.
Results suggest juvenile brook charr suffer competition from yellow perch
for food resources while perch are the primary prey item of adult charr.
This size-structured intra-guild predation interaction suggests brook charryellow perch communities may exhibit compensation-depensation dynamics.
BRUHN, JOHANN N.* and JEANNE D. MIHAIL. University of MissouriColumbia, Columbia, MO. Landscape effects of pathogens: Armillaria
root pathogens and long-term forest dynamics.
Armillaria is an agaric genus of white-rot wood decay fungi. Most Armillaria spp. can cause opportunistic lethal root disease. The host ranges of

Armillaria spp. differ yet overlap. Parasitic aggressiveness differs both

within and among species. Armillaria genets are initiated by the mating of
compatible basidiospore-derived primary mycelia. Genets are potentially
long-lived and indeterminate in size, spreading vegetatively through the
forest floor among woody food bases. The geographic distribution patterns
of Armillaria spp. depend on: 1) the influences of existing ecological landtype characteristics on vegetation pattern and each Armillaria species resource aquisition strategy; and 2) the influences of past disturbance regimes
(logging, fire, episodic climatic stress) on vegetation pattern and health and
the temporal supply of vulnerable woody foodbases. Aggressive Armillaria
genets take advantage of the interactions among predisposing site-related
factors and patterns of successive predisposing disturbance events to infect
and gradually kill individual vulnerable root systems. When conditions result in the co-occurrence of Armillaria spp., competition for substrate influences the activity of each species, and thereby the geographic pattern of
root disease related tree mortality. Because Armillaria genets are cryptic,
the spatial patterns of species and genet distributions across landscapes are
poorly understood without intensive study, and the underlying roles of Armillaria spp. as disturbance factors contributing to many forest decline
syndromes have largely gone unrecognized until recently. Armillaria root
disease is the principal mortality agent in upland Ozark oak decline, where
Armillaria spp. function both as long-lived disturbance factors and as catalysts for further disturbance, in a feedback manner. Armillaria root disease
related disturbance influences the spatial pattern of forest vegetation in
several ways. As a result, root disease helps shape the habitat for diverse
forest lifeforms, with implications for the continuity of Armillaria itself.

assumption that plant populations in different habitat fragments are demographically distinct has not been tested. To do so, I used populations of
the understory herb Heliconia acuminata located in both the fragments and
the continuous forest reserves of The Biological Dynamics of Forest Fragments Project (Brazil). Using annual censuses initiated in 1998, I constructed transition matrices for thirteen populations: four in 1-ha fragments,
three in 10-ha fragments, and six in continuous forest. These matrices were
compared with log-linear models to determine if demography varied temporally (from year to year in the same plot) and spatially (among plots in
the same year). Transition matrices for populations in continuous forest
were highly variable, differing both temporally and spatially. This contrasts
with populations in 10-ha fragments, for which temporal variation in demography was significant but spatial variation was negligible. Finally, the
fit of models for populations in 1-ha fragments was not improved by including time, location, or the interaction of the two, indicating there was
neither spatial nor temporal variation in transition matrices. These results
suggest a new way fragmentation can negatively affect plants in habitat
fragments by reducing demographic variation at the population level.
These reductions in demographic variability could severely limit the responses of populations in fragments to changing biotic and abiotic conditions, as well as increase the probability of extinctions due to landscapescale stochastic events.

BRULAND, GREGORY L. and CURTIS J. RICHARDSON. Duke University, Durham, NC. Using Mantel tests to investigate spatial variability, soil properties, and phosphorus sorption in riparian wetlands.

As residents of freshwater streams, pleurocerid snails are constantly subject

to wide variation in drag forces induced by changes in water flow rates.
Shells of these snails also serve as solid substrates upon which green algae
can grow. It is known that algae can either increase or decrease drag resistance of rocks in streams depending upon growth form, but drag effects
of algal growth on snails are relatively unknown. We sampled pleurocerid
snails which differed in their degree of algal colonization from an Ozark
river. Drag resistance on empty shells was measured in a flume in the
laboratory soon after field collection and after all algae had been scraped
from the shells. Drag resistance under both high and low flow was significantly positively related to frontal area of shells as well as the area/length
ratio (an indicator of shell shape), except in scraped shells at low flow.
Thus, we were able to detect subtle differences in flow resistance even over
small differences in size and shape of shells. While average drag was lower
on scraped shells than on shells with algae intact, this difference was not
significant, suggesting that algal growth within the ranges we studied does
not affect flow resistance, at least within the confines of the flume. Future
research will include a wider range of algal growth levels as well as more
explicit study of algal species-specific effects.

Soils of forested riparian wetlands are highly effective at phosphorus (P)

sorption. However, these soils exhibit extreme spatial variability across
riparian zones. We used a spatially-explicit sampling design in two forested
riparian wetlands in North Carolina to better understand the relationships
among P sorption, soil properties, and site variability. Our objectives were
to quantify spatial variability of P sorption and related soil properties, to
determine which soil properties best explained the variability in P sorption
after accounting for spatial autocorrelation, and to compare parametric
Pearson correlations to non-parametric Mantel correlations. We measured
bulk density, pH, soil organic matter (SOM), texture (% clay, % silt, %
sand), oxalate extractable Al (Alo), Fe (Feo), and the P sorption index (PSI).
The two sites had substantially different mean PSIs due to differences in
Alo, Feo, and % clay. We also found considerable differences in the spatial
variability of soil properties at each site. For example, pH, Alo, Feo, and
PSI exhibited autocorrelation at site one (S1), while Alo, Feo and PSI exhibited autocorrelation at site two (S2). Kriged maps illustrated that soil
properties at S1 appeared to vary at smaller scales than those at S2. After
accounting for the effects of autocorrelation, we determined Alo to have
the highest Mantel correlation with PSI at both sites. While Pearson correlations would have led us to conclude that PSI was significantly related
to most of our selected soil properties, partial Mantel correlations revealed
several of these relationships to be spurious. We believe the Mantel approach is robust and could also serve as a model for research of controls
on other biogeochemical processes such as denitrification. This type of
spatially-explicit P sorption research provides insight on how to develop
more accurate landscape-scale water quality models and on how to reproduce natural variation in mitigation of wetlands.
BRUNA, EMILIO M.1,2,* 1 Department of Wildlife Ecology & Conservation, University of Florida, Gainesville, FL, USA; 2 Tropical Conservation
and Development Program, Center for Latin American Studies, University
of Florida, Gainesville, FL, USA. Habitat fragmentation and the demographic variability of an Amazonian understory herb.
While the detrimental consequences of habitat fragmentation for species
diversity have been widely documented, the effect of fragmentation on the
demography of remnant populations is poorly understood. Some theoretical
models predict populations in fragments will have asynchronous dynamics;
this may promote persistence at the landscape level since multiple populations are unlikely to go extinct simultaneously. To date, however, the

BRUNKOW, PAUL E.* and LISA A. BAUER. Southern Illinois University

Edwardsville, Edwardsville, IL, USA. Algal growth and flow resistance
in a freshwater snail.

BRUNNER, JESSE L. and JAMES P. COLLINS. The importance of a

complex life history in the long-term persistence of a lethal virus in
seasonally abundant salamander populations.
The transmission of most infectious diseases is a density dependent process.
Below a critical host population size or density, the contact rate between
infected and susceptible hosts is too low to sustain the chain of transmission
and the parasite dies out in the host population. Sustained transmission is
least likely for virulent parasites in slowly increasing host populations (high
transmission rates and short infectious periods, coupled with low rates of
birth or introduction of susceptible hosts). Thus, virulent parasites are not
expected to persist in the small, slowly growing populations typical of
endangered species unless the parasite has an abundant, widespread reservoir host. Such reservoirs can maintain virulent parasites endemically;
reintroducing disease into smaller, vulnerable populations when they come
into contact. The ranavirus-tiger salamanders system is an exception to this
paradigm. This lethal, directly transmitted virus causes recurrent epidemics
in the dense larval segment of salamander populations. However, salamander larvae metamorphose and disperse to burrows to overwinter, precluding
much transmission. This virus degrades quickly in the environment and has
no reservoir host. Rather persistence between epidemics is achieved by
infrequent, long-lived, chronic infections in the metamorphosed salamanders, which are normally highly susceptible. Both life history stages are
apparently essential for the persistence of these lethal viruses: larvae am-

Abstracts

49

plify the prevalence of virus infection; metamorphs maintain infections

between epidemics and reintroduce the virus when they return to ponds to
breed. I will present the results of a mathematical model examining how
the complex life history of tiger salamanders helps maintain this dynamic
host-pathogen system.
BRUNO, JOHN,1,* LAURA PETES2 and ANNALIESE HETTINGER.3
The University of North Carolina at Chapel Hill, Chapel Hill, NC;
2
Oregon State University, Corvallis, OR; 3 Dalhousie University, Halifax,
Nova Scotia, Canada. Nutrient enrichment can increase the severity of
Caribbean coral diseases.

Disease is the primary cause of the decline of reef-building corals in the

Caribbean. Coral cover on most reefs has been reduced to , 5% over the
last 20 years. Several studies indicate that disease prevalence and the subsequent rate of coral mortality have increased dramatically during this period. One possible explanation is that human-induced changes in environmental conditions have impaired host resistance and/or increased pathogen
virulence. We tested the hypothesis that elevated nutrient concentrations
can increase the severity of coral diseases. We focused on yellow band
disease in two species of the reef-building coral Montastraea and on Aspergillus infections of the sea fan Gorgonia ventalina. Both diseases are
common at our study sites in Akumal, Mexico. We manipulated nutrient
concentrations on naturally and experimentally infected individual colonies
with Osmocoat fertilizer and compared the rates of disease spread and
tissue death between treatment and control colonies. For all three species,
nutrients significantly increased rates of disease spread and tissue loss by
50 to 70%. Although the dynamics and effects of coral disease epidemics
are probably controlled by a variety of factors, these results suggest that
nutrient enrichment could significantly increase the mortality of three key
Caribbean species.
BUCKLEY, LAUREN B.* and JOAN E. ROUGHGARDEN. Stanford University, lbuckley@stanford.edu, Stanford, CA. Understanding the Ecological Development of Spatial Diversity Patterns through ThermalPhysiology-Based Null Modelling.
The complexity of ecological communities discourages mechanistic understanding of how spatial diversity patterns form, yet this knowledge is needed to predict how biotas respond to perturbations. Anolis lizard communities on Lesser Antilles islands are simple enough to evaluate the influence
of both abiotic factors and species interactions on spatial diversity patterns.
On a regional scale, anoles on St. Kitts, a northern island, and Grenada, a
southern island, reveal the greatest species overlap and rates of species
turnover at differing elevations. On a local scale, anole species in the northern island group segregate habitat with respect to perch height, whereas
those in the southern island group segregate habitat with respect to microclimate. An individual-based model of anole population dynamics, which
includes species-specific thermal physiology profiles and predicts equilibrium abundance based on foraging energetics, is combined with GIS data
to predict anole abundance at each island location. Discrepancies between
the empirically and theoretically predicted distributions indicate the influence of topography and species interactions on spatial diversity patterns.
We suggest that the more mountainous topography of the southern islands
results in more broad-scale habitat segregation and divergent thermal physiologies. In contrast, the high geographic species overlap on the predominately low elevation northern islands compels partitioning along a perch
height axis within a location.
BUCKLEY, THOMAS N.,1,2,* KEITH A. MOTT3 and GRAHAM D. FARQUHAR.1,2 1 Environmental Biology Group, Research School of Biological
Sciences, Australian National University, Canberra, ACT, Australia; 2 Cooperative Research Centre for Greenhouse Accounting, Research School of
Biological Sciences, Australian National University, Canberra, ACT, Australia; 3 Biology Department, Utah State University, Logan, UT, USA. A
hydromechanical and biochemical model for stomatal conductance.
We present a new model for stomatal conductance in intact leaves. The
model is based on principles of plant water relations, leaf gas exchange,
and epidermal hydromechanics. Guard cell osmotic pressure is assumed
proportional to ATP concentration, which is calculated from a popular mod-

50

Abstracts

el of photosynthesis. Unlike other stomatal models, ours accommodates the

epidermal mechanical advantage, which causes the initial response to hydraulic perturbations to be in the wrong direction (e.g., opening when
humidity decreases), but which is overcome by hypothesizing that the sensitivity of guard cell osmotic pressure to ATP is proportional to epidermal
turgor. The model accurately predicts stomatal responses to variations in
ambient humidity, ambient carbon dioxide, ambient oxygen, irradiance, xylem resistance, soil water potential, and photosynthetic capacity. Most elements of the models structure are readily interpreted in terms of reduced
processes at the cellular level, but the model itself is easily expressed in a
simple form that is identical to the Michaelis-Menten expression for enzyme-catalyzed reaction rates.
BULTMAN, THOMAS L. Hope College, Holland, MI. Reversal of
wound-induced resistance in a grass is mediated by a fungal endophyte
and constrains tolerance.
Most plants respond to herbivore damage with elevated chemical and/or
structural mechanisms of resistance. However, some plants, particularly
those with few chemical defenses, may actually become more susceptible
to herbivores following damage; that is, damage may induce susceptibility.
We tested the hypothesis that a fungal symbiont of tall fescue grass provides induced resistance to aphids by manipulating fungal infection, plant
damage (clipping) and level of nutrient fertilizer in greenhouse/laboratory
experiments. The fungus had a negative effect on aphid reproduction that
was most pronounced following damage. This result appears due to elevated levels of alkaloids produced by the fungus after damage. Furthermore, plants lacking fungal infection were more susceptible to aphids following damage. Hence, while plant damage induces susceptibility in uninfected plants, it does the opposite (induced resistance) in infected plants.
Additional experiments showed that the reversal of induced susceptibility
is influenced by intrinsic (fungal strain) as well as extrinsic (nutrient application to plants) factors. We also found that the induced resistance conferred by the fungus comes at the cost of tolerance (reduced regrowth) in
infected plants. As a modulator of both resistance and tolerance, the fungus
plays a central role in influencing defense of host plants against herbivores
and should have important implications for plant-herbivore interactions in
natural and agricultural systems.
BUNKER, DANIEL E.* and WALTER P. CARSON. University of Pittsburgh, Pittsburgh, Pennsylvania. Experimental test of resource competition and keystone predation theories: Predicting the success of invasions and biological control.
Resource competition theory predicts that the species that reduces the availability of the limiting resource (R*) to the lowest level will win in competition. Keystone predation theory (sensu Grover) predicts that a specialist
herbivore will reverse the outcome of competition if it reduces the resource
competitive ability of its host (i.e., raises the R* of the host above that of
its competitors). We experimentally tested these predictions using invasive
purple loosestrife (Lythrum salicaria), native broad-leaved cattail (Typha
latifolia), and Galerucella calmariensis, a specialist beetle released to control loosestrife. To measure how species and herbivory affect resource
availability (R*), we established 6 cattail and 24 loosestrife monocultures
(1000 l cattle tank mesocosms), at low and high fertility (0, 16 gNm-2 year1
) (60 total). We manipulated beetle density manually, thereby creating an
herbivore damage gradient. We then measured light availability at the soil
surface to measure species and herbivore impacts on light availability.
Without herbivory, loosestrife reduced light availability to lower levels than
cattail at both low and high fertility, and therefore was the superior competitor for light. Herbivory reversed this pattern. When herbivory reached
30% (low fertility) and 60% (high fertility) leaf-area damage, cattail reduced light availability lower than loosestrife. Thus, our monocultures predict 30% (low fertility) and 60% (high fertility) leaf-area damage will reverse the outcome of competition. We tested these predictions experimentally in 48 additional mesocosms where adult loosestrife and cattail competed with and without herbivores. After two years of competition, relative
abundance of loosestrife was higher than cattail in the absence of herbivory.
Competitive outcomes were reversed when herbivory levels reached 40%
(low fertility) and 55% (high fertility) leaf-area damage, very similar to
our predictions from monoculture. Our results suggest that if Galerucella

damage does not reach these high levels in the field, biocontrol will not
succeed.
BURKE, ANNA M., MARIUS VAN DER MERWE and JOEL S.
BROWN. University of Illinois at Chicago, Chicago, IL. Effects of seasonality on foraging in urban versus natural squirrel populations.
In two populations of fox squirrels, Sciurus niger, we measured the effect
of seasonality on foraging responses to microhabitat (near and away from
cover) and cacheability of food (hazelnuts with or without shells). We chose
our populations of squirrels to reflect an urban habitat (Oak Park, IL) and
a more natural area (the Morton Arboretum in Lisle, IL). Urban squirrels
seem to have two advantages over their natural counterparts; there is more
food available and less predation. But these conditions lead to higher squirrel densities in urban areas; meaning urban squirrels should actually be
more food limited. We measured giving-up densities (GUDs) of fox squirrels in the two populations of squirrels from Winter 2001 to Fall 2002 to
test predictions about the effects of seasonality in the two different locations. We predicted GUDs would be the lowest in winter (when populations
are food limited) and highest in the fall (when natural food resources are
more abundant), and that urban squirrels would have lower GUDs. Overall,
GUDs were approximately 10% higher in summer than in fall and winter.
Interestingly, overall GUDs were lower in the Arboretum squirrel population than the urban population by 17% (about one nut per foraging bout),
contrary to our predictions. Season had opposite effects on the urban and
non-urban squirrels. Oak Park squirrels had their highest GUDs of the three
seasons in the summer, while Arboretum squirrels had their lowest GUDs
in the summer. We expected urban squirrels to show a stronger preference
for cacheable nuts and weaker distinction between safe and risky microhabitats than non-urban squirrels. Urban squirrels did have a stronger preference for cacheable nuts, but they actually showed more of a distinction
between microhabitats. Seasonality appears to influence foraging; however,
it does not necessarily have similar effects on all squirrel populations.
BURKE, INGRID C.,1,2,* WILLIAM K. LAUENROTH,1,2 JACK A. MORGAN,3 ARVIN R. MOSIER2,3 and ROGER A. PIELKE.4 1 Dept of Forest,
Range, and Watershed Stewardship, Colorado State University, Fort Collins, CO, USA; 2 Natural Resource Ecology Lab, Colorado State University,
Fort Colins, CO, USA; 3 USDA Agricultural Research Service, Fort Collins, CO, USA; 4 Dept of Atmospheric Sciences, Colorado State University,
Fort Collins, CO, USA. Carbon management, land use and global
change: feedbacks among carbon, nutrient and water cycles.
The central grassland region of the U.S. is intensively managed for livestock and grain production, resulting in a matrix of rangelands, dryland
cropping, and irrigated cropping. Each of these general classes of management has a different impact on carbon, nutrient, and water cycles, and the
distribution of the practices in time and space integrates to influence the
regional cycles. We have collected long term data on carbon, water, and
nitrogen fluxes in the shortgrass steppe portion of the central grasslands,
across multiple landuse types, using a variety of techniques including inventories of C and N stocks, estimates of trace gas fluxes, and Bowen ratio
measurements of carbon dioxide and water exchange. In general, water is
the main limitation over carbon inputs, and carbon and water availability
are the main controls over nutrient retention. In rangelands, carbon and
nitrogen cycling are enhanced by livestock grazing. Dryland cropping also
enhances carbon and nitrogen cycling, and alters the seasonality and partitioning of evaporation and evapotranspiration vs. soil water storage. Carbon losses are significantly increased by cultivation management, and nitrogen trace gas losses are also stimulated by N fertilization. Irrigated land
use management has the largest impacts on the relative magnitude of carbon, water, and nitrogen fluxes. Interestingly, although water and nitrogen
are substantially increased by human additions, and net primary production
increases in response, there is no apparent net change in carbon and nitrogen balance of the ecosystem. Biomass exports, increased volatile losses,
and enhanced evaporation all balance the increased inputs. The current
regional impact of all three landuse types is a substantial increase in nitrogen trace gas production, nitrogen leaching, and evaporation, and a decrease in methane uptake. There is strong evidence that these changes, as
well as surface albedo, have a major impact on regional mesoscale climate.

BURKEPILE, DERON E.,* JOHN D. PARKER, BROCK WOODSON and

MARK E. HAY. Georgia Tech, Atlanta, Georgia, USA. Microbial chemical defense of carcasses: Testing Janzens "Why Fruits Rot, Seeds
Mold, and Meat Spoils" hypothesis.
Interactions of microbes with higher organisms are often described as mutualistic (e.g., gut symbionts) or parasitic (e.g., pathogens), but rarely as
competitive. Microbes that colonize rich food sources such as fruits and
animal carcasses, however, should be selected to defend their food, and
thus themselves, from being eaten by larger consumers. We investigated
the possibility that marine microorganisms colonize and protect fish carcasses from marine scavengers. Rotten carrion attracted fewer consumers
than fresh carrion and was unpalatable to some consumers. In feeding assays, chemical extracts from rotten carrion were unpalatable. Microbes
probably mediate the unpalatability of carrion given that carrion "rotted"
in an antibiotic solution was palatable to consumers. Thus, microbiallyproduced chemicals are probably responsible for the chemical repugnance
of rotten carrion. Our data support Janzens (1977) untested hypothesis that,
meat spoils because that is the way microbes compete with bigger organisms. Because microbes are ubiquitous components of communities and
produce a variety of bioactive secondary metabolites, microbial defense of
resources from larger organisms may be an under-appreciated feature of
community dynamics.
BURLEYSON, TRAVIS,1,* STEPHANIE ALBRECHT,1 KURT SCHULZ,1
JOHN ZASADA2 and WILLIAM MATTSON.3 1 Southern Illinois University Edwardsville, Edwardsville, IL, USA; 2 North Central Forest Experiment Station, Grand Rapids, MN, USA; 3 North Central Forest Experiment
Station, Rhinelander, WI, USA. Leafminer damage to eastern leatherwood (Dirca palustris) in relation to plant spacing and size.
Eastern leatherwood (Dirca palustris) is a small, very shade tolerant deciduous shrub that inhabits Acer saccharumTsuga canadensis forests of
the western Great Lakes Region. Leatherwood populations tend to be
strongly aggregated owing to limited seed dispersal. With one exception,
leatherwood foliage is strikingly unattractive to mammalian and insect herbivores. Larvae of the moth Leucanthiza dircella mine the leaves during
midsummer. Leaf damage can be very extensive, resulting in the loss of
whole leaves and significant fractions of total leaf surface (.20%) by late
July. Loss of photosynthetic surface in dark forest understories has potential
impacts on survivorship, growth, and reproduction. We examined how rates
of leaf damage were affected by two variables (1) distance to nearest congener, and (2) size of the target individual. Our hypotheses were that increasing distance would be associated with lower damage rates, and that
smaller individuals would experience higher damage rates. There were
strong effects of stand identity on the role of plant spacing. In one stand
herbivory rates were negligible (,0.5% leaf area); in a second stand herbivory increased with distance; in a third stand herbivory declined with
distance. Landscape level variation in leaf miner populations apparently
overrides the influence of spacing within leatherwood populations. Herbivory rates did not vary with respect to plant size. Smaller individuals did,
however, experience a wider variance in the level of herbivory among
leaves on single shrubs. Wider variance is likely associated with larger
numbers of severely damaged leaves that may subsequently be lost.
BURTON, MICHAEL G.,1,* THEODORE M. WEBSTER,2 ERIC P.
PROSTKO,3 A. S. CULPEPPER,3 ALAN C. YORK1 and SHANNON
SERMONS.1 1 North Carolina State University, mike_burton@ncsu.edu,
Raleigh, NC; 2 USDA-ARS, twebster@tifton.usda.gov, Tifton, GA; 3 University of Georgia, eprostko@arches.uga.edu, Tifton, GA. Rapid increase
of tropical spiderwort (Commelina benghalensis L.) in herbicide-resistant crops of southeastern USA agroecosystems.
Tropical spiderwort (Commelina benghalensis L.) is a terrestrial invasive
weed species. As a consequence of multiple herbicide tolerances, it is on
the federal noxious weed list. Although previously not known to occur
north of Georgia, this troublesome weed species was positively identified
in North Carolina in September 2001. Affected states are now known to
include California, Florida, Georgia (where it is now considered the most
troublesome, although not the most common, weed species in agro-ecosystems), Louisiana, and North Carolina. This species has been observed

Abstracts

51

to increase rapidly in intensively managed, frequently disturbed, cropping

systems. Notably, tropical spiderwort has shown the most dramatic increase
where herbicide-resistant crops are used in successive years of crop rotations. Although the National Plants Database (http://plants.usda.gov) only
reports tropical spiderwort as occurring in three counties in south Georgia,
preliminary results (20% of counties responding) from a statewide survey
indicate that this species occurs in at least 29 counties, and is considered
a moderate or greater problem in $17 counties. Results from growth chamber experiments demonstrate that temperature optima for plant growth are
in the range of 30 to 35o C, which suggests that the climate throughout the
southeast USA may be suitable for rapid growth and reproduction of tropical spiderwort. In herbicide resistant cropping systems, tropical spiderwort
populations escape weed management tactics with a broad period of germination (i.e., multiple germination cohorts), rapid production of rhizomes
that can both result in vegetative reproduction and production of viable
seeds in cleistogamous subterranean flowers, and an ability to vegetatively
regenerate from cut stem sections (provided that stems are buried less than
8-cm). Tropical spiderwort also produces aerial chasmogamous flowers that
can produce viable seeds within 25-d of flowering. Population densities of
other members of the Commelinacea are also reportedly increasing where
herbicide-resistant crops are frequently used. Coordinated state, regional,
and federal efforts/funds are needed to prevent further spread and economic
injury from tropical spiderwort.
BUSHAW-NEWTON, KAREN L.,1,* DANIELLE A. KREEGER,2 SARAH DOATY1 and DAVID J. VELINSKY.2 1 American University, Washington, DC, USA; 2 Academy of Natural Sciences, Philadelphia, PA, USA.
Relative effects of dissolved organic matter derived from Spartina alterniflora and Phragmites australis on secondary production.

zones. Three year old tulip poplar (Liriodendron tulipifera L.), red maple
(Acer rubrum L.) and sweet gum (Liquidambar styraciflua L.) saplings
were exposed to a total seasonal N application of 74, 37, 18 and 0 kg N/
ha from May through October 2002. The trees were grown in pots, exposed
to full sun, and provided daily water at the USDA Agricultural Research
Service, in Beltsville, MD. Measurements of adaxial and abaxial leaf surfaces were made throughout the growing season and included fluorescence
imaging, fluorescence emission spectra (300 - 800 nm, 5 nm resolution),
reflectance spectra (350 - 2500 nm, 3 nm resolution) and various biophysical parameters. Vegetation fluorescence occurs when light energy is absorbed by a leaf surface, is not utilized for photosynthesis and is emitted
at a lower energy and longer wavelength. The actively induced fluorescence
spectra of green plants typically exhibit five distinguishable maxima centered in the UV (335 nm), blue (445 nm), green (525 nm), red (685 nm)
and far-red (740 nm) regions of the spectrum. Fluorescence emission spectra were obtained at three discrete excitations: 280 nm (280 Ex), 360 nm
(360 Ex) and 530nm (530 Ex). Significant differences among tree species
were obtained for: a) adaxial emissions centered at 360 nm and 375 nm
resulting from 280 Ex (p , 0.001); b) adaxial emissions in the blue region
from 360 Ex (p , 0.001); and c) adaxial emissions in the red peak, due
to 530 Ex (p # 0.001). The adaxial green/far-red fluorescence ratio (360
Ex) also exhibited species separation (p # 0.01). In addition, there were
significant N treatment differences in the red/far-red fluorescence ratio (360
Ex) in red maple, with the fluorescence ratio increasing with N treatment
(r2 5 0.74, p # 0.01). These findings demonstrate that individual tree
species can be discriminated using spectral signatures and that N related
differences could be determined using fluorescence measurements. These
results are encouraging, particularly in light of the fact that they are observations from the first year of a multiple year N application study.

Historically, salt marshes along the mid-Atlantic US have been dominated

by Spartina alterniflora, but presently, many of these marshes have been
invaded by Phragmites australis. Although, P. australis has high rates of
primary production, it is not known whether this species supports a marsh
food web in the same manner as S. alterniflora. Using several related photochemical and biological assays, we compared patterns of organic matter
flow of plant primary production through a key salt marsh metazoan (Geukensia demissa) using a bacteria intermediate. Whole plants were collected
from Delaware Bay salt marshes and grown in the laboratory with either
[14C]-CO2 (14C) or [15N]-inorganic N (15N). Bacterial utilization of plantderived dissolved organic matter (DOM) measured as carbon mineralization (14C), or bacterial production (15N) over time revealed that both types
of DOM were available to the natural bacterioplankton, but utilization of
Spartina-derived DOM was consistently higher than utilization of Phragmites-derived DOM. For the 14C bioassays, total carbon mineralization
after 19 days measured 35 % (+3 S.D.) for the Spartina-treatments compared with 29% (+2, S.D.) for the Phragmites-treatments. When bacteria
were fed to mussels, both plant source and sunlight regime yielded significant effects on clearance rates. Mussels cleared Spartina-derived products
at 0.38 L h-1 [g dry tissue weight] -1, more than twice that for Phragmitesderived products 0.15 L h-1 [g dry tissue weight]-1. As with the bacterial
bioassays, no clear patterns were attributable to sunlight exposure with light
positively affecting clearance rates for Phragmites-products but negatively
positively affecting clearance rates for Spartina-products. Results from 15N
analyses indicate that the level of [15N]-enrichment may not have been high
enough to supply mussels with enough isotope to resolve their N budget.
Overall, these results suggest the importance of understanding the linkages
between primary and secondary production in marsh systems.

BUTTERFIELD, SCOTT* and CAROLYN M. MALMSTROM. Michigan

State University, East Lansing, MI. Using remote sensing to assess forage
dynamics in a California rangeland restoration project.

BUTCHER, LADEAN M.,1,* ELIZABETH M. MIDDLETON,1 PETYA

ENTCHEVA-CAMPBELL,1 LAWRENCE A. CORP,1 JIM E. MCMURTREY2 and EMMETT W. CHAPPELLE.1 1 Laboratory for Terrestrial Physics, Biospheric Sciences Branch, NASA Goddard Space Flight Center,
Greenbelt, MD, USA; 2 Hydrology and Remote Sensing Laboratory, USDA
Agricultural Research Service, Beltsville, MD, USA. Spectral fluorescence signatures of tulip poplar, red maple and sweet gum under increasing nitrogen fertilization levels.

During summer 2000 I surveyed intertidal beaches of the San Juan Islands,
WA to determine the spatial distribution and abundances of native and nonnative infaunal bivalves. Three nonindigenous clam species were highly
abundant in the surveys (Venerupis philippinarum (5Tapes japonica), Nuttallia obscurata, and Mya arenaria). Venerupis philippinarum is the most
prolific of these species and might affect native clams; therefore, I conducted a manipulative experiment to examine its effect on an ecologically
similar, confamilial clam, Protothaca staminea. I manipulated Venerupis
density and predator exposure in an orthogonal design to quantify the effects of Venerupis and clam predators on the mortality, growth, and fecundity of Protothaca at six different sites throughout the archipelago. Environmental covariates such as chlorophyll, crab abundance, and sediment

An experiment was designed to characterize the spectral fluorescence and

reflectance properties of the foliage of several tree species over a range of
nitrogen (N) conditions that they may encounter in agricultural riparian

52

Abstracts

Rangeland restoration projects require information about forage dynamics

over large areas. As part of a rangeland restoration project with AudubonCalifornia, we used Landsat Enhanced Thematic Mapper (ETM+) satellite
data to quantify changes in rangeland forage over a three-year period in
the Upper Willow Slough Watershed, in Yolo County, California. We calibrated relationships between green biomass values and the normalized difference vegetation index (NDVI) using a field spectroradiometer and a
series of vegetation harvests. The biomass-NDVI relationships were then
used to produce time series maps of forage for the watershed from ETM+
imagery. The time series of forage values reveal rangeland responses to
different management techniques, including prescribed burns, prescribed
grazing, and reseeding of native grasses. An important aim of this project
is to facilitate improved management decision-making by the private landowners in the watershed. The initial reaction of most landowners to the
satellite-based forage maps was that the maps had to be wrong. After evaluating the maps carefully, however, each landowner changed his or her
mind and began to ask for more information about the forage dynamics in
each of their pastures. Remote sensing offers a powerful tool to enable
voluntary improvements in management of rangelands by private landowners.
BYERS, JAMES E.* Univ. of New Hampshire, Durham, NH, jebyers@
unh.edu. Ecological impacts of nonindigenous clams on natives in the
San Juan Islands, Washington.

organic content were also tracked at each site. Results indicate that high
densities of Venerupis had little effect on Protothaca suggesting that there
was little direct effect of competition for space or resources between the
clams. However, Venerupis grew up to five times better and suffered significantly higher mortality than its native analogue (particularly at sites with
high crab abundance)factors that are parsimoniously explained by Venerupis shallow burial depth compared to Protothaca. In a complementary
prey choice experiment conducted simultaneously in the lab, crab predators
preferred Venerupis over Protothaca. Higher production of Venerupis coupled with its high susceptibility to crab predation may be boosting regional
crab abundance and thus heightening predator-mediated competition on
Protothaca and other native bivalves.
BYRD, KRISTIN. University of California, Berkeley, kbyrd@nature.
berkeley.edu, Berkeley, CA. Linking watershed land-use change and
wetland vegetation response in a coastal California watershed.
The designation of a coastal marsh as a reserve or park does not protect it
from land use activities outside its boundary. Elkhorn Slough, inland of
Monterey Bay, supports one of the largest coastal marshes in California.
While the slough contains an ecological reserve and a NOAA research
reserve, agriculture in the watershed has increased dramatically since 1970,
especially on steep slopes adjacent to pickleweed (Salicornia virginica)
dominated salt marshes. This land use change has greatly increased sedimentation and formation of alluvial fans that have filled marshes and mudflats. Alluvial fans have historically been a small component of Northern
CA salt marsh ecosystems and supported a highly diverse plant assemblage,
but their now-greater prevalence here can potentially drive plant succession
and community dynamics along the margin of the slough. Aerial photos
were used to conduct an historic post-classification change detection analysis on 15 alluvial fans formed in coastal salt marshes. Photos from 1971,
1980, 1992, and 2001 were classified based on a modified NOAA Coastal
Change Analysis Program land-cover classification system adjusted for
fine-scale imagery. The decadal record of vegetation change indicated an
almost 50% reduction in pickleweed cover within the study areas, and an
encroachment of arroyo willow (Salix lasiolepis) into the marshes along
with an invasion of cattails (Typha spp.) in some sites. Significant differences in elevation, soil texture, salinity, and nitrate concentration between
pickleweed salt marshes and alluvial fan plant communities provide an
explanation for the vegetation shift. Plant surveys indicated that some sites
support high species diversity with composition similar to historic (1926)
accounts of local alluvial fan plant communities, though the present-day
fans are more dominated by non-native species such as poison hemlock
(Conium maculatum). While some fans can provide refuge for plant diversity within Elkhorn Slough marshes, the formation of most fans leads to a
shift in wetland type from salt marsh to monotypic freshwater marsh or
riparian willow stand. This project explains larger-scale effects on wetland
composition and structure and will guide wetland management activities
that occur in highly impacted watersheds.
BYRNE, LOREN B.,* MARY ANN BRUNS and KE CHUNG KIM. Penn
State University, University Park, PA, USA. The effects of lawn management on soil arthropods.
Turfgrass lawns are ubiquitous in urban landscapes, yet they have rarely
been studied by ecologists. The objective of our research was to examine
the effects of lawn management on soil arthropods. Arthropods were sampled using soil cores from pre-existing field locations of high- and lowmaintenance lawn treatments and unmown reference sites in State College,
PA in June 2001 and 2002. High-maintenance lawns received chemical
inputs (pesticides and/or fertilizers) and were mown regularly. Low-maintenance lawns received no chemical applications but were mown while the
reference fields received no human management inputs. Mites were nearly
twice as abundant in high-maintenance lawns compared to the reference
sites and intermediate in number in low-maintenance lawns. Entomobryomorpha collembolans were four and 15 times more numerous in lowmaintenance lawns compared to reference sites and high-maintenance
lawns respectively. Sminthurid collembolans and insects were more abundant in the unmown reference sites. Although few statistically significant
differences were found among the data, results may reflect biologically
important trends. We conclude that lawn management affects soil arthropod

taxa in different ways. Insects and collembolans appear to be negatively

affected by chemical applications but mites do not. However, data indicate
that entomobryomorpha collembolans and mites are not reduced in abundance by mowing. Opportunities exist for developing sustainable lawn
management practices that conserve backyard biodiversity. Lawns may also
serve as an effective focal point for urban ecology research and education.
HNING-GAESE, KATRIN* and NICOLE LEMOINE. Johannes GutenBO
berg Universitat, Mainz, Germany. Influence of global climate change
on species richness of long-distance migratory birds.
Global climate change has been demonstrated to lead to changes in phenology and range shifts of plants and animals. However, little evidence
exists that global climate change causes declines in populations or even
loss of species. Long-distance migratory birds might be a group of species
that especially suffer from global warming. One would expect warmer winters to lead to declines in long-distance migrants if resident birds benefit
from warmer winters and impose increasing competition on migrants. To
study the influence of global climate change on long-distance migrants, we
correlated the number of all species of land birds, and the number and
proportion of long-distance migrants, short-distance migrants, and residents
with climatic factors across 595 grid cells in Europe. We used mean temperature of the coldest month, mean spring temperature, and spring precipitation as measures of climatic conditions in winter and during the breeding
period. The number and proportion of long-distance migrants decreased
with increasing winter temperature, decreasing spring temperature, and increasing spring precipitation. We used this spatial relationship between bird
community structure and climate in Europe to make predictions for changes
in the bird communities in the Lake Constance region, Central Europe,
between two census periods (1980-1981 and 1990-1992). Winter temperature in this region increased significantly between the two censuses,
whereas spring temperature and precipitation did not change. As predicted
from the models the proportion of long-distance migrants decreased and
the number and proportion of short-distance migrants and residents increased between the two censuses. The significant declines of the longdistance migrants in the Lake Constance region are of a magnitude that
can be explained by the observed climate change. Currently, we expand
this approach to other regions in Europe to test whether our model can be
validated more generally.
CABLE, JESSICA M.* and TRAVIS E. HUXMAN. University of Arizona,
Tucson, AZ. Seasonal carbon flux dynamics in Sonoran Desert microbial crusts following precipiation pulses.
Differential responses of ecosystem components to precipitation pulses may
influence seasonal carbon dynamics in arid and semi-arid ecosystems. Microbial crust communities are important components of desert ecosystems
through their roles in nitrogen cycling and soil stabilization. Their carbon
fixation dynamics relative to ecosystem carbon fluxes following precipitation pulses are not well understood. These dynamics are important to understand because crusts may respond to different sized rain events than
vascular plants. To begin to identify crust carbon dynamics following seasonal pulses of water, a one-inch pulse of water was applied to Sonoran
desert crusts in the summer and winter. Diurnal patterns of photosynthesis
and respiration were measured. Summer photosynthetic responses only
compensated carbon loss from respiration, so net carbon gain was negligible. However, winter photosynthetic responses resulted in a net gain of
carbon due to lower respiration rates. This is because there is a single
temperature response of respiration whereas photosynthesis varies seasonally. The seasonal pattern has implications for whole ecosystem carbon
exchange. Following summer precipitation events, microbial crusts do not
show net productivity in the soil community but in the winter, there is a
greater potential for them to contribute to net ecosystem productivity.
CACERES, CARLA E.1,* and ALAN J. TESSIER.2 1 Univerisity of Illinois
at Urbana-Champaign, Urbana, IL; 2 Kellogg Biological Station, Hickory
Corners, MI. To be (dormant) or not to be (dormant): Life history
tradeoffs in Daphnia pulicaria.
It has been suggested that dormancy should be selected against in environments where a population can remain active in the water column year

Abstracts

53

round. We tested this prediction with 12 populations of D. pulicaria. Differences among lakes in basin morphometry, predators and resources create
a gradient of habitat suitability for the Daphnia. Some populations persist
year-round while others are only found in spring. For three years of field
sampling we found consistent differences among the lake populations in
investment in diapause. In some populations, only a small fraction of females switched to producing diapausing eggs each year whereas in others
the majority of the eggs produced in the late spring were dormant. In
general, populations that were only seasonally abundant had a much higher
investment in dormancy than populations that persisted in the water-column
year round, but there were exceptions. We also investigated the annual
hatching fraction of newly produced eggs in five of our 12 populations.
After being incubated at the bottom of the lakes for one year, average
hatching fraction ranged from 6% to 50% across the five populations. Our
results suggest that investment in dormancy varies considerably among
populations, but that seasonal phenology is not necessarily a good predictor
of dormancy investment. Rather, the risks associated with both the active
and dormant stage must be considered.
CADE, BRIAN S.* Quantile regression estimates of animal response
to spatially structured resources.
Estimates of animal responses to their physical environment in spatially
structured landscapes commonly have hidden biases because all factors
limiting the organism are not measured and accounted for in statistical
models. One possible approach to help account for effects of important
unmeasured factors is to include a spatial component in the model, based
on the assumption that unmeasured factors are spatially structured. A simple way to implement spatial structuring in a regression model is by including a spatial trend surface as some low order polynomial function of
latitudinal and longitudinal coordinates of sample locations. Simulations
with quantile regression demonstrate that less biased estimates of effects
of environmental predictors and more variation in animal response were
explained by models that included terms for the spatial trend. Considerable
heterogeneity in responses remain unless the unmeasured factors are strongly correlated (R . 0.9) with spatial trend and, thus, quantile regression is
useful for providing estimates of differing rates of change across the probability distribution. When there are heterogeneous responses in a regression
model there is no longer a single rate of change that characterizes how the
probability distribution is affected by covariates. Some subset of quantiles
[0, 1], typically upper quantiles near the maximum when interference interactions between measured and unmeasured factors predominate, provides less biased estimates. Quantile regression with a spatial trend surface
and physical environmental covariates is used for estimating bivalve mussel
(Macomona liliana) response to spatially structured tidal processes in a
New Zealand harbor, data previously analyzed with least squares regression
(Legendre et al. 1997).
CADENASSO, MARY L.,1,* STEWARD T.A. PICKETT1 and J. MORGAN GROVE.2 1 Institute of Ecosystem Studies, Millbrook, NY; 2 USDA
Forest Service, Northeastern Research Station, South Burlington, VT. Dimensions of ecosystem complexity: heterogeneity, connectivity, and history.
Biocomplexity metaphorically invokes connectivity among organisms and
their environment. Ecology is poised to move beyond metaphor towards a
rigorous definition of biocomplexity and model testing. We define biocomplexity as the degree to which ecological systems comprising biological,
social and physical components incorporate spatially explicit structure, organizational connectivity, and historical contingency through time. These
three dimensions of biocomplexity heterogeneity, connectivity, and history will be explored along axes of increasing complexity. Basing the
description of spatial heterogeneity on patch or continuous quantification,
complexity of spatial structure increases as quantification move from simple discrimination of patch types and the number of each type to assessment
of configuration and the change in the mosaic through time. Organizational
complexity reflects the increasing connectivity of the basic units that control system dynamics. At the simple end of the axis, the functional connectivity between units is low, and the processes within a unit are determined by structures or other processes within that unit. At the highest level
of complexity along this axis units in a mosaic interact through fluxes of

54

Abstracts

energy, matter, organisms, or information, and the structure and dynamics

of the mosaic can be altered by those fluxes. Temporal relationships in the
system exist beyond direct contemporary ones. The influence of legacies,
the existence of lagged effects, and the presence of slowly appearing indirect effects constitute increasing temporal complexity. Simplicity is the
null point on each axis of complexity. A practical goal and motivation for
understanding the dimensions of complexity is to discover the simplest
models and analyses capable of effective explanation.
CADOTTE, MARC W.* and TADASHI FUKAMI. Complex Systems
Group, Ecology and Evolutionary Biology, University of Tennessee, Knoxville, Tennessee. Community dispersal in a hierarchically-structured
landscape and the relationship between local and regional diversity.
Dispersal is thought to increase local (alpha) diversity and decrease amonghabitat (beta) and regional (gamma) diversity because extinction-prone species will maintain higher local abundances, show prolonged persistence
time, and occupy more regional sites due to a rescue effect. However,
dispersal normally occurs at multiple spatial scales in a hierarchically structured system, posing a problem in defining local and regional scales for
understanding diversity. We used freshwater microcosm landscapes each
consisting of three metacommunities each containing three local communities to examine the role of dispersal and its scale dependence. Each local
community was initially stocked with 8 of 14 potentially competing prey
species and 2 of 3 possible predators. We manipulated dispersal into three
treatments: I) dispersal within and among metacommunities, II) within metacommunities only, and III) no dispersal. At the local community scale
we found that alpha diversity was not strongly affected by dispersal. However, beta and gamma diversity were both lowered by dispersal. At the
metacommunity scale the same pattern was apparent, and higher-scale dispersal (i.e., among metacommunities) did not affect patterns of diversity
between landscapes I and II. Dispersal at the local-level appeared to be the
most important factor affecting beta and gamma diversity. Persistence time
decreased significantly with dispersal, and isolated populations were less
prone to extinction. A few species benefitted greatly from dispersal and
colonized all habitats. Overall, we attributed these patterns to the movement
of a single predator which was able to structure local communities to a
very similar state. Our results suggest that local-level processes are important in structural patterns of diversity and that the scale at which dispersal occurs need explicit consideration in understanding its effects on
local and regional diversity.
CALABRESE, JUSTIN M.* and WILLIAM F. FAGAN. Department of
Biology, University of Maryland, College Park, MD. Lost in time, lonely
and single: Reproductive asynchrony as a mechanism of the Allee effect.
Identifying linkages between life history traits and small population processes is essential to the effective conservation of threatened and endangered species. Reproductive asynchrony, which occurs when individuals
are reproductively active for only a portion of the population-level breeding
period, has not yet been analyzed in this manner. Though asynchrony has
generally been considered an advantageous bet-hedging strategy in a temporally unpredictable environment, we explored the dynamic consequences
of reproductive asynchrony as a density-dependent life history trait. We
first compiled empirical data on reproductive timing for a variety of insects
and dioecious annual plants to restrict our mathematical analyses to realistic
levels of asynchrony. We then used a general model of reproductive timing
to quantify the temporal overlap of opposite-sex individuals and to simulate
the dynamics of theoretical populations over a range of initial population
densities and levels of asynchrony. This approach allowed us to examine
quantitatively how asynchrony affects extinction risk. Finally, we considered how protandry, a life history strategy that often accompanies asynchrony, modulates the population-level effects of reproductive asynchrony.
We found that asynchrony reduces a females probability of mating by: 1)
decreasing the number of males a female overlaps with, 2) decreasing the
average probability of mating per male/female pair that do overlap, and 3)
leaving some females completely isolated in time. This loss of reproductive
potential leads to extinction via an Allee effect at low population density.
In all cases, protandry exacerbates the loss of reproductive potential, making populations exhibiting both asynchrony and protandry more susceptible

to extinction. Reproductive asynchrony, acting either alone or in concert

with protandry, should therefore be recognized as a mechanism of the Allee
effect and be included among the suite of life history characters analyzed
when determining a species extinction risk at low population density.
CALCOTE, RANDY and BRYAN SHUMAN. University of Minnesota,
Minneapolis, MN. Climate change, insect outbreaks and the mid-Holocene hemlock decline.
Fossil pollen percentages of eastern hemlock (Tsuga canadensis) decreased
dramatically about 5400 cal yrs BP and did not recover significantly for
1000-2000 years. The traditional interpretation of the decline is that hemlock populations were decimated by insect or pathogen outbreaks. Climatic
change was not considered to be a cause. Several independent lines of
paleoclimatic data now suggest that climatic changes in eastern North
America were associated with the hemlock decline. Lake level studies using
transects of sediment cores across lake basins indicate low lake levels and
dry conditions in New England, Quebec, Ontario, and Michigan from about
5300 to 3200 cal yr BP. Modern analog climate reconstructions, based on
fossil pollen assemblages near hemlocks mid-Holocene western range limit, suggest decreased winter temperatures at the time of the decline, probably in combination with periods of drought. Sites just outside the range
of hemlock, which should not be affected by insect or pathogens attacking
hemlocks, also indicate lower winter temperatures during the period of the
decline. Stable isotope studies at Crooked Pond, Massachusetts, and at
Crawford Lake, Ontario, also demonstrate that the interval of the decline
differed from previous and later intervals. The combination of temperature
and moisture conditions is likely to have contributed to insect outbreaks
and the hemlock decline either by stressing the tree populations and/or by
directly favoring insect/pathogen populations. The mid-Holocene hemlock
decline is the classic paleoecological example of a probable insect outbreak
and its effect on forests, and improvements in our understanding of the
paleo-record suggest that the interactions of multiple factors caused the
decline. The inevitability of future climate changes makes it increasingly
important to understand the combined effects of interactions.
CALEF, MONIKA P.,1,* DAVID L. VERBYLA,1 HOWARD E. EPSTEIN2
and HERMAN H. SHUGART.2 1 University of Alaska, Fairbanks, AK;
2
University of Virginia, Charlottesville, VA. Land cover estimates in
interior Alaska across classifications and resolutions.
Modeling the effects of climate change in Alaska at regional scales requires
vegetation data sets that accurately identify vegetation types across the
landscape. This study was interested in the compatibility and accuracy of
currently existing classifications for Interior Alaska and the effect of scale.
Three 1km-resolution based land cover classifications were compared with
each other and with a 25m-resolution TM image. Total overall agreements
between vegetation classifications range from 59% to 96%. Agreement between the four 1km-resolution classifications and the TM image range from
40% to 52%. Kappa values are very low, corroborating that much of the
agreement between different classifications can be attributed to random
chance. The resolution of the vegetation classification affects the representation of vegetation types: the major vegetation types eliminate the smaller
types with increasing coarseness. The land cover classifications were developed on state- and continental scale and do not accurately represent the
vegetation for a small area in interior Alaska.
CALLAWAY, RAGAN M.,* GILES C. THELEN and WILLIAM HOLBEN. Division of Biological Sciences, University of Montana, Missoula,
Montana, USA. Invasive weeds and soil microbes: negative feedback at
home and positive feedback away.
Microbial soil pathogens can regulate plant populations and escape from
microbial effects may contribute to the remarkable population explosions
of exotics. Soil microbes can also have beneficial effects on plant populations through root-fungus mutualisms and by driving the nutrient cycles
on which plants depend. Positive feedbacks occur when plant species accumulate microbes near their roots that have beneficial effects on the plants
that cultivate them, and are thought to lead to a loss of community diversity. Negative feedbacks occur when plant species accumulate pathogenic
microbes in the soils they occupy which create conditions increasingly

hostile to the plants that cultivate them. Negative feedbacks are thought to
enhance community diversity by increasing species turnover rates. We compared the effects of soil microbes collected from four populations of Centaurea maculosa L. in its native range in western Europe to the effects of
soil microbes collected from five populations in the northwestern U.S.
where C. maculosa has invaded. Sterilization of European soils caused a
107% increase in the total biomass of C. maculosa compared to a 29%
increase when North American soils were sterilized. In addition, strong
feedback loops between soil microbes and C. maculosa were demonstrated.
Centaurea maculosa plants grown alone in non-sterile French soil cultured
by conspecifics were significantly smaller than those grown in French soils
cultured by Festuca ovina. In contrast, C. maculosa planted in Montana
soils cultured by conspecifics were significantly larger than in Montana
soils cultured by F. idahoensis. Similar results were found for C. maculosa
grown in competition with bunchgrasses in cultured French or Montana
soils. Sterilization of the soils eliminated these feedbacks. Considered together, these results suggest that C. maculosa is able to modify the microbial community in invaded soils to its own advantage; but not in its native
soils.
CAMACHO, FRANK A.* and ROBERT W. THACKER. University of
Alabama at Birmingham, Birmingham, AL. Freshwater cyanobacterial
resistance to herbivory: Chemical versus structural defense.
The freshwater cyanobacterium Lyngbya wollei forms dense mats in culturally impacted aquatic ecosystems in Alabama and produces saxitoxin, a
carbamate alkaloid neurotoxin. We explored the palatability of L. wollei
compared to a filamentous green alga, Rhizoclonium hieroglyphicum, in
feeding assays using the freshwater amphipod Hyalella azteca as an herbivore. Three types of feeding assays were conducted: (a) whole mats of
R. hieroglyphicum and L. wollei in the presence and absence of H. azteca,
(b) ground L. wollei and R. hieroglyphicum, and (c) crude L. wollei extract
coated onto artificial food strips. For the whole mat assay, H. azteca exhibited strong feeding preferences for R. hieroglyphicum compared to L.
wollei treatments, significantly reducing R. hieroglyphicum growth rates.
Amphipod density on a mat was negatively correlated with mat growth
rate. Experiments using ground and extracted L. wollei showed that feeding
deterrence was not due solely to the structural effect of the calcium carbonate sheath that surrounds the cyanobacterium. These results suggest that
L. wollei may incur low rates of herbivory in freshwater ecosystems due
to both structural and chemical defenses.
CAMILL, PHIL,1,* JULIAN B. ADAMS,2 JACOB LIMMER,1 SAHIR
KALIM,1 GREG RAFERT,1 ROSALI MIDDLEMAN1 and CHRIS ANDREASSI.1 1 Carleton College, Northfield, MN; 2 University of Virginia,
Charlottesville, VA. Population, community, and ecosystem-level constraints on Picea mariana establishment in boreal peatlands following
permafrost thaw.
Because Picea mariana (black spruce) is often the sole canopy dominant
in boreal permafrost peatlands, it significantly impacts ecosystem processes, including rates of carbon cycling and permafrost formation. Classic
models of cyclic succession suggest that thawed, Sphagnum-dominated collapse-scar bogs are colonized by P. mariana from surrounding permafrost
plateaus, leading to the eventual reformation of permafrost and attendant
shifts in C accumulation. However, little is known about the processes
controlling P. mariana establishment in boreal peatlands. Our goal was to
describe demographic transitions of seed arrival, germination, and survival
as mediated by community and ecosystem-level factors for 18 thawed bog
sites in Manitoba, Canada. We collected 5-6 years of data on seed rain and
seedling germination and survival using a combination of seed traps, artificial seed additions, and seedling transplants (n 5 2160) into three treatments: water table depth (high/med/low), NPK fertilization (+/-), and vascular plant competition (+/-). Results indicate strong limitations on recruitment by population, community, and ecosystem-level processes. Seed dispersal declined with distance from permafrost plateaus, and collapse scars
received an average of 3x fewer seeds than plateaus. At the community
level, germination was severely inhibited (,0.1% after five years) by growing Sphagnum surfaces that overtopped germinants, but it was increased to
10-40% by artificially killing the moss substrate. At the ecosystem level,
seedling survival and germination were strongly controlled by water table

Abstracts

55

depth. Seedling survival after five years was limited to S. angustifolium

lawns (91% survival) and S. fuscum hummocks (96% survival) where mean
water table depth was . 10cm. Almost no seeds germinated in wet S.
riparium communities, where survival of seedling transplants was 5%.
These results suggest that P. mariana establishment is strongly controlled
by water table changes associated with peat accumulation and bryophyte
succession as well as factors that control mortality of the moss substrate.
CANHAM, CHARLES D.1,* and SETH W. BIGELOW.1,2 1 Institute of Ecosystem Studies, Millbrook, NY; 2 U.S. Forest Service - Sierra Nevada Research Center, Davis, CA. Neighborhood dynamics of nutrient redistribution via litterfall: Implications for population dynamics and ecosystem processes.
Our ability to explicitly link models of forest community and ecosystem
dynamics has been hampered by a mismatch in the spatial scales typically
considered by community and ecosystem ecologists. Most current models
of tree population dynamics focus on fine-scale neighborhood interactions,
while ecosystem models typically aggregate at a larger, plot scale. When
local species effects on ecosystem processes are non-additive, there can be
considerable error in using non-spatial models to predict the effects of
changes in species abundance. We present a spatially-explicit analysis of
redistribution of calcium (Ca), magnesium (Mg), and potassium (K) via
litterfall of the dominant tree species in a temperate forest. We used inverse
modeling and maximum likelihood methods to fit models of leaf litterfall
for individual species. In all of the species, a lognormal distribution was
the most parsimonious model of leaf litter distribution around source trees,
and there was a strong wind direction effect in 5 of the 6 species. For two
of the species, Fagus grandifolia and Quercus rubra, the modal dispersal
distances for leaves were 7 and 11 m, respectively, well outside the radius
of the crown. Leaf nutrient chemistry varied significantly as a function of
both species and soil nutrient availability. Concentrations of Ca and Mg in
litter of the 3 tree species that are typically found on more cation-rich soils
increased asymptotically as soil concentration increased, while the species
typically found on more base-poor soils had consistently low concentrations of Ca and Mg in litter. Given evidence of strong tree species effects
on soil nutrient availability in this system, and non-additive effects of mixtures of different species of leaf litter, the results have implications for the
competitive interactions between tree species and the development of
patchiness in species distribution within stands.
CAPLAN, J. S.* and J. A. YEAKLEY. Environmental Science, Portland
State University, Portland, OR, USA. Rubus discolor vigor under varying
photic and edaphic conditions in western Oregon.
Rubus discolor (Himalayan blackberry) is a major plant invader of anthropogenically disturbed habitats in the Pacific Northwest. This study used an
elevation gradient from the Willamette Valley to the western Cascade
Range in Oregon to assess R. discolor vigor in a wide range of photic and
edaphic conditions. We measured two indicators of patch vigor, mean floricane cane length and patch height, at 41 R. discolor patches along the
gradient. We measured canopy cover, slope, aspect, as well as soil particle
size distribution, color, moisture, organic matter, pH, Eh, and NO3+NO2-N.
Linear regression revealed patch height varied most strongly with canopy
cover (R250.37, p,0.01). Cane lengths were longer at sites with the highest organic matter and soil moisture content, but shorter at sites with the
highest Hurst color rating. Patches were taller at sites with the least canopy
cover, slope, sand content, and elevation, but shorter at sites with the highest silt content. We attribute superior vigor in patches with low canopy
coverage to increased light availability at those sites, while we attribute
diminished vigor in sites with low organic matter, silt, or soil moisture, and
high gravel, sand, and Hurst color to limitation from moisture availability
and low soil fertility. The growth response to increased light availability
suggests light is the predominant limiting resource to R. discolor vigor in
anthropogenically disturbed habitats.
CARDELUS, CATHERINE.* University of Florida, Gainesville, FL. Environmental determinants of vascular epiphyte distribution and abundance in a tropical wet forest.
The community composition and distribution of vascular epiphytes is examined in relation to photosynthetic photon flux density, temperature, hu-

56

Abstracts

midity and tree characteristics in the inner-crown of two emergent tree

species, Hyeronima alchorneoides and Lecythis ampla, at La Selva Biological Station, Costa Rica. In the combined sample branch area of 32 m2,
53 species were found. There was little overlap of epiphyte species among
individuals of the same tree species or between tree species. Abundance
patterns of vascular epiphytes were significantly different between the two
tree species however, Hyeronima had significantly greater abundance of
epiphytes than Lecythis, particularly for Pteridophytes. A direct gradient
Canonical Correspondence Analysis of the relationship between epiphyte
species distribution and the microenvironmental and tree characteristics explained only 12.4% of the variance in epiphyte species distributions in the
inner-crowns. Within each tree species alone, however, the microenvironmental and tree characteristics explained at least 20% of the species distributions. High diversity, combined with the lack of community structure,
suggest that tropical rainforest canopy communities are highly dynamic.
High diversity may be maintained by a non-equilibrium mechanism such
as intermediate disturbance or dispersal limitation.
CARDILLE, JEFFREY A.,* MICHAEL T. COE, MONICA G. TURNER,
STEPHEN R. CARPENTER, JONATHAN A. FOLEY and JULIE A.
VANO. University of Wisconsin-Madison, Madison, WI, USA. A generic
lake model of water balance: Impacts of climate variation and lake
type.
Lakes and terrestrial ecosystems interact through flows of water, organic
carbon and nutrients, yet understanding these interactions across spatially
heterogeneous landscapes is incomplete. We studied hydrologic fluxes in
the Northern Highlands Lake District, Wisconsin (USA), a forested landwater mosaic in which 13% of the surface area is covered by lakes. A
simple lake model that represents key fluxes and components of the integrated land-lake system was developed to address the following questions:
(1) What is the relative effect on water balance of lake type and climate?
(2) What lake types are most sensitive to climatic variation (e.g., drier vs.
wetter climate), and what lake types are less sensitive? The model simulates
hydrologic fluxes for mounded, seepage, and drainage lakes by using simple equations of groundwater and surface flows. The three lake types were
parameterized using long-term data for well-studied lakes in the region,
and basic water balance resultsmodeled volume, stream outflow, groundwater outflow, and flow timingwere compared to published data. Water
balance of real lakes under current conditions was well-represented by this
general model and a common set of parameters. Simulation experiments
were conducted for each lake type using climate scenarios that included
current average climate and the wettest and driest months recorded between
1960 and 1990. Simulation results indicate that lake response to climate
variation will differ substantially by lake type. In scenarios of increased or
decreased precipitation, lakes can change type as particular inputs and outputs emerge or disappear, and these changes can strongly affect biogeochemical cycles due to alterations of flow paths and residence times.
Through exploration of lake responses with this model, we can begin to
conceptualize the interactions between terrestrial and aquatic ecosystems
in this complex landscape.
CARDINALE, BRADLEY J.,1 ANTHONY R. IVES1 and PABLO INCHAUSTI.2 1 Department of Zoology, University of Wisconsin, Madison,
WI, USA; 2 Laboratoire dEcologie, Ecole Normale Superieure, Paris,
France. The relationship between biodiversity and ecosystem productivity: Extending our spatial and temporal scales of inference.
Research examining how species diversity influences ecosystem productivity has increased greatly in the past decade as concern about global loss
of biodiversity has intensified. Studies to date have very much improved
our understanding; yet, because most experiments have been performed at
limited spatial and temporal scales, it is unclear whether conclusions can
be readily extrapolated to the scales at which natural communities exist.
We develop a simple patch-dynamics model to examine some of the scale
dependent and independent qualities of the diversity-productivity relationship. We first simulate a typical diversity-productivity experiment and show
that the influence of species richness on productivity is temporally dynamic,
growing stronger through successional time. This holds true whether resource partitioning or a sampling effect is the underlying mechanism of
the diversity-productivity relationship. We then increase the spatial scale

of our experiment from individual patches to a region of many patch types.

The diversity-productivity relationship is not influenced by spatial scale per
se, but the mechanism producing the relationship changes from sampling
effects within individual patches to resource partitioning across patch types
composing the region. This occurs even though model dynamics are the
same at both scales, suggesting that sampling effects and resource partitioning are descriptions of the same process operating concurrently at different scales of observation. Lastly, we incorporate regional processes of
dispersal and disturbance into the model and show that these processes can
amplify the effects of species richness on productivity leading to patterns
not easily anticipated from experiments. We conclude that the relative control of community structure by local verses regional processes may be a
primary determinant of the diversity-productivity relationship. Therefore,
past experiments may not reflect patterns and processes giving rise to diversity-productivity relationships in natural communities where disturbance
and dispersal play a major role in determining the biomass of species.
CARDON, ZOE G.,1 DEBORAH TYSER1 and FRANCIS MOUSSY.2
1
University of Connecticut, Storrs, CT; 2 University of South Florida, Tampa, FL. A new, nondestructive window into dynamics of sugar concentrations around plant roots in the field.
Photosynthetic plants and algae provide the energy and molecular building
blocks necessary to support most life on Earth. In terrestrial ecosystems,
sugars (and other easily utilized organic compounds) "lost" from living
plant roots are important for supporting soil microbial activity. This microbial activity in turn influences decomposition of detritus and thus the
recycling of nutrients to support plant growth and ecosystem productivity.
Currently there is no way to continuously and nondestructively monitor
concentrations of simple organic compounds at fine spatial scales around
roots in field soils. We have developed miniaturized sensors that detect
glucose by relying on the selectivity of immobilized glucose oxidase enzyme to generate a glucose-specific electrical signal at a platinum working
electrode. The sensors, which are approximately the size of very small
thermocouples, can be used in wetland systems to measure glucose around
plant roots in water (or around algae in aquatic or marine systems); we are
continuing development of the sensors for use in soil. Ultimately, we envision sensors of this basic design could be used for spatially resolved,
continuous monitoring of concentrations of a variety of compounds in soil
and/or in water. Such nondestructive, continuous monitoring in situ is essential for advancing our understanding of links between belowground pattern and process in ecosystems.
CARNEY, KAREN M.,* BRENDAN J.M. BOHANNAN and PAMELA
A. MATSON. Stanford University, Stanford, CA, USA. The influence of
plant diversity and land-use on the diversity and function of soil nitrifiers.
Soil microbial communities mediate many critical ecosystem processes.
Little is known, however, about the factors that determine microbial community composition, and whether community composition influences process rates. Using an experimental site at La Selva Biological Station, Costa
Rica, we are examining whether plant diversity and community composition and land-use alter soil microbial communities, and whether microbial
community differences can influence transformations of carbon and nitrogen in soils. Nitrification, the transformation of ammonium to nitrate, is a
critical step in the nitrogen cycle and is performed by relatively few microorganisms, the ammonia oxidizers. We examined the molecular composition and diversity of autotrophic ammonia oxidizers in soils from plots
with varying levels of plant diversity and across land-use types. Ammonia
oxidizers, characterized by PCR amplification, cloning, and sequencing of
16S rDNA with group-specific primers, differed across the different land
use types. Using a laboratory assay that controls for factors other than the
microbial community composition known to influence this process, we also
found that nitrification potential significantly varied across the plant diversity gradient and between land-use types. We tested whether the differences
in ammonia oxidizer community were related to differences in nitrification
rates; preliminary results suggest that differences in nitrification rates may
be more related to changes in the overall abundance of soil microbes than
to their composition.

CARR, DAVID E.* and TAI H. ROULSTON. University of Virginia,

Blandy Experimental Farm, dec5z@virginia.edu, Boyce, VA. Discerning
pollinators and depraved plants: Inbreeding history alters bumblebee
visitation frequencies to Mimulus guttatus.
The majority of plant mating-systems include at least 20% selfing, and
restrictions in gene flow by seeds and pollen make biparental inbreeding
ubiquitous. Inbreeding is typically associated with reduced offspring fitness
(inbreeding depression). Previous work on the yellow monkey flower, Mimulus guttatus, had shown a 30% reduction in pollen production by offspring produced from selfing. Because pollen is an important reward for
pollinators in this species, we examined visitation patterns by a bumblebee,
Bombus impatiens, in a greenhouse population of 960 M. guttatus comprising equal numbers of outbred plants, plants from a single generation
of selfing, and plants from two consecutive generations of selfing. Eight
plants from a particular M. guttatus family and inbreeding level were
grouped together, and 24 of these octets were randomly arranged on each
of five benches. Each morning over a 17-day period we released B. impatiens from a captive hive and allowed them to forage freely on M. guttatus flowers for 75 minutes. Two observers starting at different benches
recorded visitation by following the sequence of visits of focal bees beginning with their arrival at a bench. Observers rotated to a new bench every
15 minutes. Although bees visited similar numbers of flowers when arriving
at each type of octet, bees were 21% more likely to visit outbred octets
than they were octets from a single generation of selfing and were 48%
more likely to visit outbred octets relative to octets from two generations
of selfing.
CARR, MARK H.,1 CRAIG SYMS1 and JENNIFER CASELLE.2 1 University of California, Santa Cruz, Department of Ecology and Evolutionary
Biology, Santa Cruz, CA; 2 University of California, Santa Barbara, Department of Ecology, Evolution and Marine Biology, Santa Barbara, CA.
Large-scale patterns and processes in kelp forest ecosystems.
Kelp forests along the west coast of North America are among the most
productive and species-rich ecosystems in the world. Our understanding of
the environmental and ecological processes that influence the structure and
dynamics of these shallow coastal ecosystems has been constrained by the
restricted spatial and temporal scale of ecological studies. Since 1999, we
have conducted annual surveys to quantify the structure and dynamics of
the macroalgae, macroinvertebrates and fishes that constitute kelp forest
communities along the coast of southern and central California. These
large-scale surveys indicate that different components of the kelp forest
ecosystem exhibit different patterns of spatial structure and dynamics reflecting, in part, a complex layering of responses to geologic and oceanographic processes. Macroalgal abundances differ among years in response
to the interactive effects of wave exposure, substratum type, and grazers.
Despite the high level of variability in abundance of macroalgae, which
are essential sources of habitat and food for many invertebrates and fishes,
the distribution and dynamics of invertebrate and fish assemblages have
been more constant over time. Replenishment of fish populations and their
assemblages largely reflects interannual and shorter temporal scale oceanographic processes that influence the arrival of young and their early postsettlement survival. Such patterns have directed process-oriented studies to
identify, for example, how larval recruitment of fish species respond differently to mesoscale oceanographic processes (e.g., upwelling) and how
predation contributes to dramatic post-settlement attrition. Large scale,
long-term monitoring allows us to examine the relationship between inferences drawn from small-scale process studies (e.g., experiments) to larger
scale patterns.
CARRINGTON, MARY E.* Governors State University, University Park,
IL. Aboveground vs. belowground competition for seedlings in post
fire rosemary scrub, Florida.
Belowground competition is expected to be more intense than aboveground
competition during post fire seedling recruitment in rosemary scrub, an
open, shrub-dominated ecosystem occurring in peninsular Florida on nutrient-poor, sandy soils. To test this prediction, during September 2002 I
established a field study at Archbold Biological Station to compare the
intensity of aboveground vs. belowground competition during recruitment

Abstracts

57

of sand pine (Pinus clausa) seedlings. The study is a 2 x 2 factorial design;

treatment factors are aboveground competition (+ or -) and belowground
competition (+ or -). I established treatment plots in 11 rosemary scrub
patches (balds) within a 268-ha area that burned in February 2001. In each
rosemary patch I established eight treatment plots (two replicates of each
of the four treatments) in a stratified random manner on both sides of a
44-m line transect. To eliminate aboveground competition, I clipped all
vegetation to ground level in a 2-m diameter circular area surrounding the
treatment plot. To eliminate belowground competition, I inserted a PVC
pipe 13 cm in height and 10 cm in diameter into the soil at the treatment
plot so that the top of the pipe was level with the soil surface. For all other
plots (with belowground competition), as a procedural control I inserted a
PVC pipe and immediately removed it. I then placed five sand pine seeds
within the 10-cm diameter area in the center of each treatment plot, and
covered the seeds with a 7 x 7 x 3 cm exclosure made of 1/8-in hardware
cloth to deter seed predators. Seed germination and seedling survival are
checked monthly, and plots were re-clipped in February 2003. Seedling
germination began in October 2002, with the majority of germination occurring in November 2002. Twenty-five seedlings germinated in 23 different plots by March 2003. Number of emerged seedlings did not differ by
treatment (x2 5 2.06, d.f. 5 3, P . 0.5). Although more seedlings died in
plots without aboveground competition (6) than in plots with aboveground
competition (1), the difference is not statistically significant (x2 5 1.33,
d.f. 5 1, P.0.1). Monthly monitoring of seedling survival and growth will
continue for an additional year.
CARSTEN, L D.,* T WATTS, G BOSCO and T A. MARKOW. University
of Arizona, Tucson, AZ. Gene expression patterns underlying host shifts
in Drosophila melanogaster.
The ability of phytophagous insects to switch to new hosts underlies ecological and evolutionary processes such as range expansion, niche differentiation, and speciation. However, the genetic mechanisms underlying the
ability to switch hosts are poorly understood. In this study, we used DNA
microarrays to ask about changes in gene expression patterns in response
to dietary shifts in Drosophila melanogaster, an ecological generalist. Larval diets included standard cornmeal culture medium as a control as well
as several ecologically relevant fruits, vegetables, and necrotic cactus. Dietary shifts were associated with repeatable changes in gene expression.
For example, larvae feeding on cactus exhibited transcriptional changes in
34 genes, 16 of known function, compared to those growing in control
food. We discuss the implications of these functional genomic shifts for
understanding an insects ability to adapt to new food sources.
CARTER, DERB S. Southern Environmental Law Center, Chapel Hill, NC.
Status and trends for forested wetlands in the Southeast.
Approximately one-half of remaining wetlands in the lower 48 states is in
the Southeast. Nearly one-fifth (17%) of Southeast forests is wetland and
approximately three-fourths (74%) of the wetlands in the region are forests.
Functions and values of forested wetlands are well documented and include
filtration or storage of sediments, nutrients or other pollutants; flood or
storm water storage and retention; groundwater recharge and discharge;
and habitat for plants and wildlife. In the highly altered Southeastern Coastal Plain, forested wetland tracts provide most of the large contiguous forested blocks of habitat required by several rare, declining and threatened
or endangered species. Since 1988, federal administrations have established
a national goal of no net loss of wetland acreage and function. Wetland
trend data indicate progress toward the no net loss goal. Loss of forested
wetlands remains high with most of the loss in the Southeast. Intensive
silviculture (drainage and conversion of wetlands to pine plantations) has
replaced agriculture as the leading cause of wetland loss in the Southeast.
The federal Clean Water Act exempts from permitting requirements narrowly defined silvicultural activities that have little or no adverse effect on
wetlands. Federal law prohibits conversion of a wetland to a non-wetland
without a permit. Failure to enforce this requirement of federal law on
intensive silviculture sites that convert wetlands to pine plantations severely
undermines the goal of achieving no net loss of the nations wetlands. A
recent court decision removing federal protection of "isolated" wetlands is
likely to result in significant losses of unique forested wetland types in the
Southeast since most states in the region fail to protect wetlands under state
law.

58

Abstracts

CARTER, GREGORY A.,1,* ALAN K. KNAPP,2 MELINDA D. SMITH,3

GREG HOCH2 and JIM E. ANDERSON.4 1 University of Southern Mississippi, Ocean Springs, MS; 2 Kansas State University, Manhattan, KS;
3
National Center for Ecological Analysis and Synthesis, Santa Barbara,
CA; 4 NASA, Stennis Space Center, MS. Indicators of phytodiversity in
hyperspectral imagery of the Konza Prairie LTER site.
Spectral radiance reflected from 12 watersheds on the Konza Prairie LongTerm Ecological Research site was evaluated to test the hypothesis that its
spatial variability would serve as a generic indicator of plant species diversity. A 224 channel hyperspectral data set gathered in June 2000 by the
Airborne Visible and Infrared Imaging Spectrometer (AVIRIS) provided
complete coverage of the 400 to 2,500 nm range at 10 nm per channel.
The flight altitude of 20 km yielded a ground spatial resolution of 20 m.
The data were radiometrically corrected for atmospheric absorption and
georeferenced to ground coordinates of 94 long-term transects. For each
transect location, a mean spectral reflected radiance was computed from
values within 10-20 pixels. These mean spectra, their standard deviations
and coefficients of variation corresponded with mean vegetation parameters
that were determined previously for each transect and obtained through the
Konza LTER 2000 data archive. Linear regression analysis at each spectral
band indicated no significant relationships between within-transect spatial
variability in reflected radiance and species richness or the Shannon-Weiner
diversity index. Thus, the initial hypothesis was rejected. However, several
band ratios of radiance per se in the 760-950 nm range correlated significantly (p50.01) with species richness (maximum r-squared50.42). Further
research addressing diversity-productivity interactions and temporal variation in landscape spectra will expectedly yield greater efficacy for remote
sensing in the estimation of phytodiversity.
CASE, ANDREA* and TIA-LYNN ASHMAN. Department of Biological
Sciences, University of Pittsburgh, Pittsburgh, PA, USA. Sex-specific response to resource variation alters pollinator-mediated selection via
seed fitness: lessons from experimental populations of gynodioecious
wild strawberry (Fragaria virginiana Rosaceae).
Pollinator preferences determine the reproductive success of individual
plants within populations and shape the evolution of floral design and display. Plasticity of attractive floral traits to abiotic conditions can alter plantpollinator interactions, such that the nature of pollinator-mediated selection
on flowers is context-dependent. We examined how variation in soil resource availability affects pollinator-mediated selection on seed fitness in
experimental populations of gynodioecious wild strawberry. Because pollinators generally prefer large stamen-bearing flowers in dense hermaphrodite-dominated patches, we assessed the effects of resource variation in
two sex ratio contexts: hermaphrodite-dominated (preferred) versus femaledominated. All plants increased allocation to attractive traits (petal size,
total and daily flower production), and received more pollinator visits under
high resource conditions. However, hermaphrodites invested proportionately more into attraction with increased resources, particularly with respect
to daily flower number. A unique consequence of sex-specific patterns of
daily flower production is that the effective (5flowering) sex ratio is significantly different from the actual (5plant) sex ratios in all treatments.
Futhermore, the strength of pollinator preferences were affected by each
sex ratio-by-resource context, such that the functional sex ratio differed
from the effective sex ratio. These patterns of pollinator visitation affected
levels of pollen limitation among treatments, as well as context-dependent
and sex-specific selection on attractive traits through seed fitness. These
data provide insight into the mechanisms by which differential selection
among ecological contexts can arise.
N* and ZENO
N CANO-SANTANA.*
CASTELLANOS-VARGAS, IVA
National Autonomus University of Mexico, Mexico City, Coyoacan, Mexico. Preliminary analysis of some sense organs in the oviposition site
selection of Sphenarium purpurascens.
We analyzed the role of the tarsi, the antennae and the ovipositor of the
grasshopper Sphenarium purpurascens (Orthoptera: Pyrgomorphidae) in
the oviposition site selection related to a gradient of soil particle size and
edaphic humidity gradient. We found that the blockade of the sensorial
organs (F5 3.52, g.l.5 3, 24, P5 0.015) and the date of oviposition (F5

3.24, g.l.5 8, 24, P5 0.012) but not the interaction dates 3 treatment (F5
0.85, g.l.5 24, 451, P5 0.673) have a significant effect on the site selection
related to the particle size gradient. The females with the blocked ovipositor
selected particles with sizes below to 0.1 mm, while the selection made by
females with the blocked legs and antennae did not differ from the selection
of the control females. In both cases, they selected particles sizes between
0.2 to 0.35 mm. We found that the blockade of the sensorial organs (F5
238.64, g.l.5 3, 24, P,, 0.01) and the interaction dates 3 treatment (F5
2.47, g.l.5 24, 484, P5 0.00015), but not the date of oviposition (F5 0.29,
g.l.5 8, 24, P5 0.959) have a significant effect on the site selection based
on the edaphic humidity level. In this case, the selections made by the
females with blocked antennae and ovipositor were soils with humidities
value .60%, whereas the females of control and those of the blocked tarsi
chose humidities below to 30%. We concluded that the antennae and the
ovipositor of S. purpurascens have a central role in the oviposition site
selection based on the humidity level. On the other hand, the ovipositor
also is important on the site selection in a soil texture gradient. Finally, we
demonstrated that the tarsi of the legs are the sensorial organ with less
importance in the selection based on the texture and humidity soil traits.
CATLIN, DANIEL H.1,* and DANIEL K. ROSNEBERG.2 1 Oregon State
University, Corvallis, Oregon; 2 Utah State University, Logan, Utah. Factors affecting dispersal in burrowing owl populations.
We used experimental and observational methods to examine factors affecting within and between season burrowing owl (Athene cunicularia)
breeding dispersal in two contrasting environments: extensive grasslands
and intensive agriculture. We gathered observational data from the banding
of 320 adult owls from 1998-2003, and radio-telemetry data from 62 owls
in 2002, as well as experimental data from 16 pairs of radio-tagged owls
during the 2002 breeding season. We hypothesized that lower reproductive
success, female owls, higher body condition, owls that were found a higher
average distance away from their nests prior to dispersal, lower density of
availability of mates, and later dispersal date would be associated with
longer distance dispersal movements. We compare our results with a geometric model of dispersal, which we use as our null model, where male
owls do not disperse, and female owls disperse only as far as the next
available male. We also examine how divorce rates, in relationship to nesting success and predation, affect burrowing owl movement, comparing how
far pairs and single owls disperse. The results of this study will aid in our
understanding and management of burrowing owl populations and, in general, will give us insight into how animals move through their environments.
CAVALCANTI, GUADALUPE G.* and GRAEME LOCKABY. School of
Forestry and Wildlife Science, Auburn University, Auburn, Alabama, USA.
Sedimentation influences on fine root dynamics and vegetation composition and structure in riparian forests.
One of the most important functions of riparian zones is the ability to
improve water quality by trapping sediment leaving agricultural fields and
other disturbed areas. Many studies have quantified sediment deposition
and identified sources of sediments in riparian ecosystems. However, little
information exists regarding the impacts of sediment deposition from disturbance on belowground processes within these ecosystems. As a result
of sediment deposition, one might expect a decrease in fine root production
due to reductions of soil O2, which will lead to alterations in belowground
net primary productivity. In the long-term, such alterations can compromise
nutrient levels in forest soils resulting in decreased ANPP. Studies have
also indicated that sedimentation promotes decreases in plant community
richness and diversity. This study is conducted at Fort Benning, Georgia
where intense disturbance caused by heavily military traffic has generated
significant sediment movement into riparian forests associated with ephemeral drains. Effects of sediment deposition on fine root production and,
alterations in vegetation composition and structure in riparian forests are
the focus of this study. Since February 2002, nine catchments have been
monitored. Based on vegetation inventory and visual evidence of sediment,
catchments were classified as highly or moderate disturbed and reference.
Within each drainage, samples are collected in paired plots, one in the
upper extremity (disturbed plot), which receives greater amounts of sediments due the proximity of unpaved roads and another (control plot) lo-

cated further down the catchment beyond the reach of deposition. Preliminary results show significant differences in fine root production and ANPP
between paired plots in highly disturbed catchments. No difference was
observed in moderately disturbed and reference areas.
CAYLOR, KELLY K.,1,* HERMAN H. SHUGART2 and IGNACIO RODRIGUEZ-ITURBE.1 1 Princeton University, kcaylor@princeton.edu,
Princeton, NJ, USA; 2 University of Virginia, Charlottesville, VA, USA.
Tree canopy effects on simulated water stress in southern African savannas.
A coupled energy and water balance model is used to simulate the effects
of large tree canopies on soil moisture and water stress across a series of
sites spanning a regional moisture gradient in southern Africa. The model
tracks evapotranspiration from five components of the land surface at each
site the tree canopy, grass under and between tree canopies, and bare soil
under and between tree canopies. The soil moisture dynamics are simulated
at daily time steps and driven by a stochastic model of storm arrivals and
storm depth. Evapotranspiration is modeled using the Priestly-Taylor approach, with potential evapotranspiration scaled by soil moisture availability. The soil moisture under tree canopies is compared to the soil moisture
between tree canopies, and differences in average annual soil moisture
stress conditions are analyzed at each site. The spatial distribution of large
trees has important consequences on the small-scale soil moisture dynamics
across the rainfall gradient. The results indicate that tree canopies serve to
reduce soil moisture stress of under canopy vegetation in the middle of the
rainfall gradient. At the dry end of the rainfall gradient, the effect of tree
canopies on soil moisture is dependent on the amount of yearly rainfall
received.
CHAINE, ALEXIS S.* and BRUCE E. LYON. University of California,
Santa Cruz, Santa Cruz, CA, USA. Sun baked and dry: Consequences
of drought on nesting behavior and sexual selection in lark buntings.
Variable environmental conditions can have severe consequences to the
breeding success of migratory birds. Many species considered to be
ephemeral, may in fact, alter their decision on where to breed based on
conditions when they first arrive at suitable breeding grounds. We studied
the breeding ecology of Lark Buntings, Calamospiza melanocorys, across
five years, which differed from extremes of dry and hot to wet and cold.
We show how moisture and temperature affect nest site selection, nesting
densities, and the success of individual nests in this grassland bird. Furthermore, alteration of the breeding system may have profound consequences on the choices individuals make during a breeding attempt. If
males vary in their contribution of good genes or parental care, female
choice of a mate may differ in years where the challenges of raising offspring are different. We show how sexual selection on two different male
secondary sexual characteristics, body coloration and wing patch size,
varies with environmental and social conditions in different years.
CHALCRAFT, DAVID R.,1,2,* JACK W. WILLIAMS,1 MELINDA D.
SMITH1 and MICHAEL R. WILLIG.2 1 National Center for Ecological
Analysis and Synthesis, Santa Barbara, CA; 2 Texas Tech University, Lubbock, TX. Scale dependence in the relationship between diversity and
productivity: Assessing the role of spatial and temporal turnover.
Recent reviews suggest that the shape of the productivity-richness relationship is dependent on spatial scale, but few empirical studies have examined
scale dependence or its underlying mechanisms. Using data for terrestrial
plant communities at two Long-term Ecological Research (LTER) sites, we
examined the extent to which the relationship between productivity and
richness depended on spatial (Jornada: local 5 1 m2, regional 5 3600 m2;
Konza: local 5 10 m2, regional 5 2000 m2) or temporal (short 5 average
annual richness across yrs; long 5 accumulation of species across yrs)
scale, and the role that spatial and temporal species turnover (i.e., change
in composition in space or time) played in causing scale-dependence. We
found either a weak hump-shaped relationship (Jornada) or no relationship
(Konza) between species richness and productivity at the local scale,
whereas we found significant hump-shaped relationships between species
richness and productivity at the regional scale at both sites. Scale dependence was influenced by spatial species turnover rather than temporal spe-

Abstracts

59

cies turnover. We found a significant hump-shaped relationship between

spatial species turnover and productivity at both LTER sites, but no association between temporal species turnover and productivity. Our results
document the scale-dependent nature of the relationship between species
richness and productivity, and reveals that the way in which productivityrichness relationships change with scale depend on the relation between
species turnover and productivity.
CHAMBERS, CHRIS,1,* JOHN D. MARSHALL1 and BOB DANEHY.2
1
University of Idaho, Moscow, ID; 2 Boise Cascade Corporation, Boise,
ID. Fertilizer uptake by riparian woody vegetation: A 15N tracer experiment.
The nutrient balance of streams and adjacent riparian ecosystems in the
Pacific Northwest has been influenced by the elimination of anadromous
fish runs and perhaps by forest fertilization. To improve our understanding
of nutrient cycling on land, spiraling in the stream channel, and the nutient
linkage between the two ecosystems, we fertilized two streams in central
Idaho. Two levels of nitrogen dose (224kgN/ha and 448kgN/ha) were applied on each stream. The fertilizer mix included a 15N-enriched tracer. This
presentation focuses on fertilizer uptake, expressed as percent nitrogen derived from fertilizer (%Ndff), in the terrestrial woody plant community.
Pre-treatment d15N differed between the two creeks, most likely due to
residual salmon nitrogen on one of the creeks, and this difference was still
detectable following treatment (p , .001). Although the two creeks differed
in pre-treatment d15N, %Ndff did not differ significantly between creeks,
nor was the interaction between creek and treatment significant (p 5 .15
and .13 respectively). The two dominant tree species on both sites, Abies
lasiocarpa and Picea engelmannii, displayed no significant difference in
d15N or %Ndff (difference in means: d15N 5 .39 6 .73 (se), %Ndff 5
3.23% 6 5.9). Most of the fertilizer taken up was allocated to current year
needles in conifers. 224kgN/ha and 448kgN/ha treatments responded similarly in uptake of fertilizer N (%Ndff parameter estimates 5 6.1% 6 6.9,
5.9% 6 5.1 respectively). Alnus incana, the only nitrogen-fixing shrub in
the sample, took up a much smaller proportion of fertilizer than did other
deciduous shrubs (mean %Ndff of shrubs 5 29.0% 6 2.6, A. incana 5
4.57% 6 5.12). Marginal enrichment in conifer litter (mean %Ndff 5 6.2%
6 3.3) was evident, but fertilizer N made up 25% (6 4.2%) of total deciduous shrub litter N. Thus fertilizer N appeared disproportionately in the
shrubs and in the new needles of the streamside conifers. Future work will
trace this label into litterfall and into the streams.
CHAMBERS, JEFFREY Q.,1,2,3,* LIGIA C. TOLEDO,2 ROSEANA P. DA
SILVA,2 EDGARD S. TRIBUZY,2 LILIANE T. MARTINS,2 JOAQUIM
DOS SANTOS,2 NIRO HIGUCHI2 and SUSAN E. TRUMBORE.1 1 University of California, Earth System Science, Irvine, CA, USA; 2 Instituto
Nacional de Pesquisas da Amazonia, Manaus, Amazonas, Brazil; 3 Tulane
University, Ecology and Evolutionary Biology, New Orleans, LA, USA.
Ecological controls over net ecosystem productivity in a Central Amazon forest.
We have carried out extensive field studies of net ecosystem productivity
(NEP) components in a Central Amazon forests including: (i) monthly tree
growth for over three years using dendrometer bands, (ii) precipitation and
litter quality controls over coarse woody debris decomposition and respiration, (iii) precipitation control over respiration from fine surface litter,
(iv) tree growth response to disturbance from a logging experiment, (v)
maximum tree age using radiocarbon dating, and (vi) tree allometry based
on data from hundreds of harvested trees. These data, along with additional
published data, were used to construct an individual-based stochastic-empirical model (coded in Java) that simulates the carbon cycling dynamics
of live trees, dead trees and surface litter. This model was used to explore
a number of ecological controls over aboveground NEP (ANEP). First, in
response to an assumed 25% increase in aboveground net primary productivity linked to the known and expected rise in atmospheric CO2 (fertilization response,) the forest acted as a small net carbon sink (0.2 Mg C ha-1
yr-1) for about 200 years, which is considerably less than previously reported for this forest. Second, forest carbon balance was more sensitive to
changes in ANEP driven by a simulated increase in tree mortality rates,
than a CO2 fertilization response. Third, seasonal variability in ANEP was
more strongly controlled by surface litter (coarse and fine) respiration driv-

60

Abstracts

en by moisture availability, than by seasonal changes in tree growth and

woody tissue production. In addition, tree growth exhibited a strong seasonal cycle that was only partially explained by actual precipitation variables, underscoring the importance of phenology in controlling seasonality
in ANEP.
CHAN, FRANCIS,1,2,* BRIAN GRANTHAM,1,2 KARINA NIELSEN,2
BRUCE A. MENGE1,2 and JANE LUBCHENCO.1,2 1 Oregon State University, Corvallis, OR, USA; 2 PISCO, Partnership for Interdisciplinary
Studies of Coastal Oceans, Corvallis, OR, USA. Sensitivity of marine
ecosystems to climate change: Insights from the novel appearance of
inner-shelf hypoxia in the California Current System.
Detecting the imprint of climatic change on ecosystems and causally linking observed ecological shifts to climatic changes are two fundamental
challenges in ecological research. In marine systems, our emerging understanding of the impacts of climate change have drawn most directly from
studies on temperature-mediated shifts in species abundance and range distributions and alterations of species interactions in benthic communities. In
comparisons, our understanding of the sensitivity and responses of coastal
marine ecosystems to climate-mediated perturbations in oceanic circulation
and biogeochemical cycles remains poorly developed. Here, we report the
novel appearance of an hypoxic (oxygen-depleted) zone in the inner-shelf
off Oregon9s Central Coast in the summer of 2002 and its apparent coupling to discontinuous shifts in regional oceanographic conditions. The bottom water hypoxic zone was extensive (covering ca. 650 km2) and persistent (lasting ca. 2 months), -resulting in widespread die-off and habitat
displacement of marine invertebrates and fishes. The oceanography of the
Oregon shelf is dominated by seasonal wind-driven upwelling of nutrientrich and oxygen-poor water from depth. Upwelled nutrients further fuel
intensive primary production in surface waters that are exported to and
respired at depth. Our results from cross-shelf biogeochemical and physical
transect cruises identify accelerated fluxes of nutrient-rich and oxygen poor
water as the proximate causes of hypoxia development. Analyses of nearand long-term records of oceanographic conditions at the regional and basin
(northeast Pacific Ocean) scales point to the propagation of large-scale
circulation anomalies to local but dramatic physical and biogeochemical
changes in the Central Oregon Coast in 2002. Changes in ocean conditions
may thus be a strong and surprisingly dynamic pathway by which climatic
change can impact coastal marine ecosystems. Our emerging understanding
of the coupling between oceanographic processes and local-scale ecological
dynamics further highlights the sensitivity, and potential non-linear responses of coastal marine ecosystems to climatic perturbations.
CHANDY, SHIBI,* DAVID J. GIBSON and PHILIP A. ROBERTSON.
Southern Illinois University, Carbondale, Illinois, USA. Partitioning of
species diversity and vegetation patterns at different scales in the
Shawnee National Forest.
The USDA Forest Service Research Natural Area (RNA) program has allowed the establishment of permanent study plots in forest types across the
Eastern decidious Forest. Species richness and vegetation data from these
plots provide a baseline for documenting future changes in forest health
and productivity. In southern Illinois, over 390 permanent plots have been
established and sampled in 1996-1998 from 10 RNAs encompassing over
3500 ha. A quantitative partitioning of species richness within and between
landscape units at various scales was conducted. Our results indicate that
the diversity was higher at the landscape level than within RNAs suggesting
the occurrence of heterogeneity at landscape level. We were also concerned
with whether or not vegetation-environment relationships were consistent
at multiple spatial scales. A landscape scale ordination based on tree DBH
from all RNAs indicates a predominant gradient from dry, xeric Quercus
stellata Q marilandica dominated plots to those dominated by mesic species, including Liquidambar styraciflua Platanus occidentalis Acer negungo. Nine of 10 first axis site scores from separate ordinations of each
RNA were correlated (P, 0.05, Spearman9s rank) with the first axis of the
landscape scale ordination. This analysis confirms the consistency of vegetaionenvironment relationship both at local and regional scales.

CHANETON, ENRIQUE J.,* NOEMI C. MAZIA and CLAUDIO M.

GHERSA. Faculty of Agronomy, University of Buenos Aires, Buenos Aires, Buenos Aires, Argentina. Importance of disturbance regime and
community context for exotic tree invasions in mesic grasslands.

We ran a four-year experiment to examine the impact of different disturbance regimes on the invasion dynamics of Gleditsia triacanthos in two
grassland communities of the Inland Pampa in eastern Argentina. G. triacanthos was introduced to the pampas during the mid 1800s, where it has
become a widespread invader of native and agricultural habitats. Our objective was to compare the effects of prescribed burning (infrequent, largescale disturbance) and armadillos activity (frequent, small-scale disturbance) on tree seedling emergence and survival in: (i) a native grassland
relict dominated by perennial bunchgrasses, and (ii) a 20-yr-old successional field dominated by exotic herbs. In each grassland, twenty 4x5 m
plots were arranged in a 2x2 factorial design (n55) of fire and armadillo
disturbances. Burning was applied once at the outset. Three, 0.1-m2 diggings/plot were created every two months to mimick armadillo disturbance
regimes. Every two months, over an 18-mo period, G. triacanthos seeds
were sowed at 40 seed/m2 in each plot to homogenize the propagule pressure. The old-field community was overall more susceptible to tree invasion
than the native grassland. Tree emergence in native grassland was very low
during the dry 1999-2000 season but increased by 4-fold in burned-only
plots during the wetter 2000-2001 season. Burning also favored seedling
survival over the ensuing 3-yr period. Simulated armadillo gaps had a
strong, negative impact on emergence, both in burned and unburned native
grassland, presumably due to increased seed exposure to granivores. In the
old field, armadillo-like disturbances enhanced seedling emergence during
the dry year, but still decreased survivorship in unburned plots, while burning positively affected emergence and survival rates. Thus, the combination
of both disturbance regimes promoted the highest tree invasion rates observed in the old-field community. Our results showed that disturbance
interacted with climatic conditions to determine tree invasion rates. Moreover, large-scale disturbances such as burning would have more predictable
impacts on grassland invasibility, whereas gap-disturbance effects may
strongly depend on the local community context.

CHAPMAN, CAROL* and DAVID G. JENKINS. University of Illinois at

Springfield, Springfield, IL. Metacommunity dynamics drive rockpool
community recovery after a natural disturbance.

Rock pool communities at Discovery Bay Marine Lab (Jamaica) provide

a valuable system of natural microcosms in which metacommunity concepts can be tested in a complex landscape. Following a storm surge that
flushed pools with seawater, we sampled 52 pools (by mixing and subsampling) and 34 outflows (by nets in overflows) to measure metacommunity
dynamics in 15 watersheds, then compared the results to ten years of pool
data. We also sampled vertical distributions in pools at night and midday
to determine if vertical migration affected dispersal. We hypothesized that:
pool communities would be strongly altered by the seawater flush; communities would recover to historical composition over time; and dispersal
within watershed metacommunities would drive subsequent community
structure. Freshwater communities were more severely affected by seawater
flush than brackish or saline pools. Pools with higher organic matter inputs
(mangrove litter or autochthonous inputs) rapidly produced abundant copepod nauplii, which dispersed in rainfall overflows into downstream pools.
Other organisms (e.g., ostracods, harpacticoid and cyclopoid copepods, insect larvae, turbellarians, and nematodes) also dispersed. Crab zoea (Sesarma miersi) were collected in mid- to surface samples at night, but were
not in the water column during days. Crab zoea do not disperse as readily
as other species in overflows, and may be preying on species that disperse
more readily. Therefore, prey dispersal may subsidize predator populations
in some pools, and community composition is strongly affected by dispersal after a disturbance. Rockpool communities are easily observed microcosms that enable the testing of theoretical metacommunity concepts
outside of the lab.

CHAPPELKA, ART H.,,* GREG L. SOMERS1 and HOWARD L. NEUFELD.2 1 School of Forestry and Wildlife Sciences, 108 M. White-Smith,
Auburn University, Auburn, Alabama, USA; 2 Department of Biology, Appalachian State University, Boone, North Carolina, USA. Tropospheric
ozone impacts on large, mature trees in the Great Smoky Mountains
National Park.
Great Smoky Mountains National Park (GRSM), encompassing over
206,000 hectares of area in the states of Tennessee and North Carolina, is
the most visited National Park in the United States. The Park contains a
wide diversity of plants and animals representative of a large region of the
eastern USA. Ozone exposures are high in the Park, and have increased
significantly during the decade of the 1990s, with possible detrimental effects on the vegetation, and in particular, forest trees. Yellow-poplar and
black cherry trees previously cored (1994) and identified regarding ozone
sensitivity were re-cored in 2001 at three sites within the Park. Twenty
trees/species (10 sensitive, 10 non-sensitive)/ site were cored. Some slight
differences existed between the original analysis using cores collected in
1994 compared with those collected in 2001. Based on the 1994 cores,
significant differences at p50.02 were observed for yellow-poplar regarding differences in tolerance. However, using data collected from 2001, significance was observed only at the 88% level. Reasons for these differences
are unknown, but could be due to the following factors: 1) variability due
to small sample size (not all trees measured in 1994 were included in the
2001 analysis for various reasons), 2) differences in analytical procedures
or equipment used between laboratories, and/or 3) a disparity between individuals reading samples. Growth for both species was affected by site,
but no site X sensitivity interactions were observed. Black cherry radial
growth did not differ by ozone sensitivity group during any time period
analyzed. Yellow-poplar varied by sensitivity group during 1990-1994
(ozone-sensitive trees exhibited less radial growth), but no differences in
growth were observed from 1997-2001. Yellow-poplar grew better from
1997-2001 compared with the period 1990-1994, but black cherry grew
less.
CHARRON, ISABELLE* and EDWARD A. JOHNSON. University of
Calgary, Calgary, Alberta, Canada. The scale of tree dynamics on meandering streams.
Sediment erosion and deposition patterns along a stream network are highly
variable due to changes in stream competence, sediment loads, and channel
movement. Likewise, within a deposition site, sedimentation patterns vary
both spatially and temporally due to small-scale changes in topography and
flow resistance forces. However, standard techniques for analyzing tree succession chose plots big enough so that they average out this variability.
Our purpose was to determine the consequences of this variation on trees
along second and third order meandering gravel-bed streams on a small
watershed in the Canadian Rocky Mountains. Our results show that on the
overbank flood zone, trees mainly recruit on certain surfaces that are patchily distributed both in space (at different positions relative to the river) and
in time (after different floods). Moreover, large-scale erosion of these establishment sites, due to channel movement, is rare on these small streams.
Consequently, tree recruitment is highly localized and driven by small-scale
variations in sedimentation. Finally, because of the variable nature of water
movement and sedimentation over the flood zone, the ages of the trees do
not follow a linear increase with distance from the stream. More broadly,
by linking river hydraulics to tree dynamics, the study suggests that typical
chronosequence arguments, which assume this increase in age with distance
and ignore small-scale variations in tree dynamics, lead to false succession
scenarios along meandering streams.
CHASE, JONATHAN M.1,* and TIFFANY M. KNIGHT.2 1 Washington
University, St. Louis, MO, USA; 2 University of Florida, Gainesville, FL.
Drought-induced mosquito outbreaks in wetlands.
Mosquitoes have received considerable scientific and public attention due
to their role as both nuisance and disease vectors to humans. However, the
dynamics of mosquito populations are only rarely considered within the
context of the larger community in which they are imbedded. Although
traditionally, mosquito populations are thought to be limited by abiotic
conditions such as precipitation (which creates some larval habitats), little

Abstracts

61

empirical evidence is available to support this supposition beyond a limited

time scale (weeks). Longer-term data (months or years) almost never show
a positive correlation between precipitation and mosquito abundance. Such
long-term predicative ability is essential for better-informed control measures (e.g., judicious use of pesticides), as well as preparation for higher
mosquito-vectored disease risk. We hypothesized that mosquito outbreaks
depend on the interaction between abiotic factors (habitat availability) and
biotic factors (food web interactions) in wetland habitats, which are breeding grounds for a number of important mosquito species. Wetlands that
retain water throughout the year (permanent) have lots of predators, which
limit mosquito populations. Wetlands that dry predictably every year (temporary) have many species of mosquito competitors, which resist temporal
variation and limit mosquito populations. Wetlands that only dry during
particularly dry years (semi-permanent) will have predators and competitors typical of permanent wetlands during most years, but both groups of
species will be eliminated during a drought, allowing mosquitoes to flourish. We provide support for the hypothesis that mosquitoes should outbreak
following drought years from surveys of the different wetland types, from
a long-term mesocosm experiment that manipulated wetland permanence,
and from microcosm experiments manipulating competitors and predators.
Further, we will discuss implications of this effect for long-term variation
in mosquito populations and the diseases they vector.

rescence, and stomatal porometry, and biochemical determination of the

activity of Rubisco. Light saturated photosynthesis in tall trees was approximately 40% higher than that in dwarf trees. In both cases, photosynthesis light saturated at about 500 umol m-2 s-1. Mean conductances were
similarly greater in tall trees (0.11 vs 0.7 mol m-2 s-1), however, and comparison of photosynthetic conductance responses showed no difference between the populations. Elevation of external CO2 to 1400 umol mol-1 doubled maximal net assimilation, but without completely overcoming stomatal limitations. Nitrogen fertilization of dwarf trees increased stomatal
conductance and mean net assimilation rates without noticeably stimulating
growth; phosphorous fertilization rapidly led to increased growth, based on
increased stem hydraulic conductivity, stomatal conductance and net assimilation. Rubisco activity, in vitro, was similar in controls, and N- and Pfertilized trees, but showed a statistically significant dependence on irradiance at the time of leaf harvest. The results will be discussed with attention both to photosynthesis per se and to the integration of photosynthesis
into an overall strategy of carbon gain and allocation in a harsh, oligotrophic environment.

CHAUDHARI, H. K.* and WILLIAM E. HOPPER. Florida Memorial College, Miami, Florida, USA. Preparation of a virtual natural walk and
tree labeling by a campus ecology club.

Long-term dynamics of boreal forest stands that originated from natural

fires was examined in northeastern Ontario. The objective was to determine
stand compositional development over time since fire. We used repeated
field cruises in combination of sequential aerial photographs that have been
taken from 1946 to 1990s with an interval of 8-15 years. In field measurements, basal areas by tree species were determined by 10 prism stations
along a 200-m cruise line in each stand. On the aerial photographs, a 2-ha
plot, 50 m each side of the 200-m cruise line, was mapped to estimate
stand basal area and composition following the standard forest inventory
method. Time since fire was determined from corrected tree age for stands
older than 80 years and from fire history maps for younger stands. Basal
area of the stands initially dominated by pioneer species such as jack pine
(Pinus banksiana), trembling aspen and balsam popular (Populus spp.), and
white birch (Betula papyrifera) peaked between 60-120 years of TSF and
decreased after, depending on stand composition and soil condition. Basal
area of the stands dominated black spruce (Picea mariana) did not decrease
after stands reached the peak at 100 years of TSF. Eastern white cedar
(Thuja occidentalis) dominance rarely occurred until 100 years of TSF and
its stand basal area increased with TSF. Basal area changes of white spruce
(Picea glauca) and balsam fir (Abies balsamea) dominated stands were
largely coincided with the spruce budworm outbreaks in the region. Of all
sample stands, composition of shade tolerant species increased with TSF
and eastern white cedar dominated oldest stands. Stand cover type changes
were apparent for all stand cover types except white cedar. Multinomial
response models indicated that stand cover type changes were significantly
related to TSF. The models predicted that majority of stands dominated by
pioneer species converged to stand types dominated by late successional
species through time, but divergent and cyclic pathways were also apparent,
which may be explained by the variability of species composition within
each stand cover type and spruce budworm outbreaks.

An undergraduate Ecology Club identified all major plants on the college

campus and made labels to be affixed to all trees. Digital pictures were
taken and web pages were made for each species, and then linked to a
"Virtual Nature Walk" page for use in several classes.
CHEDIACK, ARIADNA.* Bird damage evaluation in citrus plantations
in Tucuman (Argentina) and Guanacaste (Costa Rica).
Citrus growers in Tucuman (Argentina) and Guanacaste (Costa Rica) report
bird damage to citrus crops, while other countries do not consider birds a
problem. My objectives were to: measure and compare bird damage in both
provinces; evaluate economic losses (EL); and correlate bird damage with
3 plantation characteristics (age, structure, and distance to natural forest).
I surveyed bird damage in eight mandarin plantations in Tucuman, and in
four orange plantations in Guanacaste. In Tucuman, blue-fronted amazon
(Amazona aestiva), scaly headed parrot (Pionus maximiliani) and sayaca
tanager (Thraupis sayaca); and in Guanacaste, red-lored parrot (Amazona
autumnalis) cause damage to citrus fruits. Maximum fruit loss was 6.6%
from sayaca tanager and 3.3% from parrots in Tucuman, and 5.4% from
parrots in Guanacaste, while in both provinces 57.8% of loss was from
other causes. EL was 72.96-1162.73 US$/ha/yr for sayaca tanager; 0213.70 US$/ha/yr for parrots, and 88.32-2207.04 US$/ha/yr for other causes. Bird damage was higher in older plantations and varied directly with
distance to forest, but in Tucuman parrot damage was inversely correlated
with distance. Height and canopy area of citrus plants modify bird damage.
Some management recommendations are: harvesting the upper canopy first
before peak damage season, keeping or creating forest near plantations, and
cultivating seedless citrus varieties.
CHEESEMAN, JOHN M.1,* and CATHERINE E. LOVELOCK.2 1 Department of Plant Biology, University of Illinois, Urbana, IL, USA; 2 Smithsonian Environmental Research Center, Edgewater, MD, USA. Photosynthetic characterization of Rhizophora mangroves at Twin Cays, Belize
and responses to fertilization.
Rhizophora mangle (red mangrove) is the dominant tree on the mangrove
cays in Belize. These cays are composed of mangrove peat which has
accreted over the past 10,000 years in association with rising sea level.
Twin Cays is further characterized by a zone of taller trees around the
fringes of the island and creeks, and a zone of dwarf trees in interior areas
less accessible to tidal exchange. Previous studies have shown that dwarf
trees are phosphorous limited, while tall trees are nitrogen limited. As part
of a larger biocomplexity study, we have analyzed the photosynthetic characteristics of red mangrove, using in situ gas exchange, chlorophyll fluo-

62

Abstracts

CHEN, HAN.* Ontario Ministry of Natural Resources, South Porcupine,

Ontario, Canada. Stand dynamics and successional pathways and rates
of northeastern Ontarios boreal forest.

CHEN, JANET* and DANIEL L. POTTS. University of Arizona, Tucson,

AZ. Methane and nitrous oxide fluxes in a semi-arid grassland: The
role of species composition and precipitation pulses.
Methane and nitrous oxide fluxes from terrestrial ecosystems have important implications for biosphere feedback on global change. In aridland ecosystems, fluxes of these trace gases by microbial activity in soils may
depend on plant community composition and soil moisture. However, the
role of plant community composition and soil moisture on soil fluxes of
nitrous oxide and methane have not yet been characterized. Prior to and
following stimulated precipitation events, we compared nitrous oxide and
methane fluxes in plots of either native (Heteropogon contortus) or nonnative (Eragrostis lehmanniana) warm-season bunchgrasses growing on
the Santa Rita Experimental Range near Tucson, AZ. Results show a significant decreasing trend in methane flux for Eragrostis lehmanniana and
an insignificant change in flux for Heteropogon contortus through time
following a pulse. Nitrous oxide fluxes for both plant species did not

change through time during the cool season. However, this is not the case
for all seasons. We conclude that these patterns in methane and nitrous
oxide fluxes in the southwest are affected by season, species composition
and moisture availability.
CHEN, XIN,1,* GUIYE ZHI1 and SHUIJIN HU.2 1 Zhejiang University,
Hangzhou, Zhejiang, China; 2 North Carolina State University, Raleigh,
NC. Arbuscular mycorrhizal colonization and phosphorus acquisition
of plants: Effects of coexisting plant species.
Arbuscular mycorrhizal (AM) fungi influence interactions among plant species through enhancing nutrient uptake and possibly by facilitating nutrient
transport among plants. However, the effects of one species on its coexisting species on AM infection and activity are not well understood. We
examined root AM colonization and phosphorus (P) acquisition of plants
in a highly P-limiting soil near Lanxi city, Zhejiang Province, China from
the year 2000 to 2002. Three dominant native plant species, Kummerowia
striata, Digitaria ciliaris and Ixeris denticulate, were planted at a rate 300
seedlings per plot (2m 3 2m). In the monoculture, K. striata had the highest and D. ciliaris had the lowest AM infection, but plant P content was
higher in I. denticulate and D. ciliaris than in K. striata. In the mixture,
D. ciliaris and I. denticulate did not significantly affect the AM colonization or spore production, but increased the P content of the K. striata plants.
K. striata and I. denticulate significantly increased root AM colonization
of D. ciliaris roots, but had no impact on its P content. K. striata enhanced
but D. ciliaris reduced AMF infection and P content of I. denticulate. These
results suggest that the effects of one plant species on its coexisting species
in mycorrhizal infection and P acquisition may be asymmetrical and cannot
be predicted from the performance of individual species in monoculture.
CHEN, XIONGWEN,* XINSHI ZHANG and BAI-LIAN LI. Department
of Botany & Plant Science, University of California at Riverside, Riverside,
California, USA. Effect of Qinghai-Xizang plateau on the vegetation
distribution in east Asia.
Understanding the present vegetation distribution and its interaction with
climatic regularity and surface irregularity is important for predicting its
future change. In comparison with the rest regions of the world, vegetation
distribution in east Asia is unique; but the underlying causes are still not
clear. Some researches have indicated that the causes were mainly due to
the effect of monsoons; others have suggested that the direct cause was the
rising of Qinghai-Xizhang Plateau. Here we constructed a possible vegetation distribution in this area (15 degree N-60 degree N, 60 degree E-150
degree E) with assumption that there was no Qinghai-Xizang Plateau (e.g.,
its elevation was decreased to 1000 m and roughly equal to its surroundings), using a General Circulation Model and the Holdridge Life Zones
System. Our simulations showed that Qinghai-Xizang Plateau did affect
the vegetation distribution and patterns in East Asia significantly. If the
elevation of Qinghai-Xizang Plateau were about 1000 m, the area of warm
temperate forest and subtropical forest would increase about 96% and 59%,
respectively; cool temperate steppe, alpine meadow and Nival, subnival
and frigid desert would be replaced by warm temperate forest and cool
temperate forest. The development of warm temperate desert in this area
would be largely caused by the rising of Qinghai-Xizang Plateau.
CHENG, YUFU,* WALTER C. OECHEL, STEVE J. HASTINGS, PABLO
BRYANT and YONGHAI QIAN. Global Change Research Group, San
Diego State University, San Diego, CA, USA. Net ecosystem CO2 fluxes
in chaparral when grown under elevated and reduced atmospheric
CO2.
Ecosystem carbon dioxide flux was studied between 1997 and 2001 under
six different CO2 concentrations using CO2 LT (CO2 controlled, naturally
Lit, Temperature controlled) null balance chambers and Free Air CO2 Enrichment (FACE) in southern California chaparral dominated by Adenostoma faciculatum H. & A. For the null balance system, the carbon flux was
measured automatically providing replicated in situ ecosystem level measurements at the 6 different CO2 concentrations from 250 to 750 ppm, with
100 ppm in increment. The annual net ecosystem carbon flux under different CO2 treatments was calculated from daily carbon flux values. There
was a significant CO2 effect (p50.049) and year effect (p50.000) for an-

nual NEE (net ecosystem exchange) from 1997 to 2000. The effects of
elevated CO2 showed a strong seasonal pattern. In the FACE, a Clapp
cuvette was used for measuring the carbon flux at elevated CO2 (treated
by 550 ppm CO2) and ambient on a monthly basis. The effect of elevated
CO2 also showed different patterns in the dry and wet seasons. During the
growing season (April-July), the ecosystem under FACE treatment showed
less efflux than its control (P50.033), while during the non-growing season, it showed higher source (P50.041).
CHESSON, PETER* and BRETT A. MELBOURNE. Unversity of California, Davis, Davis, California. Reversal of the effects of competitioncolonization tradeoffs with environmentally-determined resource heterogeneity.
Competition-colonization tradeoffs have been proposed as a powerful
mechanism of species coexistence based on models in which space is heterogeneous with respect to the relative abundances of different species, but
not with respect to physical environmental factors, except where they are
the consequence of heterogeneous species abundances. However, a model
of explicit resource competition with fixed spatial patterns of heterogeneity
in resource supply suggests that limited dispersal favors the persistence of
an inferior competitor. Moreover, the inferior competitor can exclude a
superior species that disperses too much, with coexistence occurring in the
presence of more moderate species differences. If colonizing ability is interpreted as dispersal ability, these results are opposite to the traditional
predictions. More generally, colonizing ability involves fecundity, but higher fecundity is consonant with higher competitive ability in this model and
does not restore the traditional predictions of competition-colonization
tradeoffs. These results suggest that the predictions of competition-colonization tradeoffs need to be reconsidered with the more realistic assumptions about competition and the structure of the physical environment.
These results also suggest that fecund widely dispersive invasives may fail
to displace native species that have the good sense to remain in places that
are permanently favorable.
CHIANG, JYH-MIN,1,* KIM J. BROWN,1 LOUIS R. IVERSON,3 ANANTHA M. PRASAD3 and JIQUAN CHEN.4 1 Ohio University, Athens,
OH, USA; 3 USDA Forest Service, Northeastern Research Station; 4 The
University of Toledo. Assessing carbon stocks and productivity in current and predicted forest communities using the SHIFT and PnET-II
simulation models.
Climate is the major driving force that influences the distribution of terrestrial vegetation through time. Based on current and future climatic
change scenarios, potential changes in the next 100 years in tree species
composition were predicted using DISTRIB and SHIFT models (Iverson
and Prasad, 2001). To estimate the subsequent impacts on carbon sequestration, a physiologically based, generalized forest carbon and water balance model (PnET-II) was utilized to estimate net primary production
(NPP) and net ecosystem production (NEP) in the current and projected
species assemblages at four locations which represent "hotspots" of potential species shift in the Eastern United States. Our main questions for this
study are: What are the current NPP and NEP rates given the tree species
compositions at the four sites? Based on future shifts in tree species compositions and climate, how will productivity and carbon stocks change?
Prior and current studies at the four research sites (Howland Experimental
Forest, ME; Vinton Furnace Experimental Forest, OH; Ozark-St. Francis
National Forest, AR; and Chequamegon National Forest, WI) provide data
for the PnET-II model parameterization. Research was conducted to obtain
additional forest structure and physiological data for southeastern Ohio,
and will be presented. On the basis of the PnET-II model and data availability, current and potential future trends in NPP, NEP, and carbon storage
will be discussed. The significance of this research is the investigation of
potential climate change effects on carbon sequestration in forests in a
manner that includes not only climate changes but also potential shifts in
species compositions.
CHIARIELLO, NONA R.1,* and CHRISTOPHER B. FIELD.2 1 Stanford
University, Stanford, CA; 2 Carnegie Institution of Washington, Stanford,
CA. Seasonal response of California grassland NDVI to global change
treatments.
The Jasper Ridge Global Change Experiment studies the response of California grassland to four environmental factors changing globallywarm-

Abstracts

63

ing, nitrogen deposition, elevated carbon dioxide, and added precipitationapplied in a full-factorial design. Here we report the response of a
vegetation "greenness" index, the normalized difference vegetation index
(NDVI), to these treatments during four growing seasons. Treatments varied in the seasonal timing of their effects on NDVI and in the magnitude
of their effects in different years. Warming primarily shifted the timing of
NDVI, while nitrogen addition increased the maximum values. Elevated
carbon dioxide depressed peak NDVI but sometimes prolonged canopy
greenness as the season tapered. Watering effects varied among years.
Treatments that increased NDVI early in the season tended to also increase
biodiversity, while treatments that increased or decreased NDVI during the
peak growing season tended to have corresponding effects on annual
aboveground production.
CHIAVELLI, DEBORAH A.,1,2,* KATHRYN L. COTTINGHAM,2 CAROL S. RINGELBERG1 and RONALD K. TAYLOR.1 1 Dartmouth Medical
School, Department of Microbiology and Immunology, Hanover, NH,
USA; 2 Dartmouth College, Department of Biological Sciences, Hanover,
NH, USA. Genomic response of Vibrio cholerae to changes in aquatic
nutrient levels.
Vibrio cholerae is both the causative agent of the emerging infectious disease, cholera, and a common planktonic bacterium in aquatic ecosystems
all over the world. With a fully sequenced and well-studied genome, microarray technology can be used to study changes in V. cholerae gene
expression in varying environmental conditions, and to link these responses
to both bacterial dynamics in aquatic ecosystems and cholera epidemiology
in human populations that utilize these ecosystems. Seasonal outbreaks of
cholera in some tropical regions appear to coincide with plankton blooms,
and V. cholerae is often found attached to zooplankton and phytoplankton
surfaces, a behavior believed to enhance survival, growth, and transmission
to human hosts. Nutrient levels and the availability of attachment surfaces
are also generally important to the dynamics and productivity of pelagic
heterotrophic bacteria. We quantified the effect of nutrient conditions (three
levels of C, N, and P spanning a typical annual range in natural aquatic
habitats) on gene expression in V. cholerae at two time points after introduction into freshwater medium. Expression of many V. cholerae genes
responded to nutrient conditions, including genes regulating nutrient uptake
and metabolism, protein synthesis, cell growth and stress responses, environmental sensor systems, and flagellar structure and function. The same
behaviors (motility changes, chemotaxis, attachment), and in some cases
the same genes, are necessary for the initiation of cholera pathogenicity in
the human intestine and for attachment to planktonic substrates. These processes are hypothesized to respond to poor environmental conditions (e.g.
low nutrients) in a way that increases contact with, and attachment to,
surfaces. However, we did not see nutrient-induced differences in expression of known pathogenicity or attachment genes. This indicated that signals from a substrate may also be needed. We are currently investigating
V. cholerae gene response to the presence of zooplankton to test this hypothesis.
CHICK, JOHN H.,1,* MARK A. PEGG2 and TODD M. KOEL.3 1 Illinois
Natural History Survey, Great Rivers Field Station, 8450 Montclair Ave,
Brighton, Illinois, USA; 2 Illinois Natural History Survey, Illinois River
Biological Station, 704 North Schrader Ave, Havana, Illinois, USA; 3 National Park Service, Yellowstone National Park, Center for Resources, Yellowstone National Park, Wyoming, USA. Long term monitoring design
and spatial variation of fish communities in the upper Mississippi River
basin.
Since 1991, researchers with the Long Term Resource Monitoring Program
have collected fish abundance and composition data from six regional trend
areas (RTA) in the Upper Mississippi River System (UMRS): Mississippi
River navigation pools 4, 8, 13, and 26, the La Grange pool of the Illinois
River, and an open river reach on the Mississippi River near Cape Girardeau, MO. These six RTA were chosen to represent the range of conditions
present throughout the UMRS and are meant to allow for system wide
inferences about fish population and community dynamics. To test the adequacy of these six RTA in representing the variation of fish communities
present in the UMRS, additional sampling was conducted during 2000 from
areas above and below (out-pools) three RTA, and from navigation pools

64

Abstracts

19 and 20. Cluster analysis and non-metric multi-dimensional scaling of

species composition (presence absence) and community structure (catch per
unit effort) data revealed two major spatial groups of pools, and four subgroups. Northern and southern pools formed the two major pool groups.
The sub-groups identified included open river reaches, southern pools,
northern pools, and the La Grange reach of the Illinois River. Out pools
generally grouped with the closest RTA, but pools 19 and 20 grouped with
southern pools in terms of species composition, and with northern pools in
terms of community structure. Similarity among pools was correlated
strongly with distance between pools, suggesting that zoogeography, regional disturbance patterns, and demographic processes (immigration/emigration, source sink dynamics, etc.) have important influences on fish communities. Correlations between community structure and composition with
various habitat measures were not as strong as correlations with distance
between pools. This analysis suggests the six RTA design of the Long Term
Resource Monitoring Program may be adequate to capture the major spatial
differences in fish communities within the UMRS. Nevertheless, additional
research is needed to clarify how fish communities in pools 19 and 20
contrast with other areas in the UMRS.
CHOCZYNSKA, JOANNA* and EDWARD A. JOHNSON. University of
Calgary, Calgary, Alberta, Canada. Settlement and commercial forestry
landscapes in the boreal forest of Northern Ontario.
The two dominant human disturbances in the boreal forest - settlement and
commercial forestry - produce vastly different spatial patterns. We compared polygon characteristics of a landscape with a history of agricultural
settlement and land abandonment to a landscape that has been commercially logged. Settlement commenced in 1913 and forestry in 1923. A polygon was defined as an area of uniform land use and forest age. We found
that the settlement landscape has smaller polygons that correspond to
homestead requirements, while the forestry landscape has larger polygons
resulting from increased harvesting efficiency with an increase in clearcut
size. Low perimeter-to-area ratios were typical of small agricultural polygons in the settlement landscape and large polygons in the forestry landscape. This reflected simple shapes due to the grid survey system and
clearcutting respectively. Perimeter-to-area ratios of forested settlement
polygons were high, but decreased in years of land abandonment as agricultural land of simple shapes reverted back to forest. The age distribution
of forested polygons is consistent with land clearing and abandonment in
the settlement landscape, and product demand, labour availability, and technological advances in the forestry landscape. We conclude that human activity is evident in the resultant landscape patterns, and land clearing must
be regulated if a particular pattern (e.g. one that imitates natural disturbances) is desired.
CHOJNACKY, DAVID C., ROBERT A. MICKLER and LINDA S.
HEATH. Biomass of down woody materials in eastern U. S. forests.
Available and accessible data on components of down woody materials
(DWMs) are needed for managing global carbon storage of forests. The U.
S. Department of Agriculture Forest Inventory Analysis (FIA) program
currently measures variables related to DWMs on a subsample of plots in
several states. We tested the feasibility of linking these data to the entire
FIA database. We compiled and computed biomass on coarse woody material, fine woody material, litter, duff, and shrub/herb cover; and we developed regression models to predict DWM components for extension to
almost 100,000 FIA plots. Average biomass values for the eastern U.S. are
6.2, 4.7, 9.0, 8.8, and 2.2 Mg/ha for coarse woody material, fine woody
material, litter, duff, and shrub/herb cover, respectively. The total of all
DWM components (30.9 Mg/ha) is about 20% of the total forest biomass,
which also includes live (99.4 Mg/ha) and dead (11.3 Mg/ha) standing
trees.
CHOO, JOHANNA* and EDMUND W. STILES. Rutgers, The State University of New Jersey, New Brunswick, New Jersey. A comparison of
avian frugivory and fruit ecology between a paleotropical and a neotropical site.
We present a comparison of interactions among frugivorous birds and
fleshy-fruited plants in two lowland rainforest sites with comparable cli-

matic conditions. One site was located in the paleotropics of Sarawak

(Mulu National Park), and the other in the neotropics of the Peruvian Amazon (Paucarillo Reserve). At each site, JC collected data for one year using
systematic and comparable techniques. The comparisons we make are important because earlier pantropical comparisons of frugivory have been
based on literature review. Conclusions drawn from analysis of literature
alone may be confounded due differences in aims and methodologies of
the studies. Our comparative study addressed this problem and allowed us
to draw more rigorous conclusions based on quantitative and qualitative
flora and avifaunal datasets. We assessed both understory and canopy avifauna frugivory, monitored phenology and estimated fruit crop sizes. Fruiting plants were identified and ripe fruit samples were collected. We found
that Mulu had 40% fewer fruiting Families than Paucarillo and both locations had .20 fruiting Familes in common. In both locations, Moraceae
was an important fruiting Family with many species, but Ficus was the
only genus for Sarawakian Moraceae. Generally, fruit diversity was greater
in Paucarillo, and we examined whether diversity was correlated with fruit
production. Total avifaunal diversity was higher in Paucarillo (34% more
species than Mulu), but both sites had similar diversity of frugivore species.
Our study of both canopy and understory birds simultaneously showed that
some species of frugivores foraged across the vertical forest strata, suggesting that their classification as understory or canopy frugivores could
be misleading. In conclusion, we present the first quantitative and qualitative pantropical comparison of tropical fruit ecology and avian frugivory.
Our findings would confirm and challenge some earlier conclusions drawn
from comparative literature review, and will further our understanding of
tropical bird-fruit interactions.
CHRISTENSEN, NORMAN L.* Duke University, normc@duke.edu, Durham, NC, USA. The southeastern landscape: a history of human use
and ecological change.
The activities of humans over the past 10 millennia have done much to
shape southeastern ecosystems and landscapes. Native American impacts
on the fauna, their use of fire, and their varied agricultural practices had
much to do with the character of the wetlands, savannas and forests encountered by early European explorers. The character and scale of the conversion of forested land to agriculture by European settlers varied considerably from the seventeenth the mid-nineteenth century. Subsistence fallow-cycle farming practiced by early settlers gave way to larger scale activities dependent on regional and international markets. As erosion and
loss of nutrient capital diminished crop productivity, land owners responded
by putting more land into production, thereby increasing their dependency
on slave labor and initiating region-wide cycle of land impoverishment.
Forests not cleared for agriculture during this period were high-graded for
timber and fuel, and impacted severely by livestock. In many parts of the
coastal plain forests were managed for naval stores. Economic pressures
forced the abandonment of much of this land, beginning with the post-Civil
War Reconstruction and continuing through the Great Depression. The successional patterns and widespread reforestation that ensued were heavily
influenced by previous patterns of land use and abuse, as well as the spatial
distribution of activities on the landscapes, thus creating the so-called
southern pinery, one of the most important sources of softwood fiber in
the world. The extent of forested land in the Southeast grew more slowly
during the middle of the twentieth century, and in the 1980s actually began
to shrink with the expansion of urban centers. Where forests remain, forest
management has produced patterns of forest change quite different from
those observed in abandoned fields. Throughout this history, human use of
the land was influenced heavily by the character of the available ecosystems; those uses in turn set in motion patterns of ecological change that
greatly affected the character of ecosystem resources available to subsequent generations. Future southeastern landscapes will just as surely be
shaped by this interaction.
CHUNG, HAEGEUN* and DONALD R. ZAK. University of Michigan,
Ann Arbor, Michigan, U.S.A. Elevated atmospheric CO2 and O3 alter
soil fungal community composition and function.
Atmospheric CO2 and O3 are rapidly rising due to human activity, and this
change in atmospheric chemistry has the potential to alter the rate of carbon
cycling in forest ecosystems. Changes in plant production and litter chem-

istry under elevated CO2 and O3 may alter microbial community composition and function because soil microorganisms are dependent on plant
litter as a source of energy for heterotrophic metabolism. Because plant
litter production is enhanced under elevated CO2 and suppressed under
elevated O3, we hypothesized that microbial metabolism will increase under
elevated CO2 and decrease under elevated O3; we also expected concomitant changes in microbial community composition. We tested our hypothesis at the Free-Air CO2 and O3 Enrichment (FACE) site in Rhinelander,
Wisconsin, where Populus tremuloides, Betula papyrifera and Acer saccharum were grown under ambient and elevated CO2 and O3. We used
extracellular enzyme analysis to measure microbial metabolism and polymerase chain reaction-denaturing gradient gel electrophoresis (PCRDGGE) to determine microbial community composition. The activity of
hydrolytic enzymes that decompose cellulose and hemicellulose increased
under elevated CO2 and decreased under elevated O3. These enzymes are
produced mainly by fungi, and we observed clear shifts in fungal community composition under elevated CO2 and O3. We concluded that elevated CO2 increases fungal metabolism and O3 suppresses it, and this may
be due to change in fungal community composition.
CIPOLLINI, DON* and STEPHANIE ENRIGHT. Wright State University,
Dayton, OH. Induction of systemic acquired resistance in Arabidopsis
thaliana benefits Spodoptera exigua but not Trichoplusia ni.
The role of salicylate in plant responses to pathogens has been well documented, but the effects of salicylate on plant responses to insects are not
well understood. We examined the effects of salicylate on performance of
the generalist herbivores, Trichoplusia ni and Spodoptera exigua, using
Arabidopsis thaliana genotypes that varied in their ability to mount salicylate- and jasmonate-mediated responses. Feeding by neonates of T. ni was
highest on the salicylate deficient nahG, intermediate on the salicylate insensitive nim1-1, and lowest on the jasmonate response mutant jar1-1, the
jasmonate deficient mutant fad, and the Columbia and Wassilewskija wildtypes. Growth of T. ni larvae largely followed that pattern. Genotypes varied in defense protein expression, and only peroxidase activity responded
consistently to salicylate treatment. T. ni larvae responded positively to the
lack of endogenous salicylate signaling in their host plant, but salicylate
treatment had no effect on their performance. In a separate experiment,
growth of Spodoptera exigua larvae was highest on the Wassilewskija wildtype, intermediate on the Columbia wild-type and the fad, jar1-1, nahG
genotypes, and lowest on the nim1-1 mutant. Despite having no discernable
effect on defense protein expression, salicylate treatment increased the
growth of S. exigua larvae on all genotypes except for nahG, which converts salicylate to catechol. In this study, two generalist insects responded
differently to genetic defects in salicylate and jasmonate signaling of their
host plants, and only S. exigua was responsive to exogenous salicylate
treatment. This illustrates that potential cross-effects of salicylate on insect
resistance can occur, but are likely to vary with insect species identity.
Moreover, it illustrates that elements of endogenous salicylate and jasmonate signaling both function in resistance of A. thaliana to these insects,
perhaps through their control of plant defensive chemistry or other aspects
of the suitability of foliage for insect feeding and growth.
CIPOLLINI, MARTIN L.,* ERIC PAULK, KIM MINK, TIFFANNY FISCHER and KAREN VAUGHN. Effects of nitrogen supplementation
on growth, reproductions, and secondary chemistry of Solanum carolinense L.
The effect of variation in nutrient and water availability on growth, reproduction, and fruit chemistry of Solanum carolinense L. was examined by
focusing on three questions: 1) How does variation in N availability affect
growth, reproductive output, and fruit pulp nutrient and secondary metabolite chemistry? 2) Is N supplementation correlated with increases in Ncontaining metabolites and decreases in C-based compounds, as predicted
by the non-adaptive Carbon-Nutrient Balance hypothesis? 3) Do correlations exist between leaf and fruit chemical content, as suggested by the
non-adaptive Physiological Constraints hypothesis? To address these questions, clones of 10 maternal plants were grown for three years (1998-2000)
under two N treatments and two water treatments. At the end of each year,
aboveground plant mass, fruit number, and fruit mass was determined. For
each plant in 1999, we assayed ripe fruit pulp and leaf material for protein,

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65

sugar, phenolic, and glycoalkaloid content. We also analyzed a subset of

ripe fruits for total glycoalkaloid content in 1998. While path analysis revealed strong direct and indirect effects of N treatment on growth and
reproduction, fruit morphological and chemical variables were not strongly
affected by N treatment. Fruit pulp chemistry remained rather constant
despite the large differences in growth and reproduction that were associated with N variation. Leaf and ripe fruit chemical contents were not significantly correlated across treatments and maternal plants, and variation in
leaf chemistry did not help explain variation in fruit chemistry in ANCOVA analyses, suggesting no physiological constraints of leaf chemistry
on ripe fruit chemistry. Our results suggest that, while maternal plants may
vary somewhat in fruit chemistry, and fruit chemistry may vary somewhat
depending upon environmental conditions, levels of primary and secondary
metabolites within fruits are not best explained by the non-adaptive hypotheses. Ripe fruit chemistry remained relatively constant in the face of
drastically changing resource levels, plant growth response, and reproductive output, suggesting an adaptive function, and thus supporting the Optimal Defense hypothesis. Ripe fruit chemistry most likely reflects a balance between selection for attraction and defense.
CIRTAIN, MARGARET C.,* SCOTT B. FRANKLIN and S REZA PEZESHKI. Department of Biology, University of Memphis, Memphis, TN,
USA. Nitrogen and moisture affect Arundinaria gigantea seedling
growth.
A greater than 98% decline in Arundinaria gigantea(Walt.) Walt. Ex Muhl.
(giant cane or river cane) canebrake communities has resulted in a critically
endangered ecosystem. Historical accounts suggest loss of canebrake habitat has resulted in the extirpation (and perhaps extinction) of many species.
Thus, canebrake restoration is necessary for maintaining and enhancing
biodiversity in the southeastern United States. The purpose of this study
was to facilitate reestablishing A. gigantea canebrakes by optimizing
growth and survivorship conditions for seedlings. To this end, we developed a lab experiment to test the effects of soil moisture (periodic drought,
periodic flooding, and amply-watered, well-drained) and nitrogen on A.
gigantea seedling growth. Net photosynthesis, stomatal conductance, and
shoot and root biomass were measured to determine optimal growth conditions. A. gigantea, while able to survive both drought and flooding, had
improved growth under well-watered, well-drained conditions. Shoot
growth averaged 25.85 cm for well-drained, 22.72 cm for flooded, and
18.25 cm for drought treatments. Nitrogen application improved growth
under well-drained moisture regimes; shoot growth was 29.25 cm and 22.2
cm for nitrogen and control treatments, respectively. Transpiration was unaffected by treatments, but both stomatal conductance and net photosynthesis were affected by moisture regime. Well-drained plants had greater
stomatal conductance (mean 5 4.1 mmol m2 sec1) compared to drought
(mean 5 2.84 mmol m2 sec1) and flooded treatments (mean 5 2.82 mmol
m2 sec1). Well-drained plants also had lower net photosynthesis (mean 5
25.86 mmol m2 sec1) compared to drought (mean 5 26.38 mmol m2 sec1)
and flooded (mean 5 26.27 mmol m2 sec1) treatments. The biomass and
physiological results suggest canebrake restoration will be most successful
on well-drained fields with an ample water supply and fertilization.
CLANTON, KEITH B.* and I. J. STOUT.* Department of Biological Sciences, University of Central Florida, Orlando, Florida. Impact of habitat
structure on cotton rat population dynamics in an abandoned agricultrural field.
Inter- and intragrid variation in population dynamics of cotton rats were
studied in an abandoned agricultural field near Lake Apopka, Florida. Trapping took place from March through August 2000 within the St. Johns
River Water Management Districts 3,238-hectare North Shore Restoration
Area. Horizontal and vertical measures of habitat structure were taken. A
significant positive relationship was discovered across the study site between the rate of population growth and the spatial coverage of live herbaceous species. Significant trends between population growth and herbaceous height were not as apparent. Intragrid analyses revealed that larger
males were captured in areas with greater herbaceous height whereas smaller males were taken in areas with shorter herbaceous cover. This relationship was also influenced by the minimum height of the herbaceous layer
and the minimum patch size.

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Abstracts

CLARK, DEBORAH A.,1,* DAVID B. CLARK1 and STEVEN F. OBERBAUER.2 1 University of Missouri-St. Louis, St. Louis, MO, USA; 2 Florida
International University, Miami, FL, USA. Interannual variation in tropical rain forest carbon cycling: a long-term, intensive study (La Selva,
Costa Rica).
The responses of tropical rain forests to on-going climate change have large
implications for the rate of accumulation of atmospheric carbon dioxide.
Because there have been few long-term studies of carbon cycling in these
forests, there is limited understanding of how the net carbon balance of
these ecosystems might respond to changing climate. In an on-going team
effort (the CARBONO Project), researchers have been collaborating since
1996 to investigate the forest carbon cycle in old-growth tropical wet forest
at La Selva. We synthesize here the major findings from the first five years
of research in the CARBONO plots, a set of 18 0.5-ha plots that were sited
using the La Selva GIS so as to sample the landscapes large edaphic
gradients (slopes to alluvial terraces, 3-fold variation in soil P, K, Al) with
an unbiased, replicated design. The on-going measurements of carbon
stocks and fluxes in these plots began in September 1997. They include:
fine litterfall (biweekly), fine woody litterfall (monthly), initial stocks and
yearly inputs of coarse woody debris, aboveground biomass increment
(yearly), soil respiration (biweekly), fine root stocks (3-4 times per year),
tree dynamics and species composition (yearly), and canopy structure (annual gap-mapping). The first five years data revealed substantial interannual variation in carbon cycling in this forest, principally with respect to
CARBONO Year 1, which overlapped the 1997/8 mega-Nino event. In that
record-hot year, estimated aboveground biomass increment was 39% lower
than in the following cooler years. In Year 1 we also measured enhanced
leaf litterfall (but reductions in the other fine litter components), a large
spike in dead fine roots, greatly enhanced tree mortality (returning to baseline mortality rates by Year 4), and large inputs of coarse woody debris.
Our findings demonstrate that carbon cycling in this forest responds strongly to current-time climatic variation.
CLARK, JAMES S.,1,* MIKE DIETZE,1 SATHISH GOVINDARAJAN2
and PANKAJ K. AGARWAL.2 1 Duke University, Center on Global
Change & Program in Ecology, Durham, NC, USA; 2 Duke University,
Center on Global Change & Computer Sciences, Durham, NC, USA. Stabilizing biodiversity with random individual effects.
Ecological theory and analysis suggest that potentially high diversity of
tree species depends on tradeoffs. Colonization-competition tradeoffs involve the capacity to colonize new sites vs the ability to hold them (competition). Different successional niches involve tradeoffs between species
capable of exploiting high resource availability following disturbance vs.
others that can survive long after disturbances, when resources are scarce.
Stochasticity can promote coexistence, provided that species differ in their
colonization success, because long life span can buffer population growth
against variable recruitment (the storage effect). Tradeoffs are still implied, because stochastic recruitment only promotes diversity if correlation
among species, in terms of recruitment success, is not too large. In short,
to explain high diversity, models typically require precise parameter combinations that imply tradeoffs. In most models, species lacking these tradeoffs are rapidly excluded. We demonstrate that tradeoffs are less critical
for coexistence than is often thought, because variability within populations
can overwhelm species differences, and because this individual variation
can have a stabilizing effect on diversity. In models, stochasticity is typically implemented as variability in time that applies to entire populations.
The stabilizing effect we explore occurs when variability is packaged in
the bodies of individual organisms. Analysis suggests that the theoretical
need for tradeoffs and the empirical support for those tradeoffs can be
overemphasized. We present empirical and theoretical analysis to illustrate
both points. Using hierarchical Bayes demographic models, we demonstrate
1) that high variability within species is a source of massive stochasticity
that has low correlation among species, and 2) that parameter tradeoffs are
generally not evident among many coexisting species. Using models that
incorporate the individual effects parameterized from field data, we demonstrate its stabilizing effect on diversity.

CLARK, JAMES,1,* LORETTA JOHNSON,1 TYLER SUELTER,1 JAMES

KOELLIKER2 and ALAN KNAPP.1 1 Division of Biology, Kansas State
University, Manhattan, KS, USA; 2 Department of Biological and Agricultural Engineering, Kansas State University, Manhattan, KS, USA. Response of belowground carbon cycling to water and nitrogen in tallgrass prairie.
Atmospheric N deposition and water availability may have profound effects
on whether prairie ecosystems sequester or release C. Objectives are to
determine separate and interactive effects of water and N on belowground
carbon (C) cycling (soil respiration and root decomposition) and root biomass at Konza Prairie. We hypothesized that water will increase C cycling,
while N will be neutral or negative. Plots (2x2m) were treated with 3 levels
of water (250, 150 and 0 mm in excess of natural precipitation) combined
with 4 levels of N (0, 2.5, 5, and 10gN/m2/season). Plots were replicated
6 times in upland and lowlands (144 plots total). Root decomposition was
determined by burying 2g of Andropogon roots in mesh bags. Soil CO2
efflux was measured on 14 dates spanning two treatment seasons. After 7
months, water increased root decomposition in uplands and lowlands
(p5,.0001), but high N and high water depressed decomposition in uplands (p5.01). However, one year later, N had no significant effect on mass
loss in upland or lowland. Soil respiration was enhanced in uplands and
lowlands with water (p5,.0001) but was significantly depressed in high
N treatments. Root biomass increased in uplands and lowlands in response
to water (p,.0001) but not to N. Removal of water limitation increases
belowground C cycling in tallgrass prairie. In contrast, N addition may
depress soil C cycling and lead to greater belowground C sequestration.
CLAY, KEITH and ALISSA PACKER. Indiana University, Bloomington,
Indiana, USA. Pathogens vs. microbial mutualists.
Most plants in native systems are simultaneously infected by pathogenic,
mutualistic and commensalistic microbes. Their impacts on plants and vegetation dynamics are a function of microbial interactions and plant species
composition. A single plant-microbe interaction may range from pathogenic
to mutualistic depending on the ecological context and the criteria for classification. Long-term field studies with tall fescue grass have indicated that
fungal endophyte infection significantly suppresses plant species diversity
and reduces the rate of successional change compared to paired experimental plots with uninfected tall fescue. Intense grazing pressure by voles
favors infected fescue, resulting in greater dominance by fescue. In contrast
to the diversity-suppressing effect of endophyte infection of tall fescue,
long-term studies in forest systems focused on black cherry and soil-borne
pythium pathogens point to a diversifying effect of pathogen infection.
Pythium populations in the soil around adult cherry trees cause strong
distance- and density-mortality and growth suppression of cherry seedlings
and saplings. This impedes the development of dense stands of cherry
around existing adults. Preliminary evidence suggests that a similar process
is occurring in other temperate forest species. The Janzen-Connell hypothesis posits host-specific predators and pathogens, causing distance- and
density-dependant mortality of conspecific seedlings and saplings, contribute to the high diversity of tropical forests. We suggest that there are predictable changes in the relative balance of these positive and negative microbial interactions over large spatial and temporal scales that correlate with
observed diversity patterns. In particular, positive interactions with microbes characterize plant communities in harsh physical environments and
in younger successional habitats. Negative interactions become more important over successional time and with more productive environmental
conditions. A greater understanding of the effects of microbial interactions
with plants can enhance our understanding of vegetation dynamics.
CLELAND, ELSA E.1,2 and CHRISTOPHER B. FIELD.2 1 Dept. of Biological Sciences, Stanford Univerisity, Stanford, CA; 2 Carnegie Institution
of Washington, Dept. of Global Ecology, Stanford, CA. Selective herbivory as a mechanism to understand shifting plant community composition in response to simulated global changes.
Human activities are altering carbon (C) and nitrogen (N) cycling in ecosystems worldwide. Plant communites respond to these global changes both
physiology and with shifting species composition, potentially producing
ecosystem-level feedbacks. In California annual grasslands, slug herbivores

selectively remove species with high tissue %N early in the growing season. We investigated the hypothesis that experimentally induced global
changes would alter the extent of selective herbivory via shifts in plant
tissue chemistry, leading to functional shifts in species composition. In the
field, we planted six species in known locations both inside and outside of
herbivore exclosures, and exposed these plots to either ambient or elevated
levels of two simulated global changes: elevated atmospheric CO2 and N
deposition. Species were chosen to represent a functional axis of tissue
%N: two legume species with high tissue %N, two annual forbs with intermediate tissue %N, and two annual grasses with low tissue %N. Seedlings were monitored for germination and herbivory, and harvested to determine tissue % C and N. Under control conditions, species with high
tissue %N (legumes) were selectively removed by herbivory. In plots with
N deposition, all species responded with increased tisse %N, and herbivores
were less selective, allowing legumes to persist in the species composition.
Under elevated CO2, seedlings had lower tissue %N (n.s. trend), and herbivores were more selective, removing a greater proportion of legumes and
forbs than under control conditions. Thus, with elevated CO2
CLINTON, BARTON D.,1 JONATHAN L. HORTON,2 ERIK T. NILSEN,2,3 JOHN F. WALKER2 and COLIN M. BEIER.4 1 USDA Forest Service Southern Research Station, Coweeta Hydrologic Laboratory, 3160
Coweeta Lab Road, Otto, NC, USA; 2 Department of Biology, Virginia
Polytechnic Institute and State University, Derring Hall, Blacksburg, VA,
USA; 3 Botany Department, 3190 Maile Way Room 502, University of
Hawaii, Honolulu, HI, USA; 4 Institute of Arctic Biology, University of
Alaska Fairbanks, PO Box 75700, Fairbanks, AK, USA. The influence of
ericaceous shrubs on forest floor and soil properties in the southern
Appalachians.
In the southern Appalachians, evergreen shrub-dominated understories interfere with woody seedling establishment. Several sources of inhibition
have been studied including influences of soil and forest floor properties.
In this study we examined how soil and forest floor properties vary along
gradients of evergreen shrub basal area (BA). On three sites, we established
transects that transitioned between open understories and understories with
varying densities of Rhododendron maximum (rosebay rhododendron) and
Kalmia latifolia (mountain laurel). The three sites were a mid-elevation
ridge (DF), a low-elevation cove (BF), and a high-elevation southwestfacing slope (WS). A total of 60 2 x 2 m plots were randomly established
along transects on each site. For each plot we quantified soil chemical
properties and forest floor mass, carbon, and nitrogen as they related to
evergreen shrub presence and BA. On sites occupied by mountain laurel,
humus and litter C increased and soil pH decreased with increasing mountain laurel BA at DF, but little or no differences in soil cations were found.
Across all sites with rhododendron, soil Al was in greater supply and soil
P was in shorter supply. On BF and WS, litter and humus depth and mass,
and humus C increased with increasing rhododendron BA. On WS, humus
N and litter C increased, as well. However, litter N and soil pH decreased.
Across all sites, approximately 80% more C and 30% more N was contained within humus beneath rhododendron, and approximately 75% more
C and 20% more N in litter compared to the other two understory conditions examined. There were no differences in litter bulk density across sites;
however, humus bulk density was significantly lower by nearly 50% beneath rhododendron. These results help characterize spatial heterogeneity
in forest floor and soil prperties across differing understory conditions that
could potentially explain variation in seedling success.
COBBOLD, CHRISTINA A.,1,* MARK A. LEWIS1 and JENS ROLAND.2
1
Department of Mathematical and Statistical Science, CAB 632, University
of Alberta, Edmonton, Alberta, CANADA; 2 Department of Biological Sciences, B702, University of Alberta, Edmonton, Alberta, CANADA. The
impact of parasitoid emergence time on forest tent caterpillar outbreaks.
The forest tent caterpillar, Malacosoma disstria is a mass defoliator of
trembling aspen, Populus tremuloides and acts as a host for a number of
fly and wasp parasitoid species. Adopting the forest tent caterpillar as our
study system we have developed a discrete time model which incorporates
the relative timing of host density dependence and parasitism, allowing for
concurrent occurrence of these events. To quantify the model we carried

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67

out field experiments to measure density dependent mortality at each larval

instar. By proposing a model for the interaction between density dependence and parasitism we gained understanding of the mechanisms driving
the period and outbreak length of population cycles observed in this caterpillar species. Previous host-parasitoid models have only incorporated the
sequential order of parasitism and density dependence and do not permit
the two events to occur simultaneously. Our model, which includes the
relative timing of parasitism and density dependence frequently predicts
different dynamics compared to a model which neglects this timing. Even
when both models cycles, the period and amplitude differs significantly.
This has predictive implications for the forest tent caterpillar system, where
parasitism timing varies substantially.
COCHRAN-STAFIRA, LIANE * and EVA YAKIMOVA. Saint Xavier
University, cochran@sxu.edu, Chicago, IL; 2 Northern Illinois University,
DeKalb, IL. Sarracenia purpurea pitchers: Potential oviposition sites
for mosquito vectors of arboviral encephalitis?
1,2,

1 1

Wyeomyia smithii is the only mosquito to regularly oviposit in the waterfilled leaves of the carnivorous northern pitcher plant Sarracenia purpurea.
Furthermore, W. smithii is restricted to the geographical range of its host
plant because of its strict dependency on the plant for reproduction. The
evolutionary basis for this commensalism is not fully understood, nor is
the mechanism behind the exclusion of other container-breeding mosquitoes from these potential breeding sites. The increasing risk of arboviral
encephalitic disease transmission by container breeding taxa calls for a
better understanding of their choice of breeding habitats, especially since
pitcher plants are often abundant in regions of North America where such
diseases are becoming endemic. In a series of laboratory studies, we investigated whether the Asian Tiger Mosquito Aedes albopictus, a potential
carrier of West Nile Virus, could or would oviposit in S. purpurea pitchers.
A. albopictus females typically cling to the sides of a container habitat and
attach eggs above the water line. Females presented with intact, unaltered
pitchers and paper-lined plastic cups, chose the cups over 90% of the time.
Drowned females, but no eggs, were often found in leaves at the end of
these choice trials. Artificially enlarging pitcher openings, and providing
resting places for females inside the pitchers both increased the probability
of successful oviposition to nearly 100%. Our data suggest that pitcher
morphology, combined with female oviposition behavior, is a major factor
in deterring oviposition. We hypothesize that the small pitcher opening and
the slippery, scale-covered interior of the pitcher prevent females from
entering the pitcher and clinging to the walls. Our data also suggest that
the type of detritus in the pitcher is not involved in oviposition decisions
by A. albopictus.
COFFEY, KIM* and KATHERINE KIRKMAN.* Jones Ecological Research Center, Rt. 2 Box 2324, Newton, GA, USA. Germination phenology of several important ground cover species from the endangered
longleaf pine ecosystem.
Seeds of most temperate species have some form of primary dormancy at
the time of dispersal and may display annual cycles in germination requirements responding to seasonal temperature changes. Understanding environmental cues eliciting germination responses and the role of persistent
seedbanks is critical in developing species reintroduction strategies in a
restoration context. Little is known of dormancy characteristics of the many
ground cover species in the endangered longleaf pine ecosystem. The purpose of our study is to explore the germination phenology of several functionally or compositionally important ground cover species from this diverse ecosystem. Sixteen abundant species were selected for study (2 composites, 3 grasses, 10 legumes and 1 other forb). Nine hundred seeds of
each species (3 flats, 300 seeds/flat) were placed on potting soil in greenhouse flats under 63% shade cloth. Seed was spread and lightly pressed
into soil, ensuring contact while mimicking natural seedfall. Flats were
watered weekly and exposed to ambient outdoor temperature and light/dark
periods. Weather conditions (temperature, precipitation, etc.) were recorded
daily throughout the study. Germination was censused weekly and germinants were removed. Protrusion of the radicle was the germination criterion.
Preliminary results (first year) indicate that seed from composites, grasses,
and the other forb species have little dormancy, germinating within six
months of planting (no germination after 6 months). Legume germination

68

Abstracts

varied widely by species: two legumes (Lespedeza hirta and L. angustifolia) exhibited little germination (,10%) until well into year two, while
both species of Desmodium had over 90% germination within the first year.
For other legume species germination continued throughout the study period (24-63% first year germination). Further analyses incorporating weather data will provide insight into what environmental conditions will break
dormancy, and if pre-treatment, such as seed scarification, is required for
restoration plantings.
COLAUTTI, ROBERT I., IGOR A. GRIGOROVICH and HUGH J. MACISAAC. What determines invasion success?
Several recent attempts to predict invasions have focused on the ability of
species to invade novel habitats (i.e., invasiveness) and the susceptibility
of habitats to invasion (i.e., invasibility). We conducted a meta-analysis
of the recent (1991-2002) invasion literature to search for putative correlates of invasion success, but a lack of explicitly defined terms made generalizations difficult. In particular, the terms introduced, invasive and
naturalized and their derivatives contained implicit differences between
studies. Nevertheless, we found seven species-specific characteristics that
were always positively associated with invasiveness: invasion history,
germination success or rate, reproductive output, consumption efficiency, hermaphroditic/asexual reproduction, brain size, and small seed
size. Conversely, only one of five habitat-specific characteristics was concordant among all studies of invasibility: nutrient levels. Of particular
interest, studies examining resident species diversity found positive relationships with invasion success in ten cases, and negative relationships in
thirteen cases. Of 128 total studies, only seventeen examined propagule
pressure (i.e., variation in introduction effort) hypotheses, yet this factor
was a consistent predictor of invasion success for studies of both invasiveness and invasibility. Furthermore, correlations between propagule
pressure and other species- or habitat-specific characteristics implicate a
potential for propagule biases (artificial patterns of invasion success owing to non-random patterns in introduction effort). Since propagule pressure-hypotheses remain poorly explored, we introduce a novel framework
for the study of invasions that could aid in future attempts to predict invasions, and act as a basis for operational terms and concepts.
COLBOURNE, JOHN K.1,* and JOSEPH SHAW.2 1 Center for Genomics
and Bioinformatics, Indiana University, Bloomington, IN; 2 Dartmouth College, Hanover, NH. The development of genomic resources for measuring evolutionary responses to environmental challenges.
Until recently, an enormous effort was needed to apply genetic tools to
environmental studies, especially when striving to uncover the mechanisms
of phenotypic plasticity and the genetic basis for evolutionary adaptation.
An important consequence of the large-scale genome programs of recent
years has been the technological advancements that transform such studies
from the painstaking hunt for a few accessible genes to the rapid screening
of multiple loci for variation linked to specific phenotypic effects. By consequence, genomic resources targeting an ecologically-focused organism
are being developed by the Daphnia Genomics Consortium, whose goal is
"to develop the Daphnia system to the same depth of molecular, cell and
developmental biological understanding as other model systems, but with
the added advantage of being able to interpret observations in the context
of natural ecological challenges". While these resources are still nascent including a genomic information database and microarrays - there are sufficient data and tools to exploit the unique properties that make the freshwater crustacean Daphnia an ideal system for ecological genomic research.
Daphnia is also an established model for toxicological research. Experimental work is underway to characterize its gene-expression patterns specific to ecologically relevant levels of toxicological stress. Applications of
these tools to populations exposed to both natural and anthropogenic toxins
(which are arguably powerful agents of selection) can elucidate the potential and limits of adaptive evolution.
COLE, DENISE H.,* MEGAN BRADBURN and TIA-LYNN ASHMAN.*
University of Pittsburgh, Pittsburgh, PA. Sex-differential resistance and
tolerance to florivory in wild strawberry.
Sexual dimorphism can modify plant-animal interactions and while there
is ample evidence that the sexual morphs of dimorphic plants differ in their

attractiveness to mutualists, far less is known about their susceptibility to

antagonists (e.g., herbivores). Our understanding of either the traits that
bring about herbivore preferences or the potential for sex-differential tolerance to herbivory is still in its infancy. Using gynodioecious wild strawberry (Fragaria virginiana) and one of its major herbivores-the strawberry
clipper weevil (Anthonomus signatus), we examined sex-differential susceptibility and tolerance to herbivory. Field surveys of six populations of
wild strawberry in Northwestern PA verified that hermaphrodites experience significantly more damage by weevils than females. Data collected
from plants grown in a common garden revealed that the probability of
attack was influenced by several plant traits, including pollen and flower
production. Simulated weevil clipping was a good surrogate for natural
weevil clipping and reduced total flower number in both sex morphs. Females were significantly less tolerant than hermaphrodites in terms of fruit
and seed number. Hermaphrodites did not compensate fully in terms of
flower number but did in terms of fruit and seed production. Clipped hermaphrodites produced significantly more pollen per flower than unclipped
plants but this increase was not sufficient to offset flower losses due to
herbivory. These results suggest that clipper weevil herbivory results in
hermaphrodites whose allocation is shifted towards female function relative
to their natural state. This type of herbivore-induced plasticity has consequences for sex ratio and gender evolution.

COLLIER, SHAWN and CHAMPA SENGUPTA-GOPLAN. New Mexico

State University, Molecular Biology Program, Las Cruces, NM. Investigating plant sucrose breakdown in response to arbuscular mycorrhizal
fungi.

COLEMAN, MARK D. USDA Forest Service, Southern Research Station,

New Ellenton, SC. Total belowground carbon allocation measured with
both CO2 efflux and fine-root production.

COLLINS, BEVERLY S.1,* and LORETTA L. BATTAGLIA.2 1 Savannah

River Ecology Laboratory, Aiken, SC; 2 Southern Illinois University Carbondale, Carbondale, IL. Making it big in bottomlands: regeneration
strategies of intermediate species.

The belowground carbon compartment contains two thirds of terrestrial

carbon. Soil CO2 efflux is the major flux from that compartment, which on
a global basis is second only to the flux of gross primary production and
equal to net primary production. Yet the size of the soil carbon pool is
relatively stable indicating that autotrophic inputs are equivalent to the
large soil efflux. Of the autotrophic inputs, fine-root production may account for 10 to 50%, yet estimates are highly variable. Accurate measurement of input and efflux from the belowground carbon compartment is
required to understand soil carbon cycles. We have monitored soil CO2
efflux using infared gas analyzer methods, estimated fine root production
with minirhizotrons, trapped above ground litter inputs, developed allometric relationships for coarse root inputs, and measured changes in soil
carbon pools in loblolly pine and cottonwood plantations. Our carbon budget allows evaluation of errors, as well as quantifying pools and fluxes in
developing stands during non-steady-state conditions. Soil CO2 efflux was
larger than the combined inputs from aboveground litter fall and root production. Fine-root production increased during stand development; however, mortality was not yet equivalent to production, showing the belowground carbon budget was not yet in equilibrium and root carbon standing
crop was accruing. Belowground production was greater in cottonwood
than pine, but the level of pine soil CO2 efflux was equal to or greater than
that of cottonwood, indicating heterotrophic respiration was higher for pine.
COLEY, PHYLLIS D. and THOMAS A. KURSAR. University of Utah,
Salt Lake City, UT. Drug discovery as a conservation tool in Panama:
The use of ecological insight.
The discovery of compounds from nature that have agricultural or medicinal value often is promoted as a mechanism for obtaining tangible benefits
from biodiversity and thus helping to preserve wildlands. Here we present
data from Panama on a novel approach for drug discovery. We demonstrate
that the use of ecological theory on plant defenses increases the probability
of discovering compounds with activity in bioassays against human disease
targets (malaria, trypanosomiasis, leishmaniasis, cancer, and HIV). As predicted, extracts from young leaves were more active than those from mature
leaves, and extracts from shade-tolerant species more active than those
from gap specialists. Ferns and monocots were the least active, and basal
dicots the most active. Plant life form had little influence on activity. In
addition, a goal of our project is to provide immediate benefits to Panama
such that there are incentives for conservation of wildlands. To accomplish
this, we are conducting all of the research at Panamanian institutions, providing guaranteed and lasting benefits, even if royalties are never realized.
By conducting as much of the drug discovery research as possible in developing nations, the potential exists for providing more jobs than destructive uses of biodiversity.

Arbuscular mycorrhizal (AM) fungi consume 10-20% of the carbon fixed

by their host plants. AM fungi utilize hexose sugars that are released from
sucrose, but it is not known which plant enzyme is catalyzing this reaction.
Sucrose synthase is the main sucrose breakdown enzyme in plant sink
tissues, including the root nodule. However, I hypothesize that another enzyme, cell wall invertase, is breaking down sucrose for AM fungal metabolism. Cell wall invertase exists in the apoplastic space with the AM fungi.
In contrast, sucrose synthase is a cytosolic enzyme. Hexose sugars resulting
from sucrose synthase cleavage would require transport into the apoplast,
but no plant hexose transporters are known to export hexose sugars into
the apoplast. Instead they work in the opposite direction, moving hexose
into the cytoplasm. Although it appears that sucrose synthase transcripts
localize to plant cells containing mycorrhizal structures, I hypothesize that
this enzyme works antagonistically against cell wall invertase, directing
carbon to plant and nodule metabolism. I am characterizing the transcript
and protein levels of both enzymes in root and nodule tissues as well as
performing immunolocalization in response to mycorrhizal infection.

The dynamics of bottomland hardwood (BLH) forests are controlled largely

by plant response to heterogeneity generated by flooding and frequent
windstorms, which open the canopy and generate local microtopographic
variation. We examined regeneration strategies of canopy species relative
to flooding and light heterogeneity in BLH forests. Highly flood-tolerant
species are typically shade-intolerant and grow quickly enough to avoid
inundation. Most shade-tolerant species are flood-intolerant and restricted
to drier portions of the floodplain. In contrast, highly shade-intolerant species regenerate in large openings and colonize substrates over a range of
flooding conditions. Between these extremes, intermediate species comprise
the bulk of BLH canopy. These species may 1) have broad tolerances, with
regeneration driven by chance, dispersal, or biotic interactions or 2) exhibit
small-scale sorting over environmental heterogeneity and trade-offs in plant
traits over life history stages. We synthesized information for common
intermediate bottomland species, including Liquidambar styraciflua, Quercus michauxii, and Q. pagoda, and propose a conceptual model of regeneration strategies linked to species coexistence and regeneration patterns in
complex, highly productive bottomland forests.
COLLINS, RACHEL J.* and WATLER P. CARSON. University of Pittsburgh, Pittsburgh, PA. Do succession models predict the right pattern
for the wrong reason: shade vs. herbivore tolerance?
Succession models typically predict the replacement pattern of species following disturbance. One of the most prominent models, the shade-tolerance
model, uses species-specific tolerances of seedlings to low understory light
conditions as the mechanism to predict species replacement. A second model uses shade tolerance in combination with the timing of the disturbance
during succession (i.e., a stage-dependant model) as the mechanism for
predicting species replacement. Both of these models are based on resource-species interactions and ignore the potential effects of herbivores in
determining successional trajectories. White-tailed deer populations are at
historically high levels in many areas. By neglecting the effects of browsing, these models may either fail to predict successional trajectories or
make the correct prediction for the wrong reason (i.e., shade tolerance
instead of herbivore tolerance). We experimentally tested the predictions
of these two models with three types of disturbances: canopy gaps, understory fire, and a combination of gap and fire at two levels of herbivory. At
the Westvaco Research Forest in northeastern West Virginia, we have established 64 plots (400 m2) where we have permanently tagged, identified,
and measured nearly 17,000 seedlings, saplings, and adults of 26 species
of trees. In the absence of deer herbivory, the shade-tolerance model correctly predicted the dominance of shade-intolerant species (i.e., pioneers)

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after both fire and canopy gap formation. In the presence of deer herbivory,
however, shade-tolerant species (i.e., late succession) dominated after disturbances. The stage-dependant model did not consistently predict the dominant species. Herbivory is changing these successional trajectories, truncating succession, and preventing pioneer species from dominating during
early succession. Thus, under current conditions of overabundant deer populations that exist throughout much of eastern North America, forest succession models that fail to explicitly incorporate herbivory or tolerance to
herbivory will not consistently predict species composition and successional trajectory in forests.
COLLINS, SCOTT L.,1,* LORETTA C. JOHNSON,2 JAMES K. KOELLIKER2 and ALAN K. KNAPP.2 1 University of New Mexico, Albuquerque, NM; 2 Kansas State University, Manhattan, KS. Effects of water and
nitrogen on community structure in tallgrass prairie.
Recent evidence suggests that limiting resources may interact to affect vegetation structure in many plant communities. Tallgrass prairie vegetation
has been shown to be co-limited by nitrogen, water and light. We used two
long-term field experiments at Konza Prairie Biological Station to investigate the individual and combined effects of water and nitrogen additions
on plant community structure and dynamics in this tallgrass prairie. Results
from a 12-year irrigation experiment designed to minimize water limitation
during the growing season showed that water alone had little impact on
plant community structure, including changes in species richness or abundance of functional groups. However, abundance of Panicum virgatum increased significantly, while Andropogon gerardii abundance declined
slightly, in irrigated lowlands. In a second experiment, we added three
levels of water and four levels of nitrogen to upland and lowland vegetation. After only 4 years, forb and total richness declined with increasing N
addition, grass richness declined with increasing water addition, but there
were no significant water by nitrogen interactions. Again, P. virgatum
abundance increased, whereas abundance of A. gerardii declined, in fertilized and watered lowlands. These results show that nitrogen and water act
somewhat similarly, but independently, to shift the competitive balance
among dominant grasses in this tallgrass prairie.
COMAS, LOUISE,* HOWARD SKINNER and MATT SANDERSON.
USDA-ARS, Pasture Systems and Watershed Management Research Unit,
University Park, PA, USA. Using plant morphology, physiology and
stress response to define plant functional groups in pasture systems.
Interest in quantitatively defining plant functional groups has been increasing as ecologists strive to understand plant communities from a functional
perspective. Plant functional groups in current literature are most often
defined qualitatively. We divided 23 common pasture species from three
commonly defined functional categories (grass, legume, forb) into 8 functional groups using principal component analysis on plant growth responses
to low N and water availability determined in a greenhouse study. Sensitivity of plant biomass to drought explained similar variation between species as rooting depth. Growth of grasses was generally sensitive to N but
grasses could be separated into deep- and shallow-rooted species. Most
legumes were drought-sensitive/N-insensitive but a few were drought-sensitive/N-sensitive and drought-insensitive/N-sensitive. Shallow-rooted, Ninsensitive forbs were separated from shallow-rooted/N-sensitive and deeprooted/N-sensitive forbs. We also collected measurements of plant morphology, architecture and physiology to determine which traits were best
correlated with plant growth responses to low N and water. With the exception of several fast-growing legumes, fast-growing species in general
were the most sensitive to N and water availability. Fast-growing species
had a greater percentage of their roots at deeper depths at the expense of
shoot biomass.
COMITA, LIZA S.1,* and STEPHEN P. HUBBELL.1,2 1 Department of
Plant Biology, University of Georgia, Athens, GA, USA; 2 Smithsonian
Tropical Research Institute, Balboa, Ancon, Panama. Community level
patterns of seedling mortality in a tropical tree community.
Many of the theories proposed to explain the maintenance of diversity in
species rich plant communities hinge on mechanisms that shape spatial
patterns of seedling survival. The Janzen-Connell hypothesis (Janzen 1970,

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Connell 1971) posits that successful seedling recruitment in tropical forests

may be restricted by natural enemies, namely seed predators, herbivores,
and pathogens, that cue in on high densities of seeds and seedlings around
reproductive adults. In contrast, the regeneration niche hypothesis (Grubb
1977) suggests that plant species require differing conditions for recruitment, so the progeny of different species will survive and establish in
different habitats. In order to fully test these hypotheses, we have established a large-scale community-wide study of seedlings and small saplings
in the 50-ha Forest Dynamics Plot (FDP) on Barro Colorado Island, Panama. We have collected data on the abundance and spatial distribution of
.60,000 seedlings and small saplings of 344 tropical tree and liana species
in approximately twenty thousand 1-m2 plots distributed uniformly over the
50-ha FDP. Using data from two completed censuses of these seedling
plots, we examine patterns of seedling mortality in this tropical tree community and compare survival rates among species and among guilds. We
look at the effects of seedling size, habitat, and conspecific density on
seedling survival and examine the relative importance of these factors in
determining spatial patterns of recruitment. We use the results of these
analyses to evaluate the relative importance of mechanisms proposed to
explain the coexistence of species.
CONANT, RICHARD T. and MOFFATT K. NGUGI. Colorado State University, Fort Collins, CO, USA. Grassland management and light use
efficiency: Implications for remote sensing.
Light-use-efficiency (LUE) - the efficiency with which absorbed photosynthetically active radiation is converted into plant biomass - is a key variable
necessary for assessing aboveground net primary productivity (ANPP) in
grasslands using remotely sensed data. Quantifying LUE is important not
only for addressing scientific questions such as how management impacts
grassland biogeochemistry (e.g., C fluxes), but to enable producers to make
management decisions about livestock movement, forage stockpiling, and
herd culling. If LUE was known or well constrained, remotely sensed data
about the seasonality and amount of production could be used to answer
scientific questions or make management decisions. But management directly impacts LUE, thus precluding direct assessment with remotely
sensed estimates of productivity. One of the goals of this project is to
investigate the degree to which management impacts LUE. We used two
approaches. First, we reviewed the literature to examine how grassland
LUE varies with management across different grassland physioclimatic regions. As one component of that we estimated LUE for grasslands for
which ANPP data have been collected, thus generating independent estimates of LUE. Second, we evaluated the sensitivity of ecosystem model
production estimates to variation in LUE in order to evaluate whether incorporating more information about LUE would lead to more accurate remotely sensed estimates of ANPP. Review of the grassland LUE literature
immediately suggests that ignoring spatial, temporal, and management-induced variability in LUE will have dramatic impacts on accuracy of ANPP
estimates. Our results, in which using season, -region-, and managementspecific measures of LUE individually and in combination improve estimates of ANPP.
CONNER, CRISTIN A.,* TANA WOOD and DEBORAH LAWRENCE.
University of Virginia, Charlottesville, VA. Leaf litter addition effects
upon soil nutrients in a wet tropical forest.
Plant growth and soil organic matter in lowland tropical rain forests are
widely considered to be limited by phosphorus (P). Litterfall is a major
source of P return to the soil; hence, the relationship between litter inputs
and soil nutrient content is an important component of nutrient cycling in
these systems. We investigated soil-litter interactions in a tropical lowland
rain forest in Costa Rica. In order to evaluate this relationship, we experimentally manipulated litter along an age gradient in forest recovering from
pasture. At each of six sites, we removed litter from a 400m2 area and
added it to a 100m2 area with two replicates per site. This is equal to a
four-fold increase in litterfall. We sampled soil to 5cm depth in addition
and control plots monthly for a period of 7 months after litter addition.
Soils were air-dried and digested using a modified Kjeldahl digestion. The
digestate was then analyzed colorometrically for total P. Secondary forest
sites had significantly higher total P than mature forest sites (1.02 mg/g in
secondary versus 0.62 mg/g in mature; p,0.05). Total P was highest in

June and as the rainy season progressed, P decreased in both the control
and litter addition plots regardless of forest age. Thus, the decline in total
soil P is likely related more to environmental factors than to the amount
of litter being decomposed. Further research should focus on the influence
of environmental factors such as soil moisture and pH that might influence
this process.
CONOVER, ROSS R.,1,* ERIC T. LINDER,1 LOREN W. BURGER1,* and
TREY COOKE.2,* 1 Mississippi State University, Mississippi State, Mississsippi, USA; 2 Delta Wildlife, Stoneville, Mississippi, USA. Effects of
farm field borders on avian diversity, abundance and reproductive success.
Agricultural trends have considerably altered the amount of suitable habitat
for farmland birds. These trends may be associated with the population
declines of several wintering and breeding bird species in the southeastern
U.S. Farmland birds are expected to benefit from uncultivated field borders.
In the spring of 2002, approximately 16.5 miles of field borders were planted entirely with grasses or a grass/legume combination on seven farms
throughout the Mississippi Alluvial Valley. Thirty-eight of the planted field
borders are 10-meters wide, while six have greater widths (20-80m). We
also designated 19 control borders (10m wide), which represent traditional
ditch to ditch farming techniques. The diversity, abundance, and reproductive success of the associated avian communities were surveyed in the 2002
breeding season and the winter of 2003. Data on the reproductive success
suffered from delayed emergence of the planted vegetation. The vegetation
failed to extensively emerge until early autumn. No significance was detected between border treatments in the diversity or abundance of the avian
community, which may also be a product of late emergence. I will discuss
the reproductive success, as well as avian diversity and abundance on data
collected from the summer of 2003. Winter data suggests that 10m borders
planted with grass/legume have significantly increased avian diversity and
abundance within the border, when compared with control (unplanted) borders. Preliminary data also suggests that wide borders have increased diversity and abundance compared with treated and untreated 10m wide borders.
COOK, JESSICA L.* Michigan State University, East Lansing, MI. Effects of the pathogen Uromyces ari-triphylli on reproductive biology and
growing season of Arisaema triphyllum.
Arisaema triphyllum, Jack-in-the-pulpit, is a common spring ephemeral in
eastern forests of North America. Populations frequently contain plants
infected with the systemic rust Uromyces ari-triphylli. Previous research
demonstrated that disease reduces seed set in infected females, but results
came from a single A. triphyllum population over a single growing season.
In this study, I do a detailed comparison of flowering phenology, female
reproductive success, and length of growing period for healthy and infected
plants at four sites in Michigan. Light intensity, soil pH, soil moisture were
also measured to determine their influence on growth and flowering. Hand
pollination experiments were conducted to determine if reduced seed set is
caused by lower pollinator visitation of diseased plants. Environmental variables did not affect flowering period, emergence date, and growth period
of healthy or diseased plants, while disease influenced growing period and
flowering phenology. Diseased plants emerged significantly earlier (on average five days) than healthy plants. Growing season length was significantly shorter for diseased plants (average 41 days infected, 57 days
healthy). Average flowering period for healthy plants (28 days) was significantly longer than diseased plants (23 days). The percentage of healthy
females setting seed (71%) was also significantly higher than diseased females (17%). Hand pollination increased seed set in both healthy (34%)
and diseased females (133%). Healthy and diseased plants did not differ
in pollen viability. For females that produced seed, differences in the number of seeds produced were only slightly significant. This pathogen exerts
a strong influence on the life history of A. triphyllum by limiting flowering
time, growing season, seed production, and resource allocation to developing seeds. At the population level, this pathogen can dramatically reduce
individual plant growth and fitness, which can affect A. triphyllum population dynamics both genetically and numerically.

COOK, ROBERT1,* and KAREN NELSON.2 1 Leonard Parker Associates,

Architects, Minneapolis, MN, USA; 2 University of Maryland, College
Park, Maryland. Fully-integrated urbanism: Sustainability as inspiration.
This poster is a collaboration between urban design and ecology, presented
by an experienced architect and urban designer in collaboration with an
ecologist. We accept that, urban environments are both necessary (given
current world population levels), and desirable (to create the conditions for
exchange of information and culture). Fundamentally we seek to go beyond
the question of "how can ecologically sustainable communities be created"
to "how can the requirements for sustainability be viewed as inspiration
rather than obstacle?" In addition we seek to demonstrate urban planning
and architectural concerns for an ecological audience. Thus we demonstrate
how an urban area could be "fully integrated," creating a balance and providing a congenial environment for both humans and natural ecosystems.
We present an architectural design thesis approaching the redevelopment
of a dense core city location. The current inquiry looks at the potential for
exchanges between these two worlds in the areas of rainwater, greywater,
sewerage, smokestack emissions, solid waste, energy, air, biota, earth and
groundwater. This presentation is a creative endeavor and not a thorough
scientific analysis - in the firm belief that progress begins with inspiration.
Most importantly, we integrate much of the apparatus required to create
sustainability into the architecture in a highly visible manner, thus making
a statement that this is a place that is "infinitely sustainable as long as the
sun shines and the sky rains."
COOLEY, HEATHER S.,1,2,* WILLIAM J. RILEY2 and MARGARET S.
TORN.2 1 University of California, Berkeley, Berkeley, CA; 2 Lawrence
Berkeley National Laboratory, Berkeley, CA. Interactions between land
cover change and climate in a coupled regional model.
Regional climate affects the timing of harvest for rain-fed crops. In response to dry conditions, for example, farmers harvest crops earlier than
they do under wet conditions. This removal of vegetation alters the land
surface characteristics and may, in turn, affect regional climate conditions.
We applied a coupled climate (MM5) and land-surface (LSM1) model to
examine the effects of land cover change, i.e. winter wheat harvest, on
regional climate in the ARM-CART region of the Southern Great Plains.
We compared early and late harvest scenarios, with winter wheat harvested
on June 5 and July 5, respectively. Winter wheat accounts for 20% of the
total land area over the domain of the ARM-CART. Results showed that
regionally-averaged, 2 m air temperatures were 0.5-18C warmer in the early
harvest case compared to the nominal case, with peak warming of ca. 58C
centered over the harvested area. Soils in the harvested area were drier and
warmer in the top 10 cm. Near-surface soil water-filled pore space was
reduced by 15 percentage points, i.e., from ca. 50% WFPS in the nominal
to ca. 35% WFPS in the harvested areas. Soils were up to 108C warmer,
with area-averaged warming of ca. 0.68C at mid-day two weeks after harvest. Subsequent impacts of harvest on rainfall and latent heat fluxes were
also significant.
COOLEY, HILLARY C.* Florida International University, Miami, Florida.
Palm fuel dynamics in fire-sustained pine forests in the Florida Keys,
USA.
The relationships between time since last fire and the contribution of two
South Florida palm species (Silver thatch palm, Coccothrinax argentata
(Jacq.) Bailey and Thatch palm, Thrinax morrisii, H. Wendl.) to the burnable fuel load in seventeen sites in the lower Florida Keys were examined.
Combined, these two palms make up approximately 40 percent of understory shrub abundance. The fire history of each site was determined, and
the relationship between fuel amounts and time since last fire assessed. The
population size distribution, mortality rate of burned verses unburned sites,
fuel moisture of live and dead leaves, and growth rate of the palms were
also studied. Logistic regression models were developed to determine the
live and attached dead fuel biomass for each of the two palm species. The
equations were applied to the structural information obtained from each
site in order to estimate the average live and dead attached dry biomass
contributed by size category at different sites. Equations with crown area
and number of leaves as independent variables best predict the amount of

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71

biomass contributed by the palms. Live biomass continued to increase for

approximately 3 years after fire before leveling off, while dead attached
biomass continued increasing for about 11 years after fire. T. morrisii contributed a greater amount of total burnable fuel and also retained dead
leaves longer than did C. argentata. The average moisture content of the
dead leaves was much less than that of live leaves (35% and 106%, respectively). Plant material with lower moisture content will burn more readily and intensely, contributing to higher fire susceptibility. Knowledge
about the nature of attached palm fuels and the amount of time post-fire
before it stabilizes will be useful in developing ecologically-based management criteria in this fire adapted ecosystem.
COOPER, ROBERT J.* and JILL J. GANNON. School of Forest Resources, University of Georgia, Athens, GA, USA. Response of Prothonotary
Warblers to timber harvest and hydrology in a bottomland hardwood
forest.
The Prothonotary Warbler (Protonotaria citrea) is widely regarded as an
indicator of the health of forested wetland ecosystems in the sotheastern
United States. Bottomland forests in the lower Mississippi Alluvial Valley
are threatened by deforestation and altered hydrology; those in public ownership often are actively managed by timber harvest. We investigated the
effects of alternative timber harvest methods in bottomland forests of the
White River National Wildlife Refuge, Arkansas, USA on the productivity
of Prothonotary Warblers. Started in 1994, the study was a replicated, largescale manipulative experiment with a before-after, controlled intervention
design. Treatments (single tree selection, small patch cuts) reflected those
used on the refuge and were designed to mimic natural disturbances. Treated plots had fewer territories on them, but nest success (probability of
fledging at least one conspecific young) and productivity (fledglings produced per pair) was unaffected by timber harvest. Plot productivity (number of fledglings produced per plot) was less on treated plots than on control
plots. Nest success showed significant annual variation primarily due, we
believe, to annual variation in flooding. The major cause of nest failure
was nest depredation by a variety of predators. Years with normal flooding
resulted in higher nest success, and dry years, or dry periods within wet
breeding seasons, resulted in increased nest depredation. We believe that
in wet years, protection is afforded to nests built over water, and that alternative prey are available to primary nest predators. In dry periods, nests
are relatively unprotected and alternative aquatic prey are not available to
predators. Results suggest that: (1) the Prothonotary Warbler is a potentially
useful indicator species for bottomland forest ecosystems, (2) active forest
management is not a serious threat to Prothonotary Warblers in this system,
and (3) proposed water management projects on the White River, which
would affect hydrology, are a significant threat to this system.
COOPERBAND, LESLIE R. University of Wisconsin-Madison, Soil Science Department, Madison, WI. Biogeochemical cycles of urban and
livestock production wastes: Why do we continue to mine nutrients in
a world awash in surplus?
There are over 142 million Mg of municipal organic wastes or by-products
produced annually in the U.S., or 75% of the total amount of municipal
wastes generated. They include sewage sludge or biosolids (7 million Mg),
food scraps (20 million Mg), paper (73 million Mg), textiles (7 million),
wood (10 million Mg) and yard trimmings (25 million Mg). Of this total,
only 32% are recovered for reuse and over 60% are land filled. Almost 9
million Mg of carbon and 325,000 Mg of nitrogen are produced in yard
trimmings alone. Livestock wastes include animal manure, livestock mortalities and meat processing by-products. Total manure produced in the US
for confined livestock facilities exceeds 144 million Mg, representing 7
million Mg of N and 1 million Mg P. Trends toward concentration and
vertical integration of livestock production and processing enterprises have
resulted in large disparities between where organic livestock wastes are
generated and where they are needed for land application. In this presentation, I will describe current fates of these waste streams and the implications for atmospheric C and N losses and P loss to surface waters. I will
contrast the current unidirectional flow of mined nutrients (including C)
that are used in typical livestock and crop production systems with scenarios that promote recycling and soil sequestration of waste-derived sources of C, N and P. Lastly, I will link organic waste recycling to building

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Abstracts

organic matter in grass and shrub lands and describe the potential for use
of organic wastes to restore degraded, desertified grass and shrub lands.
COPENHEAVER, CAROLYN A. Virginia Polytechnic Institute and State
University, Blacksburg, VA. Influence of topographic position on dendroclimatic responses in white and chestnut oak in the southern Appalachians.
In the southern Appalachians topography influences vegetation distribution.
The ridge tops are characterized by chestnut oak (Quercus prinus), Table
Mountain pine (Pinus pungens), and hickories (Carya sp.). The valley forests are dominated by white oak (Quercus alba) and the cove forests by
eastern hemlock (Tsuga canadensis). The objective of this study was to
evaluate whether the dendroclimatic responses of white oak and chestnut
oak change with topographic position from ridges to valleys. Tree-ring
master chronologies were developed from five stands: a white oak valley
stand, a chestnut oak valley stand, a white oak ridge stand, a chestnut oak
north-facing ridge stand, and a chestnut oak south-facing ridge stand. All
stands were old-growth with the oldest stand being the white oak valley
stand where the oldest tree dated to 1722. The youngest stand was the
chestnut oak stand on the north-facing ridge stand where the oldest tree
began growing in 1743. Temperature and precipitation were correlated with
a tree ring width index from each stand to identify differences in relationships with growth and climate across sites and species.
CORBIN, JEFFREY D.,1,* TODD E. DAWSON,1 MEREDITH A. THOMSEN1 and CARLA M. DANTONIO.1,2 1 University of California, Berkeley,
CA; 2 USDA-ARS, Reno, NV. Summer water use by California coastal
prairie grasses: Fog, drought, and community composition.
Plant species in arid and semiarid habitats have been shown to rely on a
variety of character traits to survive prolonged drought conditions. Recent
advances in the use of stable isotopes have shown that, while many perennial plants in dry conditions access water deep in the soil profile, many
species also support shallow roots to take advantage of sporadic precipitation events. Coastal fog can be an important moisture source for California coastal vegetation, though its role in influencing species distributions
is poorly understood. We sampled the stable H and O isotope ratios of
water in the soil and roots of a variety of perennial grass species in four
sites in northern California in order to determine the relative importance
of fog and rain throughout the year. The relationship between dH and dO
provided evidence that evaporation of water in the soil had taken place
prior to uptake of water by vegetation. To account for this, we employed
a novel correction for evaporative fractionation, and then applied a twosource mixing model. Our results indicated that 20-50% of the water in
vegetation during the spring and summer came from fog rather than from
residual water from winter rain. Fog use decreased as distance from the
coast increased, and species differed in their use of each water source. We
did not find consistent differences between species whose distributions are
limited to coastal habitats versus species with broader distributions, but
rather found that species responded individualistically to summer fog and
exhibited varying degrees of dependence on fog. Native perennial grasses
in coastal habitats can be strong competitors against exotic annual grass
species. We believe that summer moisture inputs from fog may lengthen
the growing seasons for these perennial grasses, giving them an important
advantage over annual grasses.
CORNELISSEN, TATIANA G.* and PETER STILING. University of
South Florida, tgcornel@chuma.cas.usf.edu, Tampa, FL. Fluctuating
asymmetry of leaves affects herbivory by leaf miners on two oak species.
Fluctuating asymmetry (FA) represents small, random variation from symmetry and can be used as an indicator of plant susceptibility to herbivory.
We investigated the effects of leaf FA of two oaks species, Quercus laevis
and Q. geminata, and the responses of three species of leaf miners, Acrocercops albinatella, Brachys tesselatus, and Stilbosis quadripustulatus to
random variations in leaf morphology. Thirty plants of each species were
followed from leaf flush until mine termination and two aspects of leaf
quality were assessed: tannin concentration and water content. To examine
FA, 40 leaves from each plant species were collected, and FA was calcu-

lated as the unsigned difference between right and left widths, after digitalizing each leaf. Two indices of FA were calculated and plants were
divided into three classes of asymmetry: low FA, medium FA, and high
FA. Differences in growth of leaf miners between leaf types were determined by tracing five mines on 20 plants of each species on a 2-day interval. Asymmetric leaves contained significantly lower concentrations of
tannins than symmetric leaves for both plant species, but no differences in
water content were observed between symmetric and asymmetric leaves.
FA accounted for 57% of the variation on the attack rates of herbivores on
Q. geminata and for 54% of the attack rates on Q. laevis, and plants with
higher levels of FA were more attacked by Brachys and Stilbosis than
plants with lower levels of asymmetry. Although Brachys and Stilbosis
mines were smaller on asymmetric leaves, no differences in mine survivorship were observed between symmetric and asymmetric leaves. This
study indicated that leaf miners might use leaf FA as a cue to plant quality
and selection for oviposition sites, although consequences for offspring
performance in terms of differential survivorship were not observed.
CORNWELL, WILLIAM K.* and DAVID D. ACKERLY.* Dept. of Biological Sciences, Stanford University, Stanford, CA, USA. Assembly of
woody plant communities: Patterns of functional trait diversity.
The controls on functional diversity of communities are crucial to ecosystem function and have received relatively little attention. The persistence
of a species at a site is determined by the species functional traits, the
abiotic conditions, and species interactions. If a trait is strongly associated
with habitat tolerance, only species that possess a narrow range of that trait
value can establish and persist. Alternatively, if a trait is associated with
resources that are being divided among speciesa niche axisspecies with
similar values of that trait will be excluded and the spread of trait values
will be large. Here, we quantify the functional diversity of communities
using the variance in functional trait values of species. We present a study
of woody plant communities across a topographically-mediated gradient in
water availability in coastal California. We measured leaf and xylem functional traits for every woody species that occurred at our sites (including
measures of intraspecific variation across sites for widespread species). We
then constructed a null model in which we selected species randomly from
the pool of species in the study. The variance of the observed distribution
of functional trait values, relative to the null model, provides a test for nonrandom assembly processes. Overall, the observed variances of leaf and
xylem traits were significantly lower than in the null model, suggesting a
dominant role of habitat tolerance. However, there were contrasting trends
in trait variance across the abiotic gradient. At the dry end of the gradient,
variance of wood density decreased and variance of leaf size increased
suggesting inverse shifts in the relative importance of habitat and niche
partitioning for each trait. Future work will incorporate multivariate models
examining shifts in the distributions of trait combinations associated with
different plant strategies.
CORP, LAWRENCE A.,1,* ELIZABETH M. MIDDLETON,1 JAMES E.
MCMURTREY,2 JACOBY CARTER,3 PETYA K. CAMPBELL1 and
MARYN BUTCHER.1 1 Biospheric Sciences Branch, NASA Goddard
Space Flight Center, Greenbelt, MD; 2 Hydrology & Remote Sensing Laboratory, USDA Agricultural Research Service, Beltsville, MD; 3 Biological
Resources Division, USGS National Wetlands Research Center, Lafayette,
LA. Steady-state fluorescence sensing of wetland vegetation.
The disappearance and degradation of wetlands, and the means of restoration is a problem of great national importance and concern. In coastal
areas the distribution of submerged aquatic vegetation (SAV) communities
are controlled by a variety of abiotic factors that will be affected by global
change processes. These processes range from global changes in climate
due to anthropogenic factors, to changes in regional biophysical regimes
such as nutrient regimes and hydrology because of human modifications
of the landscape. Since the aquatic environmental gradient spans fresh,
brackish, and high salinity, changes such as sea level rise and precipitation
patterns can locally affect nutrients, salinity, and light availability. These
factors determine which species of SAV, including exotic and invasive species, can potentially grow and establish in an area. Several wetland species,
Scheonoplectus robustus, S. californicus, Phragmites australis and Distichlis spicata, were investigated with respect to their fluorescence spectral

properties. On plants acquired from natural settings, characterizations were

performed within the visible emission spectrum (400 - 760 nm) with ultraviolet excitation at 355 nm. We were able to identify spectral characteristics that differed among the three genera, and to separate the two species
of the genus Scheonoplectus, based on spectral characteristics. To assess
the responses of SAV communities in changing micro-environments, a salinity experiment was conducted with potted plants placed in large fiberglass tanks. The experiment was conducted as a 3 x 2 factorial design, with
three levels of salinity and two water depths. The most definitive results
were obtained with Phragmites australis, for which the red and far-red
chlorophyll fluorescence bands produced significant interactions between
water depth and salinity. Plants grown at a salinity of 10 ppt had significantly higher fluorescence intensity (p , 0.05) than those grown at 0 ppt.
Additionally, a significant increase in red fluorescence emission intensity
occurred for plants placed in deep water vs. shallow water conditions.
COSGRIFF, ROBERT J.* Great Rivers Field Station, Illinois Natural History Survey, Brighton, IL, USA. Predicting seed/seedling dynamics: The
non-Tarot card approach.
A seed/seedling dynamics study was conducted from 1996-2001 on five
key pools of the Upper Mississippi River System (UMRS). Twenty-four
species were identified on the five pools in which Populus deltoides, Betula
nigra, Acer saccharinum, Fraxinus pennsylvanica, Ulmus americana, and
Acer negundo were the most common species encountered. River pool,
forest community, species, and month were significant predictors of seed
density. Year was not a significant predictor of seed density. River pool
and germination year were significant predictors of seedling density, whereas forest community, germination month, and species were not. Although
B. nigra produced the greatest number of seeds and had high reproductive
power, this species had poor representation in the seedling cohorts of the
sites studied (germination ranged from 0.0-0.15%). Likewise, P. deltoides
had no seedling germination in any of the sites studied. The percent germination of A. saccharinum varied from 0.35% in the wet forests of the
La Grange Pool to 218.2% in the wet forests of Pool 13. Fraxinus pennsylvanica and U. americana seed germination ranged from 0.0-25.3% and
0.92-85.0%, respectively. Hard-mast species (Quercus and Carya spp.) had
much lower germination rates than what was expected (0.0-25.3%). Due
to low numbers of seedlings in all but five species, survivorship analysis
was conducted for F. pennsylvanica, A. saccharinum, U. americana, Rhamnus cathartica, and Zanthoxylum americanum. Seedling survival was
strongly related to species, river reach, and day of mortality. Fraxinus pennsylvanica, Z. americanum, and U. americana had greater long-term survivorship than did A. saccharinum and R. cathartica. Survivorship within a
species also varied according to forest community. Fraxinus pennsylvanica
had a greater chance of surviving on dry forests than wet (Chisquare532.74, df51, p,0.001) whereas A. saccharinum had a greater
chance of survivorship on wet forests (Chi-square541.76, df51,
p,0.0001). All river reaches other than Reach 4 and the La Grange Reach
showed significant differences in seedling survivorship curves. Likewise,
there were significant differences in survival curves between forest communities. Dry forests had greater seedling survivorship than wet forests.
COSTANZO, KATIE S.,* BANUGOPAN KESAVARAJU and STEVEN
A. JULIANO. Illinois State University, Normal, Illinois, USA. The effects
of habitat drying on competition of two container mosquitoes: Aedes
albopictus and Aedes aegypti.
The Asian Tiger mosquito, Aedes albopictus, was introduced into the U.S.
from Asia, and has become the most abundant container-dwelling mosquito
in regions of the country. In the southeastern U.S., there has been a decline
in the abundance or local extinction of Ae. aegypti following the invasion
of Ae. albopictus. Interspecific competition among larvae appears to be the
mechanism producing these negative effects on Ae. aegypti. However, the
geographic persistence of Ae. aegypti in urban and southern regions of
Florida associated with warmer, seasonally dry climates, suggests that abiotic factors may have an effect on competition between the two species.
We tested the hypothesis that habitat drying reduces or eliminates the competitive advantage of Ae. albopictus over Ae. aegypti. The laboratory experiment began with the following ratios of larval cohorts (Ae. albopictus:
Ae. aegypti): 52:0, 52:52, and 0:52. Larvae were distributed among four,

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73

150-ml cups in each cage, containing 120 ml of deionized water. Cages

were randomly assigned one of two simulated precipitation regimes: Fluctuating - water in the cups evaporated to 50% of the initial water volume
before refilling with deionized water; Drying - water in the cups evaporated
completely, and remained dry for two weeks before refilling. Adults were
enclosed into cages where they mated, took blood meals, and oviposited
in the cups. We analyzed adult production within cages in three 36-day
periods of the experiment using MANOVA. There was a significant interaction of species combination and treatment for adult production of both
species. In two-species cages, the advantage of Ae. albopictus over Ae.
aegypti in adult production was significant and increased over time in the
fluctuating environment. In the two-species cages in drying environments,
Ae. albopictus had a slight advantage over Ae. aegypti in adult production
early, but later the two species produced roughly equal numbers of adults.
The abundance of larvae was significantly affected by treatment for Ae.
albopictus and species combination for Ae. aegypti. These results provide
some support for the hypothesis that habitat drying may promote local
coexistence of these species.
COSTELLO, ANDRIA M.,1,* MELANY C. FISK,2 INES OTZ1 and TIM
J. FAHEY.3 1 Syracyuse University, Syracuse, NY, USA; 2 Appalachian
State University, Boone, NC, USA; 3 Cornell University, Ithaca, NY, USA.
Fungal decomposers of fine roots.
Fine root turnover is a large input of carbon to forest ecosystem detrital
systems, yet the identity of organisms responsible for decay is largely unknown. We explored the species composition of fungi (basidiomycetes and
ascomycetes) on decaying roots (0.2 - 0.3 mm diameter) of sugar maple,
yellow birch, and American beech in a northern hardwood forest in New
Hampshire. We detected from 1 - 10 fungal species on individual dead fine
roots, using terminal restriction fragment length polymorphism (t-RFLP)
analysis. Taxa were not consistently abundant across all dead roots samples.
Some species were abundant on 50% or more of root samples, but overall
there was high variability among roots in the numbers and species of decomposers. The fine root substrate appears to support a spatially heterogenous fungal communitiy, which may contribute to widely varying decay
rates that have been observed among individual fine roots.
COTTINGHAM, KATHRYN L.* Dartmouth College, Hanover, NH. Is
stability theory derived for removal perturbations relevant to a broader class of perturbations?
Two key measures of ecosystem stability are sensitivity, the magnitude of
response to perturbation, and return time, the time it takes a system to
recover after a perturbation. Ecological theory predicts that sensitivity and
return time depend on community structure and system productivity. However, this theory was developed primarily for catastrophic removal perturbations (e.g., hurricanes, stream scours), and it is not yet known whether
it is applicable to other common perturbations such as additions of nutrients
or exotic species. I am using simulation studies and field experiments to
evaluate the generality of removal-based theory using freshwater plankton
communities. Extensive numerical experiments with published simulation
models indicate that expectations derived from removal perturbations work
reasonably well for addition perturbations so long as the perturbation size
is below some threshold. Intriguingly, this threshold depends on community
structure (zooplankton size) and system productivity (phosphorus loading
rate) in both models. To begin testing these results, I quantified the effects
of zooplankton community structure and nutrient supply rate on phytoplankton responses to removal and addition perturbations in a mesocosm
experiment during summer 2002. The factorial experiment contrasted two
community structures (copepod-dominated and Daphnia-dominated) X two
nutrient supply rates (ambient vs. enhanced) X three perturbation treatments (control, 33% removal of seston biomass, 50% increase in available
nutrients). Responses to the nutrient addition were generally consistent with
expectations from theory and previous experiments: increased abundance
of Daphnia and small cladocerans reduced sensitivity and decreased return
times. However, the effects of the removal treatment were too modest to
estimate sensitivity and return time. Together, these studies suggest that
stability theory derived for removal perturbations is relevant for some, but
not all, addition perturbations. The next step is to clarify exactly when
current theory breaks down, and construct new theory for these situations.

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Abstracts

COULSON, TIM.* University of Cambridge, Cambridge, CB2 3EJ. Nonlinear and stochastic population dynamics and selection in large mammals.
Quantifying the impact of density, extrinsic climatic fluctuations, and demography on population fluctuations is a persistent challenge in ecology.
I analyzed the effect of these processes on fluctuations in population size
of ungulate populations that have been the focus of long-term individualbased studies. Because the age and sex structure of these populations can
fluctuate independently of population size, and because animals of different
age and sex respond in different ways to density and weather, identical
weather conditions can result in different dynamics within populations of
equal size. In addition, the strength of density-dependent processes is a
function of the distribution of weather events. I extended these stochastic
demographic models to incorporate associations between phenotypic traits
and vital rates. These models captured observed weak and fluctuating selection on several traits. Finally, I explore the potential impact of global
climate change on population dynamics and trait selection within these
populations.
COVICH, ALAN P.1,* and TODD A. CROWL.2 1 University of Georgia,
Athens, GA, USA; 2 Ecology Center, Utah State University, Logan, UT.
Fragmentation of river networks, freshwater shrimp distributions and
Caribbean climate change.
Long-term data on rainfall suggest that perennial rainforest streams are
rarely subject to drying of riffles or pools. However, droughts coupled with
increasing urban demands for freshwater are decreasing stream flow and
can alter some species ability to migrate upstream from estuaries to headwater pools. Decreased flows and pool depths restrict habitat, diminish
water quality and can disrupt chemical communication that influences predator-prey dynamics. This disruption of connections within river drainage
networks is likely to occur more often as drought frequency and intensity
are predicted to increase over the next several decades while human populations also continue to increase their needs for freshwater. As part of our
long-term studies of the effects of variable flow regimes on benthic detritivores, we sampled shrimp populations bimonthly using baited traps in
pools along 1200 m (from 305 to 480 m in elevation) of Quebrada Prieta,
a second-order headwater stream in the Luquillo Experimental Forest (Caribbean National Forest). Unusually low rainfall in 1994 caused some headwater riffles to dry out completely, resulting in isolated pools, reduced pool
volumes, and loss of access to microhabitats. Following contraction of
high-elevation pools, densities of the numerically dominant shrimp (Atya
lanipes) increased from 22 to 75 shrimp per m2 of pool area and a second
common species (Xiphocaris elongata) increased from 5 to 14 shrimp per
m2. A smaller percentage of adults of both species was gravid during the
drought. Following the 1994 drought (1995-1998), densities of both shrimp
species and reproductive activity of Atya returned to pre-drought (19901993) levels. Following the drought, Xiphocaris reproductive activity remained lower than in the pre-drought period. Predatory shrimp (Macrobrachium carcinus) moved farther upstream into the headwaters during
drought conditions presumably because of decreased habitat availability at
lower elevations. These increased concentrations of predator and prey densities along with degradation of habitat quality may increase vulnerability
of shrimp populations, especially in rivers where harvest regulations are
lacking.
COX, STEPHEN B.* and KEN R. DIXON. The Institute of Environmental
and Human Health, Lubbock, Texas. Spatially explicit models of bubonic
plague in black tailed prairie dogs.
A stochastic modeling approach has been successfully applied in an epidemiological setting to understand the dynamics of infectious diseases.
However, limitations on computing power pose a significant obstacle to
incorporating spatial dynamics into even relatively simple epidemiological
models. One approach to overcome this obstacle has been to represent
model dynamics via cellular automaton approximations, where fully stochastic model dynamics are approximated using cell transition probabilities,
or within relatively small grids of populations with nearest neighbor coupling. However, these approaches ignore many host and vector demographic processes that may be important in predicting disease dynamics. We

utilized a high-performance, parallel computing environment to explore a

stochastic Susceptible-Infected-Recovered (SIR) model of prairie dog and
flea dynamics in a truly spatially explicit manner. Simulation results suggest
that because prairie dogs are highly susceptible to plague, persistence of
plague in a regional metapopulation is highly dependent on prairie dog and
flea movement rates between individual colonies (i.e., subpopulations). Because prairie dog colonies increasingly are associated with urban environments, information on the degree of domestic animal contact with prairie
dogs and their fleas, as well as the degree of colony isolation, are critical
to managing potentially devastating epizootic outbreaks of plague.
COYLE, DAVID R.,1,* MARK D. COLEMAN,1 JOHN T. NOWAK,2,*
CHRISTOPHER J. FETTIG3,* and DONALD C. BOOTH.4,* 1 USDA Forest Service, Southern Research Station, New Ellenton, SC, USA; 2 USDA
Forest Service, Forest Health Protection, Asheville, NC, USA; 3 USDA
Forest Service, Pacific Southwest Research Station, Davis, CA, USA;
4
Bartlett Tree Research Laboratories, Charlotte, NC, USA. Testing the
plant vigor hypothesis in an intensively-managed forest.
The plant vigor hypothesis (PVH) states that herbivores preferentially feed
on more vigorous plants or plant parts. This presents a dilemma in agricultural or intensively-managed forest systems, as silvicultural, chemical,
and genetic manipulations are often employed to increase plant vigor and
ultimately production. Plant vigor is directly related to plant growth and
productivity in intensively-managed forest systems. Few studies have examined the PVH in this environment. The Short Rotation Woody Crops
Cooperative Research Program operates a 55 acre fertigated tree plantation
in South Carolina. Pest feeding and the resulting plant injury from several
arthropods were evaluated on trees receiving one of four silvicultural treatments: irrigation only, fertilization only, irrigation + fertilization or an untreated control. Arthropods studied included the cottonwood leaf beetle,
cottonwood leafcurl mite, and various ambrosia beetle species on eastern
cottonwood; and the Nantucket pine tip moth on loblolly pine. In general,
pests of cottonwood supported the PVH while the Nantucket pine tip moth
did not. Due to the unnatural environment created, intensively-managed
forests are more susceptible to pest outbreaks than are natural ecosystems.
A thorough understanding of pest populations and injury as they relate to
tree vigor is essential for effective management of these systems.
CRAIG, RACHAEL G.* National Science Foundation, Directorate for
Geosciences, 4201 Wilson Blvd., Arlington, VA, 22230. Growing Opportunities in Biogeosciences at NSF.
Growing recognition of the importance of interdisciplinary approaches in
answering scientific questions has led to creation of a Program at NSF that
spans all major disciplines of Geosciences and links closely with Biosciences. The program officially began in 2002 and by working closely with
scientific societies and interested scientists through workshops and working
groups is developing a series of funding opportunities, the first of which
(geomicrobiology) appeared in fall of 2002. Biogeosciences at NSF focus
upon themes that span disciplinary boundaries, cross many spatial and temporal scales and recognize the increasingly human-modified landscape. The
Biogeosciences Program emphasizes rigorous understanding of complex
systems and seeks to integrate the geologic and biologic viewpoints. Maturation of this field requires clarification of vibrant connections between
many disciplinary approaches: present and past, nano- to planetary, physical, chemical and biological. Collaboration with core programs, such as
Biological Oceanography, Geology and Paleontology and the NCAR Biogeosciences Initiative, allow development of complementary approaches.
Annual announcements, reflecting a growing budget that began at $5M,
will maintain previous themes and add elements as appropriate themes are
identified your input is welcome. Other opportunities for Biogeoscientists
are envisioned as this field grows and could include new disciplinary foci
within Divisions of the Directorate.
CRAINE, JOSEPH M.,1,2 WILLIAM G. LEE,2 WILLIAM BOND,3 PETER
B. REICH1 and SCOTT OLLINGER.4 1 University of Minnesota, Saint
Paul, MN, USA; 2 Landcare Research, Dunedin, 00, New Zealand; 3 University of Cape Town, Cape Town, 00, South Africa; 4 University of New
Hampshire, Durham, NH, USA. The resource economics of chemical and
structural defenses across nitrogen supply gradients.
Plant defenses are a fundamental regulator of the transfer of energy and
materials to higher trophic levels and can require a substantial fraction of

the resources acquired by plants. In order to better understand the role of

nutrient supplies in determining the prevalence of plant defense types, we
investigated the theoretical relationships between ecosystem N supply and
the net C gain of shoots that were undefended or defended in one of three
ways: by N-free chemical compounds, N-containing chemical compounds,
or structural defenses. We extended economic models of shoot resource
balance to include the relative value of C and N (derived from a model of
ecosystem C and N uptake) as well as depreciation, and amortization. In
general, the relative net C gain of plants along N supply gradients were
dependent on the type of defenses present and similar to changes in their
generally understood abundance along an N supply gradient. For example,
at low N supply, the additional C acquired when investing N in photosynthesis and C in defense is much higher than investing N in defenses. Only
at high N supply is it better to invest large quantities of N in defense rather
than additional photosynthesis. In a sensitivity analysis, net C gain of
shoots was most sensitive to factors that affect the relative value of C and
N and the rate of herbivory. This work expands our understanding of the
determinants of the distribution of chemical defenses and extends into
structural defenses, whose resource economics have largely been ignored.
CRAWFORD, CHRISTOPHER B.,* JOHN J. DILUSTRO, BEVERLY S.
COLLINS and LISA DUNCAN. University of Georgia, Savannah River
Ecology Lab, Aiken, SC, USA. Soil response to prescribed fire in mixed
pine-hardwood forests at Ft. Benning, Georgia.
Short-term response of soil quality and nitrogen availability were examined
in mixed pine-hardwood forests in the first season following prescribed fire
to determine if changes in nitrogen availability were influenced more by
nitrogen loss due to combustion or altered soil inorganic nitrogen transformations. Stands (16) were burned during winter-spring 2001-2. Soil temperature returned to pre-burn temperature within several hours and was
related to the time of day of the burn. Fire minimally consumed the soil
organic layer, which ranged from 2% to 36% of the pre-burn mass. Fire
effects on mineral soil extractible total nitrogen were variable, ranging from
a 54% decline to a 327% increase; however, all values were low (,6 mg
N/g-1 soil). Our data indicate the prescribed burns were low intensity and
had little or variable effect on soil nitrogen. Detailed longer-term studies
are needed to understand prescribed fire effect on soil nitrogen dynamics
and sustainability of upland mixed pine-hardwood forests.
CROMARTIE, WILLIAM J.* and JASON R. GLIDDON. Richard Stockton College, jamie.cromartie@stockton.edu, Pomona, NJ, USA. Biomonitoring in New Jersey blackwater streams: recalibrating an invertebrate bioassay.
In 2001, the New Jersey Department of Environmental Protection (NJDEP),
using the Ambient Biomonitoring Network (AMNET) protocol for invertebrate bioassay, designated several streams in the NJ Pine Barrens impaired water quality. These designations conflicted with assessments carried
out by the NJ Pinelands Commission, using fishes, macrophytes, pH, specific conductance, and percent developed land as criteria. In 2002, NJDEP
changed all sites in the Pine Barrens listed as impaired based on biological
sampling to insufficient scientific information. We have been working with
the NJDEP Great Egg Harbor River (GEHR) Watershed Management Area
project, the County of Atlantic and the Pinelands Commission to eliminate
conflicts between AMNET and other predictors of water quality. We analyzed existing water quality and biotic data and developed GIS coverages,
showing percent developed land in each sub-watershed. From these data,
we developed a conceptual model of water quality in the GEHR basin. We
selected sub-watersheds representative of conditions in the GEHR for preliminary study, sampled macroinvertebrates from gravel, mud and woody
debris and measured pH and specific conductance. Invertebrates were identified to genus or family level. Multivariate analyses were used to determine
relationships between invertebrate samples and other indicators. Invertebrate assemblages of the GEHR resemble those in blackwater rivers further
south on the coastal plain, which helps explain why NJ AMNET criteria,
developed for cobble-bottomed trout streams, work poorly. Strong correlations exist among the environmental variables, with percentage of developed land the best predictor. Invertebrate fauna is clearly affected by these
variables. In streams with impaired water quality, species not found in
pristine Pine Barrens streams increase. It appears that the macroinvertebrate

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75

fauna on woody debris in the streams is the best target for biological sampling. By studying this assemblage intensively, we hope to develop biological indices that can predict water quality change in the Pine Barrens.
CROPPER, WENDELL P.,1,* NICHOLAS B. COMERFORD2 and HUA
LI.2 1 School of Forest Resources and Conservation, Institute of Food and
Agricultural Sciences, University of Florida, Gainesville, Florida, USA;
2
Department of Soil and Water Science, Institute of Food and Agricultural
Sciences, University of Florida, Gainesville, Florida, USA. Optimizing
simulated fertilizer additions using a genetic algorithm with a nutrient
uptake model.
Intensive management of pine plantations in the southeastern United States
is rapidly increasing in importance. This management regime typically includes suppression of competitors and additions of nutrients to promote
fast growth. In order to help reduce the impact of this management system,
it is useful to minimize nutrient additions while meeting the demand imposed by rapid growth. We simulated P uptake for the initial four years of
loblolly pine plantation development using the SSAND (Soil Supply And
Nutrient Demand) model. To optimize P addition we used a Genetic Algorithm (n 5 250 model runs per generation, 25 generations) with fitness
based on minimizing the difference between observed nutrient demand and
simulated uptake. Optimal day of nutrient addition and amount added was
sensitive to the root density inputs. An optimal P addition of 14.5 kg P/ha
at day eight was found with no simulated mycorrhizae, but with mycorrhizae the optimal solution shifted to 6.7 kg P/ha added on day 576 and
7.0 kg P/ha added on day 1012. Examination of the fitness surface around
the optimal solutions indicated a complex topology with some broad areas
of near optimal fitness.
CROSS, ALISON,1,* KATHLEEN K. TRESEDER1 and JAMES T. RANDERSON.2 1 University of Pennsylvania, Philadelphia, PA; 2 California Institute of Technology, Pasadena, CA. Preliminary estimates of mycorrhizal biomass by region.
While laboratory assays address small-scale activities of mycorrhizal fungi,
global biomass estimates are needed to scale up to regional and global
functioning. In a literature search, mycorrhizal root colonization data were
compiled from 80 published field studies in natural ecosystems, including
both arbuscular mycorrhizae (AM) (as % root length) and ectomycorrhizae
(ECM) (as % root tips). Global means of 3502 data points were calculated
for 11 biomes dominated by AM or ECM fungi. Neither AM (4062) nor
ECM (46610) percent colonization varied significantly by biome. We used
a previously published model of hyphal architecture in conjunction with a
global database of root length to calculate global mycorrhizal biomass.
Moderate investment in AM biomass (6.03109 Mg, versus 13.73109 Mg
for live fine roots) yields a large return in hyphal length (39.731015 km
hyphae versus 0.8831015 km roots). ECM biomass estimates agree with
those obtained from chloroform fumigation methods. Preliminary estimates
of global mycorrhizal biomass should be useful in calculating the proportion of the global C budget and C fluxes attributable to fungi.
CROWDER, LARRY B.,1 WILL F. FIGUEIRA1,* and FELICIA COLEMAN.2 1 Duke University Marine Lab, Beaufort, NC, USA; 2 Florida State
University, Tallahassee, FL, USA. Assessing the impact of commercial
versus recreational marine fisheries catch: The importance of who,
when, and where.
Many of the federally managed marine fish stocks are considered to be
either "fully-" or "over exploited". The blame for this has typically fallen
on commercial fisheries due to the perceived industrial nature of their activities and the notion that with such limited catch, recreational fishing
simply cannot compare to the commercial catch. In fact, for all marine
finfish fisheries in the United States, the recreational take is certainly quite
low, on the order of 2 to 5%. However fisheries are not managed as one
lump unit but rather on a species by species or stock by stock basis. When
viewed on this level, there can be dramatic differences in the impact of
commercial versus recreational fishing. In this study we use data from the
National Marine Fisheries Service on commercial and recreational catch of
marine finfish stratified by fishery, fishing mode, and time to analyze the
relative impacts of the two fishing types on the scale relevant to manage-

76

Abstracts

mentthat of individual species. Results for specific species are discussed

individually; however, in general we found that conclusions about the relative impacts of commercial versus recreational fishing were in fact species
specific and temporal analysis allowed us to look at these changes over
time. Such information is very valuable for making decisions on how to
best protect our marine resources and makes especially clear the need to
consider management options on either commercial or recreational fishers
on a fishery by fishery bases rather than attempting to establish any blanket
policy adressing who deserves regulation and who does not.
CROZIER, LISA G. University of Chicago, lcrozier@uchicago.edu, Chicago, IL. Predicting range shifts in response to climate change: studying a butterfly using convolution integrals.
Climate change is already causing a majority of species to shift their geographic ranges in expected directions. However, our ability to predict the
timescale of individual responses is limited by a widespread assumption
that environmental conditions at the range edge should remain constant.
Experimental work on Atalopedes campestris, the sachem skipper butterfly,
suggests that either summer or winter temperature can limit the northern
edge of the range. I here describe a model that demonstrates how the rate
at which it responds to climate change will depend on the rate of warming
in different seasons. This approach reveals some of the complexities inherent in environmental constraints and the importance of considering flexible population dynamics within the broader context of climate change.
CRUMRINE, PATRICK W.* University of Ketucky, Lexington, Kentucky.
Examining the role of size structure on intraguild predation in larval
odonates.
The direction and intensity of intraguild predation (IGP) interactions are
heavily influenced by size differences among interacting species. Asymmetric IGP is common when IG predators are larger than IG prey, and
many empirical studies on IGP include predators with these relative size
relationships. In addition to size differences between species, individuals
in most natural populations of predators and prey vary in size, but few
studies specifically test how size variation within species influences IGP
interactions. The aim of this study was to investigate how IG predator size
structure influences the survival and behavior of IG prey and shared prey.
I tested for these effects in a larval odonate system by exposing shared
prey (Ischnura verticalis) to the presence or absence of IG prey (Pachydiplax longipennis) and two size classes of IG predators (small or large
Anax junius) in a 2x2x2 factorial design. Both size classes of A. junius
significantly decreased I. verticalis survival and activity level, but there
were no significant effects of P. longipennis on I. verticalis survival or
behavior. P. longipennis responded differently to the two size classes of A.
junius. P. longipennis survival was significantly lower when exposed to
only the large size class of A. junius, and P. longipennis also decreased
their activity level in the presence of large A. junius. Survival of the small
size class of A. junius was also lower in the presence of large A. junius.
These results demonstrate that different size classes of IG predators can
impose varying levels of risk on IG prey and shared prey, but these effects
depend on the assemblage of predators in the system. Within-species size
structure adds significant complexity to IGP interactions, and studies incorporating this feature should provide a more complete understanding of
how IGP influences community structure.
CUMMING, GRAEME S.* University of Florida, Gainesville, FL. Correlates of fish species richness: Environment, low-head dams, and questions of scale.
The grain and extent at which observations are made will determine the
likelihood that the effects of particular drivers of biotic pattern will be
detectible. In many instances, regional patterns are not the sum of local
patterns. In this paper I ask whether the relatively small-scale mechanism
of the effects of low-head dams on fish communities can be scaled up from
local to regional extents. I examine the relationships between local and
regional environmental variables, the number of downstream dams, and
local fish species richness in first-order streams in Wisconsin, USA at several different extents and the grain of a 1:100,000 map. The analysis is
based on an extensive database of 13,628 localities and c. 180,000 indi-

CUMMING, JONATHAN R.1 and HEIDI B. LUX.1,2 1 West Virginia University, Morgantown, WV, USA; 2 Marine Biological Laboratory, Woods
Hole, MA, USA. Nitrogen-Aluminum Interactions in Mycorrhizal Tulip Poplar.
Elevated nitrogen deposition is contributing to soil acidification in forest
ecosystems, potentially mobilizing soil aluminum (Al). Al is phytotoxic
and may cause nutrient deficiencies in forest trees. Further, the form of N
available, ammonium or nitrate, and preferentially taken up by plants affects the uptake of nutrients through alterations of the cation:anion balance.
Symbioses between plant roots and mycorrhizal fungi facilitate greater uptake of nutrients, as well as potentially conferring tolerance to Al in the
root zone. However, the amount and form of nitrogen available are significant factors in the ability of mycorrhizae to successfully colonize plant
roots. Liriodendron tulipifera (tulip poplar), an important southern Appalachian tree species, is known to benefit from mycorrhizal symbioses, and
has demonstrated sensitivity to Al at concentrations as low as 50 mM. In
a previous study with an acid-tolerant community of arbuscular mycorrhizal
(AM) fungi, L. tulipifera seedlings demonstrated only limited symptoms of
Al toxicity when exposed to Al concentrations up to 200 mM. In the present
study, tulip poplar seedlings inoculated with field-trapped mycorrhizal inoculum from the Fernow Experimental Forest were exposed to 0 or 100
mM Al, high and low levels of nitrogen, and high and low ratios of ammonium to nitrate in sand culture for 7 weeks. L. tulipifera seedlings exhibited reduced root mass, increased root:shoot ratio, and decreased mycorrhizal colonization when exposed to 100 mM Al, and reduced Mg, P,
and Fe nutrition. Ammonium nutrition also inhibited colonization, and this
effect was coupled with decreases in growth. Seedlings exposed to high
levels of NH4-N accumulated N at concentrations higher than their NO3treated counterparts, suggesting that tulip poplar preferentially takes up
NH4-N, but that ammonium nutrition disrupts the uptake of other nutrient
cations, likely due in part to decreases in mycorrhizal colonization.
CURRAN, MARY CARLA.* Savannah State University, Savannah, GA,
USA. Preliminary evaluation of brown marsh near Savannah, GA.
Dead or brown marsh is indicated by large bald patches devoid of Spartina,
and has been observed in coastal Georgia. The Savannah State University
Marine Ecology Class is evaluating this phenomenon. The purpose of the
study is to provide an opportunity for students to collect data valuable to
a topic of current ecological concern. To assess the marsh areas, the densities of live and dead Spartina shoots were determined using transects
performed through live and dead marsh areas. Shoots were counted and
measured in 0.5 m2 quadrats. Preliminary results indicated that dead shoots
outnumbered live shoots in both the live and dead marsh areas. Some dead
marsh areas had no live shoots. Furthermore, dead shoots tended to be
taller than live shoots. Preliminary evaluation of the sediment indicated
that more meiofauna, primarily nematodes, were found in live marsh. In
the future, we will evaluate whether these dead patches are expanding.
CELINE, BOUTIN1,* and MARTIN PAMELA.2 1 Environment Canada,
Ottawa, Ontario, Canada; 2 Environment Canada, Burlington, Ontario, Canada. Patterns of earthworm distribution in crops and woody hedgerows
in organic and conventional farms.
The increasing intensity of agriculture and urbanization in southern Ontario
has resulted in a drastic decline of many species of wildlife. Often the only

D
R
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remaining habitats in the agricultural landscape for wildlife preferring

wooded areas are the narrow woody hedgerows that remain between fields
on some farms. Even these small fragments may be further altered through
the regular use of pesticides that may inhibit the growth of native plants
and reduce the abundance and diversity of invertebrates inhabiting them.
Earthworms represent essential elements of soil dynamics in many ecosystems, including agricultural systems. We compared the diversity and abundance of earthworms in organic and conventional farms in Peterborough,
Ontario, Canada. The objective was to examine the importance of field
crops and adjacent woody hedgerows as refuge for earthworms, and to
assess the impact of long term routine use of agrochemicals (pesticide and
fertilizer) and other management practices (e.g., cultivation, manure) within
these habitats. Earthworms were collected in the spring of 1999 (n56), in
the fall of 1999 (n516) and 2000 (n511), using the hot mustard extraction
method. Worms were identified to species and weighed to obtain an estimate of biomass. Soil characteristics were measured and management practices recorded. Species richness was similar among habitats and farming
types. Number of earthworms was higher in hedgerows than in fields.
Abundance and biomass was higher in organic than conventional hedgerows but the opposite was found in cropfields due to difference in management practices. For example, ploughing fields in the fall decreased the
number of earthworms considerably. This is part of a larger project that
investigated differences in biodiversity and abundance of native plants,
invertebrates and birds, in conventional versus organic agriculture, taking
also into account the landscape matrix.

W
IT

vidual collection records. The environmental variables include climate, elevation, distance from nearest town, and network-specific measures such
as link number (the number of first-order tributaries upstream of the current
reach), downstream link number, stream order, and fractal dimension. For
these data. the link number of the downstream reach is the strongest broadscale correlate of fish species richness in first-order streams. In general,
while numerous environmental variables (including the number of downstream dams) are correlated with fish species richness, I found that the
abiotic-biotic relationships in streams at this scale are weak; any regional
effect of dams on fish species richness can not be distinguished from natural, environmentally-driven variations in species richness. The picture is
further confused by the non-random placement of dams along environmental gradients. Collection of a more temporally competent data set would
provide a more rigorous test of this result.

DABUNDO, DENISE M.* and DAVID J. LONGSTRETH. Louisiana

State University, Baton Rouge, LA. Effects of submergence and hypoxia
on root elongation and aerenchyma development in rice (Oryza sativa
L.) seedlings.
Aerenchyma - a specialized tissue with abundant, interconnected gas spaces
- facilitates oxygen transport from shoots to roots in many emergent wetland plants. The importance of oxygen transport to root elongation and
aerenchyma development in rice seedlings was studied. Imbibed seeds of
Lemont and red rice were positioned at 4.5, 2.5, or 1.0 cm below the
surface of aerated or hypoxic water to control the duration of leaf submergence and the availability of oxygen. Daily measurements of primary
root and leaf lengths revealed that root elongation rates 1) were slower in
hypoxic water than in aerated water and 2) increased after leaves emerged
from 1.0 and 2.5 cm of overlying hypoxic water but not after leaves
emerged from 4.5 cm of overlying hypoxic water. In aerated roots, little
gas space was present in the first 10 mm behind the root tips and the
increase in gas space to a maximum occupied over half the roots lengths.
In roots of emergent seedlings that grew in hypoxic water, gas spaces began
to form 1 to 4 mm behind the root tips and near-maximum amounts of gas
space were found along over three-quarters of the roots lengths. The results
suggest that when seedlings grow in hypoxic water, oxygen transport can
1) enhance root elongation when water is not too deep and 2) promote a
root anatomy that favors efficient oxygen transport to the root tip.
DAVANZO, CHARLENE.* Hampshire College, Amherst, MA. Research
on learning: Little impact on ecology teaching.
For some scientific disciplines recent advances in research on learning has
lead to significant changes in teaching. For example, studies of expert vs.
novice learners of physics suggest that experts organize knowledge into
"chunks" (schemas) of readily accessible patterns of information. As a result, physics faculty have developed practices that help learners repeatedly
identify, recall, and apply a few fundamental schemas. In contrast, application of cognition research to college ecology teaching is rare. Examples
of how cognitive research would be useful for ecology faculty include
teaching ecosystem processes and altering misconceptions studenta have
about ecology. For ecology teaching to change, access to research on ecology learning must be as easy as access to ecology research, teaching reform
must be rewarded, and ecologists and cognitive scientists should work together to study ecology teaching and learning.

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77

DSA, JEAN V.1,* and KELLY S. JOHNSON.2,* 1 Environmental Studies,

Ohio University, Athens, Ohio, U.S.A; 2 Department of Biological Sciences, Ohio University, Athens, Ohio, U.S.A. Effects of acid mine contaminated sediments on Ctenodaphnia magna in laboratory bioassays.
Acid mine drainage or AMD is a major worldwide environmental problem
that adversely affects both surface and ground waters. AMD streams are
not only characterized by low pH but also by high concentrations of heavy
metals. Although the effects of metals in the water column on macroinvertebrates have been reasonably well investigated, the effects of metal
contaminants in the sediments are less well understood. Understanding the
conditions under which sediments may or may not retain toxicity to biological communities is important for predicting the success of specific remediation efforts. Laboratory bioassays using Ctenodaphnia magna (Order:
Cladocera) were performed to test the AMD contaminated sediments for
toxicity and sediment transplantation field experiments were carried out to
determine the residual toxicity of these sediments over time. Sites (around
Athens, Ohio) with reduced pH levels, visible metal precipitates and elevated water column concentrations of iron, aluminum, and manganese were
identified, and 7-day laboratory bioassays in static containers using C. magna were carried out. Sediments from all four of the AMD impacted sites
were found to be highly toxic to the daphnia. Toxicity was correlated to
reduced pH levels but was not correlated with any specific metal content.
In another experiment, AMD contaminated sediments were filled in baskets
and transplanted into the bed of a healthy stream for a week. Washed
sediments of each site were collected on the 2nd, 5th and 7th day and were
monitored for toxicity by performing 5-day laboratory bioassays using C.
magna. After 7 days in the clean stream, the same aluminum and ironcontaminated substrates were no longer found to be toxic to the daphnia.
These results suggest, that sediments at these sites will not retain toxicity
after remediation of the AMD input.
DAILEY, THERESA B.,* PETER E. SCOTT and XIAO-CHUAN LIU.
Indiana State University, Terre Haute, IN. Nectar sugar composition and
quantities in insect-pollinated Midwestern spring herbs.
To compare the nectar resources offered to flies and small, solitary bees
by native flowers and exotic weeds, we studied the woodland flowers Erigenia bulbosa, Claytonia virginica, and Cardamine concatenata and the
agricultural weeds Stellaria media and Barbarea vulgaris. One-day nectar
accumulations in covered, unvisited flowers were analyzed via high performance liquid chromatography (HPLC) to determine sugar identity. Sugar
quantities were determined from HPLC data and large additional samples
(40 flowers per species per day) were analyzed by spectrophotometry. Fructose, present in 85-100% of all samples, was the most commonly found
sugar in all species, and was used in making standards for spectrophotometry. Glucose and/or sucrose were found in each species, but never consistently. Despite the tiny quantities and multi-step measurement process, we
obtained repeatable mean nectar sugar accumulations for each species. Erigenia bulbosa and Stellaria media (means 5 24 and 32 micrograms) offered the smallest amounts of sugar, while Cardamine concatenata (mean
5 172 micrograms) offered the largest. Claytonia virginica and Barbarea
vulgaris offered similar amounts of sugar (means 5 78 and 80 micrograms). Thus, both habitats had a low-reward and a moderate-reward flower. The most rewarding flower was a woodland species, but it was less
common than the other 4 species. This suggests that per-flower nectar rewards in midwestern woodlands and adjacent agricultural fields may be
similar in spring.
DAMM, MARY C.* and WILLIAM D. BOWMAN. University of Colorado, Boulder, CO. Alpine plant biogeography and species endemism in
the Rocky Mountains.
The distribution of vascular alpine plant species in the Rocky Mountains
was examined over a latitudinal gradient. Studies in mountain systems in
the northern hemisphere indicate that the number of alpine species with
wide-ranging, circumpolar distributions is greater in areas closer to the
arctic tundra, while the number of species with narrow ranges is greater in
mountains further south. The pattern of alpine vascular plant species distributions was identified in alpine floras over fifteen degrees of latitude
from the Front and Main Ranges in Alberta, Canada, to the Sangre de

78

Abstracts

Cristo Mountains in New Mexico. The percentages of widely and narrowly

distributed species were compared among the northern, middle, and southern Rocky Mountain physiographic regions. The geographic distribution of
alpine plant species is significantly related to latitude. Species with wideranging circumpolar and North American distributions are greater in the
northern Rocky Mountains. In contrast, the percentage of narrow-ranging
Rocky Mountain and endemic species is greater in the southern Rocky
Mountains.
DAMSCHEN, ELLEN I.* and NICK M. HADDAD. North Carolina State
University, Raleigh, NC. Effects of landscape corridors on plant species
richness.
Habitat degradation and loss are the most important factors causing the
extinction of species. Corridors, linear strips of habitat connecting two otherwise isolated patches, have often been implemented to minimize the negative effects of habitat fragmentation. To date, corridor effects have been
frequently documented in animal populations, while the effects on plants
and communities have remained largely unexplored. At the Savannah River
Site in South Carolina, we assessed plant species richness in a replicated
landscape-level experimental system that controls for patch size and shape
while testing for effects of corridors. These sites were created in the winter
of 1999-2000. The presence or absence of all plant species within our
experimental landscape was recorded in June of 2001 and 2002. According
to metapopulation theory, we predicted that connected patches would have
higher species richness than unconnected patches. In 2001, there was no
difference in species richness between connected and unconnected patches
(p50.29). However, results from 2002 indicate that there is a trend toward
higher species richness in connected patches (p50.06). These findings indicate habitat connectivity may influence species interactions and successional processes.
DANE, LAURA,1,* DAVID LIPSON,1 SPRING STRAHM,1 RICHARD
WILSON,1 RAMESH MURTHY2 and GREG BARRON-GAFFORD.2
1
Department of Biology, San Diego State University, San Diego, CA; 2 Columbia University, Biosphere 2 Research Center, 32540 S. Biosphere Road,
Oracle, AZ. Effects of elevated atmospheric CO2 on soil microbes in
the Intensive Forestry Biome of Biosphere2.
We studied microbiological properties of soils from three atmospheric CO2
treatments (400, 800, and 1200 ppm) at the Intensive Forestry Biome of
Biosphere2 research center. We measured substrate induced respiration
(SIR), enzymatic activities, and microbial biomass (by fumigation-extraction and microscopy). There was high spatial variability within the CO2
treatments for most of the variables measured. Microbial biomass carbon
(C), bacterial numbers, and glucose SIR all tended to increase with increasing CO2 concentration. This trend was significant for glucose SIR (P50.04)
and was marginally significant for microbial biomass C (P50.075). The
community may also be functionally different between CO2 treatments; the
proportion of the the microbial community that degraded various substrates
varied significantly. The ratio of glycine SIR to glutamate SIR was lowest
in the 800 ppm treatment (P50.02), whereas the ratio of salicylate SIR to
glucose SIR was highest in the 800 ppm treatment (P50.04). These data
indicate that the microbial community changes nonlinearly in response to
CO2, with some effects being more pronounced at intermediate levels of
CO2. We attempted to explain spatial variation in soil microbial properties
by comparing soils collected from the north, middle and south portions of
the plots, and by measuring soil organic matter (SOM) in each soil sample.
SOM showed a marginally significant increase with increasing CO2
(P50.065), possibly indicating increased plant root production. However,
unlike most natural ecosystems, correlations between SOM and soil microbial properties were generally weak and non-significant. The middle portion
of the plots showed some marginally significant differences from the north
and south ends in SIR of salicylate (0.077), glucose (0.044) and glycine
(0.065), and had somewhat higher moisture contents (P50.078). We conclude that there are differences in microbial biomass and activity resulting
from the CO2 treatments, but that spatial variability must be addressed in
further studies.

DANG, QING-LAI* and SONG CHENG. Lakehead University, Thunder

Bay, Ontario, Canada. Effects of soil temperature on ecophysiological
traits in seedlings of four boreal tree species.
One-year old seedlings of black spruce (Picea Mariana (Mill.) B.S.P.),
white spruce (Picea glauca (Moench) Voss.), jack pine (Pinus banksiana
Lamb.) and aspen (Populus tremuloides Michx.) were exposed to 5, 10,
15, 20, 25, 30 and 35 oC soil temperatures for three months. Net photosynthesis, stomatal conductance, mesophyll conductance, midday xylem
water potential, photosynthetic water use efficiency, the internal to ambient
CO2 ratio (Ci/Ca), and specific leaf area were evaluated. The ecophysiological variables generally showed a bell-shaped response to soil temperature. The relationships between soil temperature and the ecophysiological
variables (except Ci/Ca) were modeled using 3rd-order polynomials. The
models suggest that the average optimal soil temperatures for the physiological traits were 21.8, 19.6, 22.4 and 26.1 oC, respectively, for aspen,
white spruce, black spruce and jack pine. Photosynthesis in aspen and jack
pine was progressively more limited by mesophyll conductance (relative to
stomatal limitation) as soil temperature increased from 5 to 35 oC whereas
in the spruces, mesophyll conductance was the most limiting to photosynthesis at much lower soil temperatures (10 to 20 oC). Aspen was generally
more sensitive to low soil temperatures than the conifers while the conifers,
particularly black spruce, were more sensitive to high temperatures. Most
of the conifer seedlings in the 35 oC soil temperature treatment died at the
end of the experiment while all the aspen seedlings survived.
DANZ, NICHOLAS P.,* JIM LIND, JOANN HANOWSKI, GERALD
NIEMI and MALCOLM T. JONES. Natural Resources Research Institute,
University of Minnesota Duluth, Duluth, MN, USA. Forest bird monitoring in northern Minnesota and Wisconsin 1991-2002.
The western Great Lakes region has among the highest richness of forest
breeding bird species in the continental US and is often considered a bird
population source for agricultural landscapes in the lower Midwest. A longterm regional monitoring program was implemented in 1991 to address
concerns over potential declines of forest breeding species in the region.
About 1400 permanent sampling points were located in four study areas
so that the sample was representative of the forest within each area. Birds
were surveyed annually by experienced observers using 10-minute point
counts. To characterize population trends, which are directional changes in
population size over a specified time period, we used simple linear regression to model an index of abundance as a function of time. Trends were
computed for species having a minimum level of occurrence; the number
of species tested per study area ranged from 40 to 53. The ratio of increasing:stable:decreasing species was consistent at about 1:3:1 across the four
study areas, although the identity of species in each trend direction category
changed somewhat across the areas. Six species showed consistent increases across the areas: Least Flycatcher, Red-eyed Vireo, Black-capped Chickadee, Cedar Waxwing, Chestnut-sided Warbler, and American Redtart. Nine
species showed consistent declines: Eastern Wood-Pewee, Winter Wren,
Hermit Thrush, Ovenbird, Connecticut Warbler, Mourning Warbler, Common Yellowthroat, Song Sparrow, and Brown-headed Cowbird. With the
exception of nesting strategy, there were no common threads to explain the
pattern of increasing and decreasing species. No species that nest on the
ground had increasing trends, while seven of the eight declining species
that build nests do so on the ground. We are investigating the hypothesis
that decreased abundance of ground nesting birds is related to a recent
regional increase in forest edge and a concomitant increase in mammalian
nest predators surrounding edges.
DAOUST, ROBERT J.* and JAMES T. MORRIS. Department of Biological Sciences, University of South Carolina, Columbia, SC, USA. Nutrient
controls on oligohaline marsh macrophyte community zonation and
production (Plum Island Ecosystem LTER).
Distinct zonation of marsh macrophyte communities is a prominent feature
of estuarine wetlands. These zonal patterns exist along strong gradients of
salinity stress which co-occur with ecosystem-wide nutrient limitation of
macrophyte production. Past research has explicitly sought to delineate the
relative importance of nutrient availability and salinity stress as factors in
controlling these zonation patterns. This research, however, has been fo-

cused within saltmarshes. Macrophyte community zonation also occurs in

upstream, oligohaline marshes but little research has been done in these
systems. Macrophyte dynamics within oligohaline marshes is, however, inherently more complex than in downstream systems since nutrient availability varies in response to fluctuations in freshwater discharge and marine
inputs. We utilized in situ reciprocal transplants of Typha angustifolia and
Spartina patens (two co-occurring dominant macrophytes in an oligohaline
New England marsh), nested within a factorial nitrogen and phosphorus
fertilization experiment, to elucidate mechanisms which control the zonation patterns observed within oligohaline marshes. Species-specific responses to nutrient additions exist with T. angustifolia responding to phosphorus and S. patens to nitrogen, indicating that both of these nutrients are
local determinants of macrophyte production. The reciprocal transplants
further show that while both species are able to survive within their neigbouring community, when nutrient availability is manipulated T. angustifolia outperforms S. patens. These results imply that changes in nutrient
loading to estuaries will directly affect the distribution and zonation patterns of macrophyte communities within oligohaline marshes. Furthermore,
this study indicates that the role of phosphorus in estuarine systems may
be more important than has previously been suggested.
DARCY, TARA L.* W. K. Kellogg Biological Station, Michigan State
University, tdarcy@kbs.msu.edu, Hickory Corners, MI. Linking algal species traits to environmental contexts: A test of the keystone predation
hypothesis.
Species face ecological and evolutionary tradeoffs that greatly impact their
survival and abundance in different environments. For example, the "keystone predation hypothesis" posits that traits which confer greater competitive ability also may make species more vulnerable to predation. Thus,
along gradients of increasing productivity and predators, the dominant species within communities shift from superior competitors to predator tolerant
species. To determine whether such "keystone predator" tradeoffs operate
within a system, one must investigate the correlations between species
traits and their predicted environmental associations. During 2002, I conducted a series of mesocosm and microcosm experiments to explore the
traits of benthic algae that confer success in various environmental contexts. I used the fourth-corner matrix analysis, a recently-devised and powerful statistical approach, to study the correlation matrix of algal traits
(superior competitive ability for limiting resources and predator tolerance)
and environmental variables. Our results indicate that the tradeoffs assumed
in the keystone predation hypothesis are valid for our system. For example,
the ability of benthic algae to compete in a phosphorus limiting environment is significantly negatively correlated with environmental productivity,
whereas the ability to resist predation by a grazing snail (Physa spp.) is
significantly positively correlated with environmental productivity. These
results demonstrate the usefulness of the fourth-corner analysis in studying
species tradeoffs and they illustrate the value of a mechanistic approach to
understanding the turnover of species across environmental gradients.
DARNELL, TRACI M.* and KATHRYN E. SIEVING. University of Florida, Gainesville, FL, USA. Habitat permeability and corridor use by
endemic south-temperate rainforest birds.
Our research examines corridor use and matrix permeability for a fragmentation sensitive bird, the Chucao Tapaculo (Scelorchilus rubecula), endemic to South American temperate rainforest. Probability of inter-patch
dispersal by Chucaos is being assessed directly, through translocation experiments, and indirectly, based on landscape-scale patterns of patch occupancy. Translocation experiment results indicate that Chucaos are behaviorally resistant to crossing open habitats (pasture), but will move through
forest corridors and matrix dominated by dense shrubs. Preliminary analysis of patch occupancy data (275 patches) also shows a strong landscapecontext effect, with patches surrounded by open habitat having lower probability of occupancy than those connected by corridors or surrounded by
permeable (shrubby) matrix. Thus, all results available to date support the
hypothesis that matrix types differ in resistance to Chucao movement, influencing patch occupancy patterns at the landscape scale. These empirical
data will also be used to analytically derive coefficients for scaling colonization probability curves for each of the dominant matrix types. The
colonization curves will be used to parameterize a graph-theory model for

Abstracts

79

predicting metapopulation persistence under various potential landscapeplanning scenarios.

DARROUZET-NARDI, ANTHONY,1,* CARLA DANTONIO1,2 and ERIC
BERLOW.3 1 University of California, Berkeley, Berkeley, California;
2
USDA-ARS, Reno, Nevada; 3 University of California, San Diego, White
Mountain Research Station, Bishop, California. The effects of invading
young sagebrush (Artemisia rothrockii) on the water status of an herbaceous montane meadow in the Eastern Sierra Nevada.
Rothrock sagebrush (Artemisia rothrockii) invades montane meadows on
the Kern Plateau in the Eastern Sierra Nevada. Sagebrush is typically associated with drier terraces or drier soil patches within more mesic terraces.
It is not clear whether it contributes to soil drying after it invades. We
hypothesized that young sagebrush shrubs contribute to soil drying by their
transpiration, thus accelerating the conversion of herb-dominated sites to
sagebrush patches. To test this, we manually removed young sagebrush
shrubs from 3.5 x 3.5 m plots in an area with 4-8 year old sagebrush plants.
Time domain reflectometry measurements of soil moisture in the removal
and control plots showed no difference between treatments at any soil depth
down to 60 cm over the entire growing season. We also measured transpiration rates of sagebrush and meadow herbs. These corroborated the soil
moisture data by showing that young sagebrush shrubs are not transpiring
enough water to dry out meadow soils. Meadow soil moisture levels are
highly variable seasonally and across sites in this study. We conclude that
the effect that young sagebrush have on soil moisture is small compared
to this variability. Large-scale hydrological changes such as a falling water
table and smaller-scale fluctuations in underground water sources are likely
more significant factors than sagebrush transpiration in determining water
availability.
DAS, ABHIK,1,* MICHAEL R. RIGGS,1 MICHAEL C. ZICUS2 and DAVID P. RAVE.2 1 Research Triangle Institute, 6110 Executive Blvd., Suite
420, Rockville, MD, U.S.A.; 2 Minnesota Department of Natural Resources,
Wetland Wildlife Populations and Research Group, 102 23rd Street, Bemidji, MN, U.S.A. Space-time modeling of a discrete ecological response: Characterization of Mallard nesting in Minnesota.
Discrete responses in the form of binary, categorical or count data, distributed over space and time, are common in ecology. For example, the U.S.
Prairie Pothole Joint Venture seeks to increase waterfowl populations by
reducing nest loss to predators. Mallards nesting in constructed nest structures often experience higher nest success than those nesting on the ground.
Occupancy of these structures can be thought of as a binary (yes/no) outcome. Identifying environmental features (such as land use and cover attractiveness) responsible for the geographic distribution of nests over time
is crucial to understanding the ecological mechanism affecting mallard nest
distribution. Using this knowledge to predict occupancy of structures would
be particularly useful for waterfowl management, since it would optimize
deployment of structures where probability of use was greatest. In this
study, we developed a generalized linear mixed model (GLMM) to characterize the spatio-temporal distribution of Mallard nests in 2 types of nest
structures. Our approach used logistic regression, which is natural for binary data, while extending its scope to accommodate both spatial structure
and temporal trends. Moreover, in order to identify the size of the area
surrounding the nesting structure that had the most influence on nest occupancy, we present a likelihood-based procedure for model selection in a
GLMM. The results show that, even after adjusting for spatio-temporal
effects, ecological features such as nesting cover attractiveness type of nesting structure, and size of open-water area in deployment wetlands were
significantly associated with nest occupancy. Our approach can accommodate space-time modeling for any discrete outcome. Thus, given the
prevalence of such data in ecological studies, we believe these methods are
broadly applicable to a variety of ecological research questions.
DASZAK, PETER.* Consortium for Conservation Medicine, Palisades,
NY, USA. The emergence of a new genus of lethal zooonotic paramyxoviruses in Australia and Malaysia.
Emerging infectious diseases (EIDs) are the key future threat to global
human health through the emergence of previously unknown or novel zoo-

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Abstracts

notic agents such as A.I.D.S. Zoonotic viruses (transmissible between humans and animals) make up 75% of known human EID agents. Therefore,
predicting their emergence is a holy grail of EID research so far impossible
to achieve. In the current talk, I focus on a new genus of viruses (Henipavirus) that emerged during the 1990s in Australia and Malaysia. These
viruses have high human case fatality rates (.40%), no known therapies
or vaccines, and a lack of knowledge of their range, biodiversity and ecology. The first Henipavirus to emerge, Hendra virus, killed 15 horses and
2/3 people in three separate outbreak events in Australia during the 1990s.
The second, Nipah virus, was more devastating, killing over 100 people in
Malaysia and Singapore during 1998-1999. Both viruses have fruit bat
(Pteropodid) reservoir hosts and appear to require a domestic animal intermediate host (horses and pigs respectively) for transmission to humans.
By analyzing recent changes in land use, agricultural patterns, climate, fruit
bat migration and distribution, we are testing hypotheses on the anthropogenic factors that drove Henipavirus outbreaks. We have developed a
preliminary matrix model for Henipavirus emergence that will be parameterized using these data to ultimately assess the risk of future outbreaks.
Recent reports of Nipah-like virus outbreaks in Bangladesh and India, and
the presence of antibodies in fruit bats of other southeast Asian countries
suggest a wider biodiversity and distribution of this genus. Our approach
will be directly applicable to predicting emergence throughout the Henipavirus range as future viruses are discovered. Finally, the emergence of
Henipaviruses highlights the double impact of some anthropogenic environmental changes that both reduce available wildlife habitat and promote
the interspecies transmission of microbes.
DAUGHERTY, MATT* and CHERIE BRIGGS. University of CaliforniaBerkeley, Berkeley, CA. Incorporating predator phytophagy into intraguild predation theory.
Thus far, intraguild predation (IGP) theory has focused on small subunits
of food webs, consisting of a top consumer, an intermediate consumer, and
a shared resource. In many systems, IGP interactions occur in the context
of a more complex food web, with multiple possible prey items for predators. Polyphagous predator populations are supplemented by feeding on
prey outside of the IGP module. This is especially true for highly omnivorous predators, such as predatory heteroptera, who are involved in IGP
with other predators, and also feed on multiple herbivores and plants. We
investigated the effect of a trophic supplement, predator phytophagy, on a
3+ level IGP model that includes a plant, an herbivore, and two predators
(IG prey and IG predator). Predator phytophagy affects the ability of the
predators to persist along a productivity gradient and to suppress the herbivore population, and the effects are dependent on which predator is plant
feeding. Phytophagous IG prey can invade at lower plant quality, persist
at higher plant quality, and suppress herbivores better than non-phytophagous IG prey. A phytophagous IG predator invades at lower plant quality,
it displaces the IG prey at lower plant quality, and it disrupts herbivore
control more than a non-phytophagous IG predator. Thus, IG prey phytophagy increases the potential for predator coexistence and pest suppression,
while IG predator phytophagy reduces coexistence and pest suppression.
Yet, even small amounts of phytophagy tend to destabilize the dynamics,
and extinction of one of the predators due to unstable dynamics is likely
over much of parameter space.
DAVELOS, ANITA L.1,* and ANDREW M. JAROSZ.2 1 University of
Minnesota, St. Paul, MN; 2 Michigan State University, East Lansing, MI.
Demographic analysis evaluating the effectiveness of biocontrol of
chestnut blight.
The largest stand of blight-free American chestnuts remaining in North
America was located in West Salem, WI until 1986 when chestnut blight
was discovered. Since 1991, the stand has provided the unique opportunity
to study the dynamics of a blight epidemic and serve as a site for evaluating
hypoviruses as biological control agents. A transition matrix model has
been utilized to determine if hypovirus infections of the chestnut blight
pathogen, Cryphonectria parasitica, allow American chestnut trees to recover with regard to growth, survival and reproduction. The degree of
recovery at a population level has been assessed by comparing the diseasefree, epidemic (new infections not treated with hypovirus), and old epidemic (hypovirus application) areas of the West Salem population. If hy-

poviruses are acting as an effective biological control agent, transition matrix models should predict similar demographic parameters within the disease-free and old-epidemic areas.
DAVIDSON, ANA D.1,2,* and DAVID C. LIGHTFOOT.1,2 1 Department of
Biology, University of New Mexico, Albuquerque, NM, USA; 2 Sevilleta
Long-Term Ecological Research Program, Department of Biology, University of New Mexico, Albuquerque, NM, USA. Effects of prairie dogs and
banner-tailed kangaroo rats on desert grassland vegetation in the
northern Chihuahuan Desert.
Prairie dogs (Cynomys spp.) and banner-tailed kangaroo rats (Dipodomys
spectabilis) are keystone species in semi-arid grasslands. Their burrowing
and foraging activities are known to effect community composition and
structure. In the American Southwest where these species co-occur, the
relative effects of their activities on grassland ecosystems are unknown.
Although prairie dogs and kangaroo rats appear to have similar functional
roles, these species are ecologically different and have unique effects on
grassland communities. This research evaluated their comparative and interactive influences on vegetation at the Sevilleta National Wildlife Refuge
(SNWR), New Mexico and near Janos, Chihuahua, Mexico. At the SNWR,
Gunnisons prairie dogs (Cynomys gunnisoni) created diffuse disturbance
patches on the landscape, and kangaroo rats created more intense localized
disturbance patches. The overall degree of disturbance was enhanced where
the species co-occurred. At the Janos research site, the effects of blacktailed prairie dogs (Cynomys ludovicianus) and kangaroo rats on vegetation
were less apparent; their impacts appeared to be overridden by intensive
cattle grazing. Interestingly, the exclusion of prairie dogs and kangaroo rats
from plots at the SNWR resulted in a significant increase in the exotic forb
species, Russian thistle (Salsola kali); this general pattern was qualitatively
observed at the Janos site.
DAVIDSON, CARLOS. Environmental Studies Department, California
State University, Sacramento, Sacramento, CA, USA. Historical pesticide
use and California amphibian declines.
Pesticides have long been proposed as a possible cause of amphibian population declines, but due to a number of challenges there has been relatively
little research on pesticides and amphibian declines. This study examined
the association between the spatial pattern of declines for five California
amphibian species and historic pesticide use in California from 1974 to
1991 based on Department of Pesticide Regulation records. Information on
declines was derived from maps of historic sites and current population
status for the Yosemite toad (Bufo canorus), California red-legged frog
(Rana aurora draytonii), foothill yellow-legged frog (R. boylii), Cascades
frog (R. cascadae), and the mountain yellow-legged frog (R. muscosa).
Generalized additive models were used to analyze the relationship between
site status (i.e., presence/absence of each amphibian species) and total upwind pesticide use, upwind pesticide use by 66 pesticide classes, and covariates including spatial location (longitude and latitude), precipitation,
elevation, and surrounding urban and agricultural land use. Total upwind
pesticide use was a strong, significant predictor of amphibian presence/
absence. A model based on total upwind applications of cholinesteraseinhibiting pesticides fit the data significantly better than models based on
total pesticides or on any other individual class of pesticides. This is the
first study in which population declines of multiple declining species have
been associated with historic pesticide applications.
DAVIDSON, JENNIFER M.,1,2,* PATRICIA E. MALONEY,2 ALLISON
C. WICKLAND2 and DAVID M. RIZZO.2 1 Pacific Southwest Research
Station, Forest Service, USDA, Albany, CA, USA; 2 University of California-Davis, Davis, CA, USA. Forest composition influences transmission of Phytophthora ramorum (Sudden Oak Death) in California oak
woodlands.
Phytophthora ramorum, the causal agent of Sudden Oak Death, is now
known to infect over twenty native plants in California oak woodlands.
The potential for P. ramorum to spread in these forests depends on its
ability to form reproductive spores on hosts during winter rains and to
survive within hosts during hot, dry summer months. We have begun to
investigate sporulation and survival of P. ramorum in two common forest

types in Northern coastal California: mixed-evergreen forest (dominated by

coast live oak (Quercus agrifolia) and bay laurel (Umbellularia californica)) and tanoak-redwood forest. To date, studies placing infected plant
tissue in moist chambers show that spores are produced on bay leaves from
mixed-evergreen forest, and bay leaves, tanoak (Lithocarpus densiflorus)
leaves, and tanoak twigs from tanoak-redwood forest. We initiated additional studies on the survival of P. ramorum in bay leaves because the
pathogen sporulates prolifically on this host, which may be key for spread
of P. ramorum. Monitoring infected bay leaves in both forest types indicated that infected bay leaves were significantly more likely to abscise in
drier mixed-evergreen forest as opposed to tanoak-redwood forest. Furthermore, leaf abscission and forest type influenced survival of P. ramorum. Recovery of P. ramorum from attached, infected bay leaves at the
end of summer was 24% and 60% from a mixed-evergreen forest and a
tanoak-redwood forest, respectively. In contrast, recovery of P. ramorum
from abscised bay leaves collected during summer months from leaf litter
in a mixed-evergreen forest was nearly zero. In tanoak-redwood forests,
higher survival of P. ramorum in bay leaves throughout the summer and
additional spore production from dominant hosts like tanoak may contribute to differences in levels and timing of inoculum produced in these two
forest types during the winter rainy season.
DAVIDSON, R. SCOTT1,* and HARALD STEEN.2 1 University of Otago,
Dunedin, Otago, New Zealand; 2 University of Oslo, Blindern, Oslo, Norway. Seasonal changes in density dependent population growth rate of
feral house mouse populations.
Density dependence is difficult to detect in wild populations. The effect of
density on population growth rates can easily be obscured by habitat heterogeneity and environmental stochasticity. By measuring vital rates on
replicate populations under a variety of conditions we are better able to
take these factors into account and isolate any effect of density. We examined the relationship between density and population growth rate in
seasonally breeding populations of feral house mice (Mus musculus) living
in coastal dune vegetation near Dunedin, New Zealand. Our study design
consisted of eight live-trapping grids that we trapped every second month
from February 2000 to June 2002. The experimental design allowed us to
measure vital rates with both temporal and spatial replication. Mark-recapture data were analysed with program MARK. Mouse apparent survival
was estimated using the Cormack-Jolly-Seber model, and recruitment into
the population was estimated using the Pradel models. For each grid at
each trapping session we produced a population projection matrix from the
survival and recruitment estimates, and calculated the population9s asymptotic growth rate. We fitted a generalised linear model to the growth rates
to analyse how they varied with density and the time of year. While there
was no overall relationship between the population9s growth rate and the
density of the mice, there was a seasonal density-dependent effect. The
population9s growth rate during winter was inversely related to the log of
population density. This occurred early in the non-breeding period just after
the mice reached their seasonal peak population density in autumn. This
result is consistent with mortality from starvation following the depletion
of summer foods. We did not find a significant relationship between the
population9s growth rate and the population9s density at any other time.
DAVIES, KENDI F.,1,* PETER L. CHESSON,2 SUSAN HARRISON1 and
BRIAN D. INOUYE.1,3 1 Department of Environmental Science and Policy,
and Center for Population Biology, University of California, Davis, CA;
2
Section of Evolution and Ecology, and Center for Population Biology,
University of California, Davis, CA; 3 Department of Biological Science,
Florida State University, Tallahassee, FL. Scale matters: How spatial
scale alters the diversity-invasibility relationship.
Elton suggested that diverse communities should be less susceptible to
invasion. In agreement, models and experiments at small spatial scales have
tended to show a negative relationship between native diversity and the
number of invaders. In contrast, studies at large spatial scales have tended
to find positive relationships between native and exotic diversity. This paradox is likely the product of different processes operating at different spatial scales: interactions between species at small scales and environmental
constraints at large scales. Shea and Chesson (2002) formalized this with
a conceptual model in which the negative slopes of the relationships be-

Abstracts

81

tween mean native and mean exotic richness for multiple sites at local
scales, form a cloud of points that make up the relationship between mean
native and exotic richness at a larger spatial scale. We point out that most
empirical studies at larger scales contrast the total rather than the mean
native and exotic richness. Thus there is a mismatch between an elegant
concept and empirical practice. We show that if beta diversity is not constant, the slope of the relationship between mean native and exotic richness
is not the same as the slope of the relationship between total native and
exotic richness. As a case study, we examined the relationship of native
and exotic diversity for four nested spatial scales in a California grassland
community. For means, the relationship between native and exotic richness
was negative for all four scales. However, for totals, the relationships at
the two smaller scales were negative but the relationships at the larger
scales were positive. In the literature, nested spatial data have been treated
in many different ways, often with the wrong approach used to address the
intended hypothesis. To test hypotheses that match processes to spatial
scale, we need to specify statistical models that properly partition processes
to the relevant scales.
DAVIS, ANDREW K.,* SONIA M. ALTIZER and NICHOLAS VITONE.
Dept. of Environmental Studies, Emory University, Suite E510, 400 Dowman Dr., Atlanta, GA, USA. Geographic variation among monarch butterfly populations: Migratory behavior and the evolution of wing morphology.
Native and introduced monarchs populate islands and continents worldwide, including North and South America to Hawaii, Australia, and New
Zealand. In parts of North America and Australia, monarchs undergo an
annual migration (traveling up to 2000 km to overwintering sites), whereas
monarchs in resident populations breed continuously and do not migrate.
For decades, scientists have assumed that monarchs from these different
regions were genetically distinct, yet almost no empirical studies have addressed this issue directly. We examined geographic variation in wing morphology among monarch butterfly populations in 2 migratory and 2 nonmigratory populations in N. America. We used both wild-caught monarchs
and captive-reared progeny from eastern N. America (migratory), western
N. America (migratory), southern Florida (resident), and Hawaii (resident).
We scanned the forewings of monarchs from each of these populations and
quantified size, color and shape characteristics using image analysis software. Based on comparisons of migratory and non-migratory bird species,
we predicted that monarchs from migratory populations will have more
sharply angled and more pointed forewings, while non-migratory populations would have more rounded wings. Consistent with our predictions,
monarchs from resident populations had shorter forewings (and smaller
wing areas) than monarchs from migratory populations. We further detected
significant population-level differences in wing shape, with the highly migratory eastern monarchs having the most angular and pointed wing design,
similar to that found in migratory bird species. Finally, we found that
monarchs from both resident populations had significantly darker forewings
than monarchs from migratory populations. Comparison of wild and captive-reared butterflies indicates that these differences are not solely environmentally driven and are heritable between generations. This study increases our understanding of the genetic basis of population divergence
among insect populations separated by geographic barriers or migratory
divides.
DAVIS, DAYTON K.,1,2,* BOB PARMENTER2 and RUDY BUENO.3
1
Unversity of New Mexico, Albuquerque, New Mexico; 2 University of
New Mexico, Albuquerque, New Mexico; 3 Albuquerque Bio-Diseases
Management, Albuquerque, New Mexico. The geographic disturbution
and temporal behavior of various mosquito species In New Mexico.
Due to the recent impact of vector borne diseases, such as the West Nile
Virus, across the nation, stem the emphasis for a better understanding of
the geographical and temporal behavior of various mosquito species. Recently we have started investigating the geographic distribution of mosquitoes, species and number, wise, along the Rio Grande valley in Bernalillo County. There is a correlation between the different habitats and the
presents of various amounts of mosquito species present in the different
areas.

82

Abstracts

DAVIS, HEATHER G.* University of California, Davis, hgdavis@

ucdavis.edu. Evolution of self-compatibility in an invasive plant is not
suppressed by inbreeding depression.
This study compared patterns of self-compatibility and rates of inbreeding
depression between the self-incompatible, perennial Spartina alterniflora
from the Atlantic coast of North America with an invading, inbred, population on the Pacific coast. Invasive plants were more self-compatible with
twice the self-pollinated seedset (50%) than plants from the native range.
Outcrossing with pollen from either the Atlantic natives or Pacific invaders
produced a more than 60% increase in seedset over selfing rates for the
native plants but no differences in the invasive plants. Native selfed seed
germinated at only 75% the rate of those from the Pacific in preliminary
data analysis. Invasive plant seeds outcrossed with invasive pollen had
twice the germination rate (70%) as the outcrossed seeds of all other combinations. This caused a higher measure of inbreeding depression for invasive plants (0.4) compared to native (0.07) despite the selfed invasives
higher germination rate. To investigate the mechanism of change in the
invaders self-compatibility, I contrasted plants from multiple invasive subpopulations representing the time since colonization (old . 60 yrs; intermediate 10-60 yrs; young , 10 yrs), equivalent to plant age, with the
younger the plant the more generations there were since initial colonization.
Young plants had twice the seedset when selfed, at 40%, than old plants.
Inbreeding depression of the young plants (0.2) was nearly half that of the
old plants. These results suggest the exotic Pacific population, that at the
low densities at the leading edge of the invasion suffers from greatly reduced fecundity, has evolved from the predominantly self-incompatible
state of the dense, competitive marshes on the Atlantic coast. Extreme
inbreeding has not hindered this invaders developing self-compatibility
since the inbreeding depression is not large and is decreasing over generations.
DAVIS, MARK A.* Macalester College, Saint Paul, MN. C3 grasses replace C4 grasses and reduce oak establishment in the prairie-forest
border.
As a result of changing land-use patterns and introductions of new species,
C3 grasses have replaced C4 grasses in many grasslands along the North
American prairie-forest border. Since C3 grasses are generally not as well
adapted to dry conditions, the replacement of C4 grasses by C3 grasses
may alter the competitive dynamics between grasses and woody plant seedlings and hence may affect the establishment success of woody plants. To
test this hypothesis, a multi-factorial field experiment was conducted in
which water, nitrogen, and grass functional group were manipulated in a
study of germination, survival, and growth of oak seedlings. The experimental design also permitted the separate quantification of root and shoot
competition by the grasses. The study showed a sharp reduction in oak
seedling survival under drought conditions, and to a lesser degree under
medium water conditions, for seedlings growing in C3 grasses. Analyses
of root and shoot competition showed that the cause for the reduced survival in C3 plots was due primarily to underground processes. These findings confirmed the initial hypothesis that replacement of C4 grasses by C3
grasses would reduce the establishment success of woody plants. However,
the explanation for the reduced oak seedling survival in C3 grasses is not
clear since soil water levels (measured weekly) did not differ between C3
and C4 plots and other resource levels measured (light and NO3 and NH4)
were higher in C3 plots. It is possible that the differential survival patterns
could be due to short-term fluctuations soil water levels that were not detected in the weekly soil water measurements. Or, the differential survival
might be due to other underground processes, such as herbivory, allelopathy, or mycorrhizal interactions, that might have differed between the C3
and C4 plots. Whatever the cause, the results show that the replacement of
C4 grasses by introduced C3 grasses in water-limited environments may
alter successional patterns by delaying or impeding the conversion of grassdominated environments to woody-dominated systems.
DAVIS, MICHELLE L.* and JIM BERKSON. Virginia Polytechnic Institute and State University, Blacksburg, VA. Red Porgy population status
determination: Optimizing data during data-poor periods.
The majority of marine stock assessments rely on some combination of
fishery-dependent and fishery-independent data. However, when a mora-

torium is in effect, the flow of fishery-dependent data is interrupted. With

the elimination of what is often the dominant data set, fishery-independent
data may drive assessment efforts, potentially affecting the determination
of key population parameters such as the intrinsic rate of increase, carrying
capacity, and individual growth rates. Effective sampling strategies for fishery-independent data are essential, but seldom investigated. Off the coast
of the southeastern United States, one fish of concern is red porgy (Pagrus
pagrus, Family Sparidae), a protogynous reef fish important to both recreational and commercial fisheries. Stock assessments performed on this
species since 1985 have shown a population in drastic decline, and, as a
result, a number of management actions were put into place, including a
moratorium in 1999. In order to determine the potential effects on our
ability to estimate key population parameters, the most recent assessment
model (a forward-projecting age-structured model) was modified to investigate a number of scenarios, including the loss of commercial and recreational data during a moratorium and increased fishery-independent sample
sizes. The results of this work will provide a framework for determining
key population status parameters when important data components are lacking.
DAWSON, AMY C. and SHELLEY E. ARNOTT. Queens University,
Kingston, Ontario. The effects of intra-annual pH fluctuations on recovering zooplankton communities in Swan Lake, Sudbury, Ontario.
Very little is known about the role that environmental variability plays in
the recovery of damaged ecosystems. The environmental variability characterized by pH variability within one year can be very high in lakes recovering from historical acid deposition. Swan Lake, a small lake that was
historically acidified by mining-related sulphur dioxide emissions, is beginning to show chemical recovery (i.e. increased mean annual pH) in
response to reduced emissions. Despite improvements in water quality,
there has been little recovery of the biota. One explanation is that pH
fluctuations prevent re-colonization of sensitive species. These environmental fluctuations may play a significant role in the recovery of species
that have been extirpated by historic stresses. To test whether or not short
term pH fluctuations influence the ability of a zooplankton community to
recover from acidification, pH levels were experimentally manipulated
within large epilimnetic enclosures (2800L). A two-factor experimental design was established to test the effects of small magnitude (pH fluctuation
from 6.5 to 5.8) versus large magnitude (pH fluctuation from 6.5 to 5.0)
pulses and short frequency (3-day) versus long frequency (9-day) pH pulses. Several species showed decreased abundance for the large magnitude
pulse (lowered to 5.0), when compared to the small pulse (lowered to 5.8),
but the frequency of the fluctuation had no effect. Other species showed a
similar response to magnitude, but also responded positively to the frequency of the pulse (i.e. higher abundance in 9-day treatments). Finally,
pulse magnitude and frequency had an interactive effect on some species.
These different species responses to short term pH fluctuations suggest that,
in lakes recovering from acidification, this variability may influence the
recovery trajectory of individual species, shaping zooplankton community
dynamics and ultimately recovery.
DE LUCIA, EVAN H.1,* and WILLIAM H. SCHLESINGER.2,* 1 University of Illinois, Department of Plant Biology, 265 Morrill Hall, Urbana, IL,
USA; 2 Duke University, Nicholas School of the Environment and Earth
Sciences, A 237 Levine Science Research Center, Durham, NC, USA. Respiratory control of the carbon cycle in a changing environment: synthesis and discussion.
Oxidative respiration from terrestrial ecosystems releases approximately
120 gigatons of carbon annually to the atmosphere, roughly 20 times the
amount from the combustion of fossil fuels. The difference between photosynthesis and the sum of respiratory processes determines the rate of
ecosystem carbon storage. For example, in a young pine forest in the Piedmont of North Carolina, autotrophic and heterotrophic respiration were
1704 and 216 gC/m2/y, respectively, or 71% and 9% of gross primary
production. After respiratory losses, the rate of carbon accumulation in this
forest was approximately 428 gC/m2/y. Fundamental questions remain
about how respiration will respond to the components of global change
and about the effectiveness of current methodologies for estimating respiration at different spatial and temporal scales. In this symposium we will

discuss the relationship between respiration and increasing levels of carbon

dioxide in the atmosphere. We will critically examine the potential limitations of extrapolating rate-based measurements to total ecosystem respiration. We will also discuss isotopic methods for partitioning respiratory
fluxes between autotrophic and heterotrophic components and evaluate the
importance of mean residence time as a control of carbon cycling in the
soil. This symposium will contribute to a new vision for improving estimates of ecosystem respiration.
DEATON, RAELYNN,* MELODY A. BROOKS and EDIE MARSHMATTHEWS. University of Oklahoma, Norman, Oklahoma. Effects of
food availability on embryo mass and female condition in a livebearing
fish.
Theory suggests that maternal provisioning to embryos should be influenced by resource availability. We examined effects of food availability on
embryo mass and female body condition in the largespring gambusia, Gambusia geiseri. This study is part of a larger project investigating factors
affecting matrotrophy (mother-to-embryo transfer of nutrients) in live-bearing fishes. Fish were collected from Anson Spring, TX and returned to a
greenhouse laboratory at the University of Oklahoma Aquatics Research
Facility. Females were randomly assigned to groups of 15 individuals in
four feeding treatments, with four replicates per treatment. Feeding treatments were: once every three days, once every two days, once a day, and
twice a day. At the end of 30 days, females were euthanized and dissected.
Embryos were removed from the ovary, dried and weighed. Female condition was assayed by extraction of fat with multiple washes of petroleum
ether. Body condition varied significantly among treatments: females in
treatments with low resources weighed less and were in poorer body condition. Body condition did not affect mean embryo mass. Further, embryo
mass did not decline with developmental stage, suggesting that maternal
transfer of nutrients was sufficient to counter loss of embryo mass due to
yolk metabolism. These results suggest that decreased resources did not
compromise matrotrophy, despite a clear reduction in female body condition.
DEBELJAK, MARKO.* Nova Gorica Polytechnic, marko.debeljak@
p-ng.si, Nova Gorica, Slovenia, Slovenia. Exergy as indicator of ecosystems integrity.
Environmental sciences and sustainable management with renewable resources require development of integrated ecosystem theory which can be
applied to explain scientifically our observations of nature by few basic
natural laws. Comprehensive thermodynamic hypothesis about the processes of ecosystems growth and development can make significant contribution to this attempt. The research has tested the ability of exergy indicators to assess forest ecosystems development cycle consist of exploitation, conservation, destruction and renewal development stags. Exergy
dissipation was calculated from data obtained with thermal remote sensing
of studied plots. Calculation of exergy storage based on genome size of
vascular plants of particular development stage. The results have shown
that exergy dissipation successfully indicates differences between initial
development stages, while exergy storage indicates differences between
mature stages of forest ecosystems development. According to the results,
new exergy indicator was proposed: maximum dissipation specific storage
of exergy. It is combination of exergy dissipation and exergy storage and
gives thermodynamic explanation for complete cycle of forest ecosystem
development. Results have been compared with the results of parallel conducted biometric research of the same study plots. The outcome showed
supplementing of both results. This was used for evaluation of tentative
ecological low of thermodynamic formulated thus: If a system receives a
through flow of exergy the system will utilise this exergy flow to move
away from thermodynamic equilibrium, and if more then one pathway is
offered to move away from thermodynamic equilibrium, the one yielding
most storage exergy by the prevailing conditions will have a propensity to
be selected. The rigorous examinations of the results have proofed the
existence of proposed law. Research has made a new contribution toward
formulation of ecosystem theory and integrated development of sustainable
management with renewable resources.

Abstracts

83

DEFOREST, JARED L.,1,* DONALD R. ZAK1 and KURT S. PREGITZER.2 1 University of Michigan, School of Natural Resources & Environment, Ann Arbor, MI; 2 Michigan Technological University, School of Forest Resources and Environmental Science, Houghton, MI. Chronic NO3deposition and lignin degradation in northern hardwood forests.
Human activity has globally doubled N input to terrestrial ecosystems. We
reasoned that chronic N additions will alter the flow of C through the

84

Abstracts

A
W

There have been many efforts recently to connect the work of scientists
and educators. This has become policy at the National Science Foundation
and at many other agencies that fund scientific research. Most often these
efforts are cast as social outreach activities where the knowledge and expertise of scientists is to be shared with the public. Much of this outreach
is focused specifically on the K-12 educational experience, with a hope of
improved teacher education, higher quality instructional materials, and
more effective classroom activities, but higher education can benefit from
these efforts as well. However, little thought has gone into how, exactly,
this translation from scientific research to the schools, higher education, or
the general public is to happen. In this talk I will explore some of the ways
that science researchers can collaborate with K-12 educators to help improve science education and how this approach can be adapted for other
audiences as well. A key strategy is through focused discussions of the
content that the students are expected to learn. At Project 2061 we have
developed what we call strand maps, which show the conceptual linkages
between sets of content-based learning goals. Through discussions of this
content and these linkages, scientists can come to understand the learning
expectations that educators have for K-12 students, their developmental
trajectories, and the level of preparation they should have when they enter
higher education. At the same time, the scientists can learn about a goalsbased model of content instruction that can be adapted for use in higher
education or the general public. A second way for scientists to collaborate
with K-12 educators is for them to offer phenomena that they work with
as possible new contexts for teaching important concepts, and to offer
visual models and representations that they believe can be used to clarify
difficult concepts for students. When scientists think in terms of the specific
learning goals of their audience, then their outreach efforts can be more
focused and effective. The purpose of this talk is to discuss some of the
ways to accomplish this.

Conservation biology has long searched for a predictive framework for

extinction. This study examines one novel approach, behavioral plasticity.
Pursuant to a hypothesis of behavioral plasticity, the degree of phylogenetic
constraint on an organism9s behavior determines, in part, its response to
environmental change. Species with a higher degree of plasticity would be
characterized by higher persistence during periods of change. These predictions were tested on 12 lemur species in habitat fragments in southeastern Madagascar. First, the degree of behavioral plasticity for each species was estimated, and tested as a predictor of fragment incidence. Further,
the incidence of lemur predators, the fragment9s botanical makeup, the
availability of lemur foods, historical species incidence, and hunting were
tested as alternative causal agents. Species were found to strongly differ in
their response to fragmentation. These responses were non-random, highly
nested, and reflected the relative degree of plasticity for a species. Second,
if behavioral plasticity was the primary mechanism behind the differential
persistence of these species in fragments, one would expect that behavior
would change for fragment populations in a manner that mirrors the degree
of plasticity of a species. To adjust to the reduced levels of resources within
fragments, those species with a high degree of behavioral plasticity were
predicted to increase their foraging effort. Conversely, those species estimated to be more behaviorally constrained were predicted to have no
change in foraging effort, but consequently to suffer physiological and
demographic declines. These predictions were tested for two of the 12
species, Eulemur fulvus rufus, and Propithecus diadema edwardsi, which
exhibited greater and lesser degrees of plasticity, respectively. In fragmented habitats, Propithecus populations did not modify their foraging
effort, and consequently declined significantly in abundance and body
mass. Eulemur populations, in contrast, increased their home range and in
abundance, and maintained body mass.

DEBOER, GEORGE E.* Education Linked to Scientific Research: the

Education Perspective.

DEHGAN, ALEX O.1,2,* 1 The University of Chicago, Committee on Evolutionary Biology, 1025 E. 57th Street, CH402, Chicago, IL, USA; 2 The
Field Museum of Natural History, Division of Mammals, 1200 South Lake
Shore Drive, Chicago, IL, USA. Predicting the extinction proclivity of
rainforest lemurs in S.E. Madagascar during fragmentation through a
hypothesis of behavioral plasticity.

IT

Microcrustaceans are a diverse and abundant component of the animal

communities of isolated depressional wetland ponds on the southeastern
Coastal Plain. They play an important role in the trophic structure. Some
species are temporary wetland pond specialists, thus they may be useful
indicator species. Twenty isolated wetland ponds are part of the Carolina
Bay Restoration Project on the Savannah River Site (SRS) in South Carolina. All of these ponds had been ditched for agriculture, well before 1951.
Since 1951, all have become forested, mainly with bottomland hardwoods
and pine. The natural hydrology was restored in 16 of the ponds by plugging their ditches during 2000. These ponds were also logged. Half were
re-planted with herbaceous vegetation, and half with pond cypress and
black gum. The remaining four remained untouched to act as controls. We
sampled the ponds for three yrs before the ditches were plugged, and we
gave continued sampling for .2 yr post-treatment to characterize microcrustacean responses. Most of the pre-treatment communities supported
fairly rich microcrustacean assemblages. Species richness was a function
of hydroperiod length. No significant changes in species richness were
observed in the first 2 years of post-treatment, indicating that the treatments
had no detrimental effects. As suggested by a previous restoration of another SRS pond, we predict that new species will colonize slowly over at
least the next 5-6 yrs. Analysis of data from a set of 88 SRS reference
ponds suggests that the ponds with herbaceous plantings will have richer
assemblages than those planted with trees. Also, if the restoration eventually produces permanent inundation, temporary pond specialists, such as
the phyllopodous branchiopods, will be lost.

microbial foodweb by inhibiting the activity of lignin-degrading soil fungi,

which, in turn, control ecosystem-level patterns of C and N cycling. We
tested our hypothesis in a mature northern hardwood ecosystem in Michigan which has received experimental N additions (30 kg NO3--N ha-1 y-1)
since 1994. In a laboratory study, we amended soils with 13C labeled vanillin, a product of lignin degradation, and tracked the flow of 13C through
the microbial community. In addition, we simultaneously measured phenol
oxidase activity, which is responsible for degrading lignin and vanillin.
Phenol oxidase activity was significantly suppressed by 133% in N amended soils. In addition, we found N additions increased the recovery of 13C
in fungal PLFA by 82%, but had no significant effect on recovery of 13C
in bacteria PLFA or microbial respiration. Results suggest that chronic N
additions suppressed lignin degradation, while it increased the ability of
soil fungi to metabolize the products of lignin degradation.

DEBIASE, ADRIENNE E.* and BARBARA E. TAYLOR. Savannah River

Ecology Laboratory, University of Georgia, Aiken, SC, USA. Microcrustacean responses to hydrologic restoration of isolated wetland ponds.

DEL GROSSO, STEPHEN J.,1,* ARVIN R. MOSIER,2 DENNIS S. OJIMA,1 WILLIAM J. PARTON1 and CINDY KEOUGH.1 1 Colorado State
University, Fort Collins, CO; 2 USDA - Agricultural Research Service, Fort
Collins, CO. Using the DAYCENT ecosystem model to assess greenhouse gas emissions from agricultural systems in the US.
As a signatory of the Framework Convention on Climate Change the US
is required to conduct annual greenhouse gas (GHG) inventories. Agriculture is not a strong source of CO2 emissions, but is responsible for a significant proportion of anthropogenic emissions of N2O and CH4. Although
the atmospheric concentrations of N2O and CH4 are small compared to
CO2, they have a higher global warming potential than CO2 on a per molecule basis. Currently, GHG emissions from the agricultural sector are estimated using the Intergovernmental Panel on Climate Change (IPCC)
emission factors. For example, 1.25% of total N fertilizer applied to crops
is assumed to be lost as N2O emissions. To improve GHG flux estimates,
we combined climate, soils, and land management data from various sources to make input files required by DAYCENT. DAYCENT is an ecosystem
model used to investigate the effects of changes in land use and climate

on C and N fluxes. The US was divided into 63 regions and the most
common agricultural practice within each region was represented in the
model. Direct N2O emissions from soil as well as indirect N2O emissions
associated with NO3 leaching, NH3 volatilization, and NOx emissions were
summed for each region. Simulated crop yields, N gas emissions, and NO3
leaching were compared with field data to test model performance. Comparisons of DAYCENT simulated N2O emissions with site level data show
that DAYCENT matches observations better than N2O emissions calculated
using the IPCC method for most sites tested. Model results suggest that
corn/soy cropping in the Midwest stores 25g C m-2 annually and wheat
cropping stores about half as much C. However, the C equivalents of N2O
emissions are higher for corn/soybean rotations (20g CO2-C m-2 yr-1) than
wheat cropping (15g CO2-C m-2 yr-1).
DELL, ANTHONY I.* Department of Zoology and Tropical Ecology,
James Cook University, Townsville, Queensland, Australia. Do aquatic
and terrestrial food webs really differ?
While comparisons of empirical food webs from a diversity of environments around the globe have revealed general consistencies in web properties, there is recognition that webs from similar environments may be
more structurally alike. At present, however, there is little understanding
of what these patterns might exactly be. This failure to clearly identify
habitat-specific web properties, or indeed disprove their existence, has been
most strongly hindered by the ways in which web comparisons have been
made. Published webs vary so greatly in aspects of data collection and web
construction that comparisons of multiple empirical webs are subject to
many confounding sources of variation that may mask or alter real patterns.
Experimental investigation of a series of drying intermittent pool beds allowed me to closely monitor changes in web structure in a community that
shifted from being composed of aquatic species to one that was composed
of terrestrial species. Comparison of time-specific webs throughout this
transition were largely immune to the problems that have troubled previous
workers as all webs occurred in the same space, at about the same time,
and were equal in all other aspects of data collection and web construction.
There is general recognition that theory is fast outstripping data in food
web research. The results I present here represent a significant advance in
our understanding of the underlying structure of biological communities
from aquatic and terrestrial environments.
DELPH, LYNDA F.* Indiana University, ldelph@indiana.edu, Bloomington, IN. Elaborate mating displays and the physiological costs they
incur.
Genetically based variation exists in the number and size of flowers produced among populations of the dioecious plant Silene latifolia, which also
exhibits strong sexual dimorphism in floral traits (males make many, small
flowers compared to females). Artificial selection experiments have revealed that flower size/number is strongly genetically correlated with vegetative and physiological traits. Direct selection on floral display comes
with correlated responses for a host of other traits, including leaf traits,
overall plant size, and metabolism. For example, plants that make many,
small flowers are smaller and produce smaller, thinner leaves with higher
rates of photosynthesis and respiration than do lines selected for making
few, large flowers. These results suggest that selection on floral traits cannot
be independently selected without also indirectly selecting for physiological
traits, and vice versa. Hence, the potential exists for the abiotic environment
to constrain selection on floral traits via mating success or pollinator-mediated selection. Clearly, an understanding of why populations of this species vary in their floral traits requires knowledge of the genetic correlations
among traits.
DENNO, ROBERT F.,1,* CLAUDIO GRATTON,2 HARTMUT G. DOBEL1
and DEBORAH L. FINKE.1 1 Department of Entomology, University of
Maryland, College Park, Maryland, USA; 2 Department of Entomology,
University of Wisconsin, Madison, Wisconsin, USA. Predation risk influences relative strength of top-down and bottom-up impacts in phytophagous insects.
Elucidating the relative strength of top-down and bottom-up forces in communities of phytophagus insects has been a major historical focus. Current

consensus is that both forces play a role, but it is poorly known if these
forces act differently on herbivores in the same community. Using manipulative experiments with an assemblage of sap-feeding phytophagous insects (six species of planthoppers, leafhoppers, and heteropteran bugs) inhabiting intertidal Spartina marshes, we examined the association between
herbivore behavior, risk of predation, and ultimately the relative impact of
top-down (wolf spider predation) and bottom-up factors (host-plant nutrition) on the population density of each sap-feeding herbivore. Bottom-up
effects prevailed in this community, whereby the density of all six sapfeeders increased when the nitrogen content of their Spartina host plant
was elevated. By contrast, wolf-spider addition significantly suppressed
populations of only Prokelisia planthoppers, and had little impact on the
other four sap-feeder species in the community. Functional response experiments and behavioral studies revealed that certain species (Prokelisia
planthoppers) were at much higher risk of attack by wolf spiders than other
sap-feeders in the assemblage, and that risk of predation was associated
with the particular escape-defensive behavior of a species. Moreover, risk
of spider predation was linked to the strength of top-down impacts in the
field, because species with ineffective escape behaviors and a high risk of
spider attack (Prokelisia planthoppers) were the only sap-feeders whose
populations were suppressed by spider predation in the field. Thus, specific
behavioral characteristics of the sap-feeders on Spartina influenced risk of
predation and the relative strength of top-down and bottom-up impacts on
their population dynamics. These results call into question the overall pervasiveness of top-down forces and underscore the primacy of basal resources in structuring this community of phytophagous insects.
DENSLOW, JULIE S.,1 CURT C. DAEHLER,2 SHAHIN ANSARI2 and
HUANG-CHI KUO.2 1 USDA Forest Service, Hilo, HI; 2 University of Hawaii, Honolulu, HI. Predicting pest plants in Hawaii and other tropical
Pacific islands.
More than 1000 species of non-native plants have naturalized in the Hawaiian Islands. While only a small proportion cause significant ecological
harm to native ecosystems, most that do were intentional introductions and
often are still in the commercial plant trade. We sought a mechanism to
rapidly and efficiently evaluate exotic species for potential invasiveness on
tropical islands. Oceanic islands are particularly vulnerable to establishment
of exotic pests and thus present a challenge to effective screening programs.
We adapted the Australian Weed Risk Assessment protocol for use in tropical high islands. This system uses 49 questions about plant invasive behavior where introduced elsewhere, life history traits, and climate suitability to develop an index of invasiveness. We evaluated 172 non-native species long in the trade in Hawaii and the Pacific and selected to represent
a wide range of plant families and growth forms. Independently we solicited the expert evaluations of 25 botanists with extensive field experience
in the tropical Pacific. The system returned clear evaluations for 76% of
the species submitted. Addition of a secondary screen to evaluate undecided species increased the percent evaluated to 92%. Agreement with expert
opinion was high: 95% of major pest plants, 67% of minor pests, and 76%
of non-pests of native ecosystems were correctly identified. Information on
invasive status elsewhere was an important component of the screen, but
by itself identified only 81% of major pests. Invasive behavior in native
and managed ecosystems were significantly correlated. We suggest that this
system can be a useful tool to rapidly evaluate large numbers of actual and
potential introductions of plants for their potential pest status.
DEPRIEST, TIMOTHY T.* Cornell University, Ithaca, NY. High school
students as forest ecologists: Learning the nature of science and the
science of nature through inquiry.
Scientists have the capacity to play a key role in the education of secondary
school students through engaging them in research activities. However, to
be effective at meeting new standards in science education, the learning
experience must include more than following a research protocol. As a
participant in a NSF program that places graduate students in school settings to teach science, I attempted to integrate my own experiences in forest
ecology research into an educational setting for the purpose of meeting
current reform standards, which emphasize learning the nature of science
(NOS) and skills in scientific inquiry. In one project we carried out investigations into local old-growth forests through a structured inquiry in which

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I defined the class research goal of establishing a null model for native
forest ecosystems based on specific biological (tree size class and species
composition) and physical (amount of course woody debris and leaf litter
depth) characteristics. The curriculum then follows a pedagogical model
that reflects a progression from structured inquiry to guided, and eventually
open-inquiry through a process of shifting the identification of the research
question and research process from the instructor to the student. In another
project, we applied a research protocol to investigate the genetic variability
of sugar levels in the sap of maple trees in an active sugarbush. My goal
for this project was to investigate methods of incorporating aspects of the
NOS related to the research process into a structured inquiry model. Learning and teaching the nature of science, as reflected in reform literature,
took place through reflective discussion of the NOS aspects experienced
by the students in the research process. From these experiences I learned
that attempting to meet science education standards through facilitating
student research involved not only an integration of skills in ecological
research processes, but also a critical reflection of my own conceptions of
the NOS.
DERMODY, ORLA, STEPHEN P. LONG and EVAN H. DELUCIA. University of Illinois, Urbana -Champaign, Illinois. Canopy development of
soybeans grown in elevated [CO2 ] and [O3].
Canopies are the interface for energy, CO2 and H2 O exchange between
the atmosphere and the terrestrial biosphere. Despite their importance few
studies address how canopy structure and function may be impacted by
altered tropospheric chemistry. Increased photosynthetic efficiency under
elevated [CO 2] may lower the light compensation point of shade leaves.
Longer retention of these leaves may then increase canopy size under elevated [CO2]. Rapid senescence in elevated tropospheric [O3] may however
reduce canopy size. In a soybean (Glycine max) free air concentration enrichment system (SOYFACE), leaf area index (LAI) was measured in plots
(252m diameter) exposed to ambient air (;370ml l 21) and elevated
(;550ml l21) [CO2]. LAI was also measured in plots at 1.2*ambient [O3].
LAI was consistently greater in soybeans grown at 550ml l21 [CO2]. Measurements of the LAI profile and canopy depth indicated that leaves were
not retained deeper in the canopy at 550ml l21 [CO2]. Maximum quantum
yield (Fmax) was greater for soybeans exposed to 550ml l-1 [CO2]. Cumulative damage by [O3] caused early and rapid senescence, which decreased LAI. No effects were detected on Fmax. Remobilisation of nitrogen from senescing leaves to the upper canopy may counteract retention
of shade leaves under elevated [CO2]. Increased senescence in elevated [O3]
may however offset any gains in productivity predicted with increased tropospheric [CO2].
DERR, KATHARYN D.* and STEVEN L. MATZNER. Effects of fire
intensity on understory vegetation diversity in the Black Hills of South
Dakota.
Fire has become an increasingly important issue in western states because
of the frequency and severity of many recent wildfires. Years of fire suppression policy have resulted in higher fuel loads that cause fires to burn
more intensely. This paper presents a study on the effects of different fire
intensities on the understory vegetation in a Pinus ponderosa community
at the Jasper Burn Area in the Black Hills of South Dakota. It was hypothesized that high intensity fires would be more damaging to forest understory recovery, while low intensity fires might lead to increased diversity by stimulating the growth of more fire tolerant native species. It was
also hypothesized that the low intensity burn sites would have fewer invasive species compared to the high intensity burn or unburned sites. The
results however did not conform to our original hypotheses. Species richness and diversity were not significantly different between high and low
intensity burn sites. In fact, the unburned sites had significantly higher
species richness and diversity compared to either the low or high intensity
burns. Analysis of individual species changes revealed that fire of any
intensity resulted in the loss of fire intolerant species, but that fire did not
appear to be stimulating growth of fire tolerant native species. The absence
of an increase in fire tolerant natives may be due to a lack of seed source
or individuals to resprout. Recovery of natives in this system may occur
more slowly than originally thought and may require additional burn events
before recovery is complete.

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DERRICKSON, ELISSA M.* Loyola College, ederrickson@loyola.edu,

Baltimore, MD. Four years to develop a scientist: Accelerating an understanding of the process of science in undergraduate biology majors.
For the past several years the Biology Department at Loyola College has
been revising the curriculum for our undergraduate majors. One of our
goals has been to improve the ways that students learn the process of
science. We constructed a sequence of three courses that build skills and
use the inquiry method to facilitate student growth as scientists. As students
progress through these courses, they acquire increasingly specialized
knowledge. We have replaced our introductory labs for first year students
with a 3-credit course that introduces students to the process of science.
All majors take a second course that emphasizes experimental design, analysis, and scientific writing. Finally, upper level students choose a course
in experimental methods within a field in biology. The upper-level course
uses student-designed, multi-week experiments to learn the methodology
of a particular sub-discipline of biology. I am constructing a methods
course in ecology that ran for the first time last fall. I will describe the
structure of this course, how this course fits into our overall curriculum,
and some of the problems that I experienced. I will also discuss how our
experiences might be translated into changes at other institutions.
DESROCHERS, RACHELLE E.* Laurentian University, Sudbury, Ontario, Canada. Partititioning the components of diversity along ecological
perturbation gradients.
Moments of Renyi9s generalized entropy, such as Shannon diversity and
Simpson9s index, have long been used to quantifying diversity and have
found use in the assessment of ecological recovery. Similar measures, Kolmogorov complexity, quadratic entropy and taxonomic entropy, have been
introduced to expand upon Renyi9s generalized entropy and incorporate
relevant properties such as structure and taxonomic relatedness. These measures were applied to vegetation data from recovering plant communities
along a gradient of perturbation by industrial pollution in northern Ontario,
Canada. Perturbation was most intense near the source, the decommissioned nickel and copper smelters, and decrease with distance. Diversity
should be greater in communities subjected to less intense perturbation and
represent a more advanced level of recovery. As hypothesized, Shannon
diversity increased with increasing distance from the smelters. Despite the
incorporation of structure or taxonomic relatedness, the three new measures
showed similar patterns of increase thereby providing little additional insight as to the nature of community recovery. Each measure can be partitioned into components representing a moment of Renyi9s generalized entropy and the structural or taxonomic information gain. Interestingly it is
the dynamics of the partitioned components of each measure that best characterize the nature of recovery along the gradient. The allocation of information into these components is also better able to reveal differences in
recovery among the naturally recovering communities and rehabilitated
communities found along the gradient. To determine if this is a local phenomenon, the analysis was applied to similar vegetation data from southwestern Finland near a still active nickel and copper smelter complex.
While the pattern of diversity along the gradient and the allocation of
information differ from the Canadian case, the partitioned components of
diversity remain key in understanding the nature of recovery along the
gradient.
DEVANNA, KRISTEN,* NICHOLE RUBECK-SCHURTZ, AMY L.
DOWNING and LAURA TUHELA-REUNING. Department of Zoology,
Ohio Wesleyan University, Delaware, OH. Response of pond biodiversity
and ecosystem processes to the insecticide Sevin.
Pesticides introduced on land are often carried into aquatic systems becoming chemical pollutants. In this study we explored the sensitivity of freshwater pond communities and ecosystem processes to Sevin, a widely used
insecticide that is particularly toxic to aquatic invertebrates. We constructed
artificial pond mesocosms adjacent to a local pond on the Ohio Wesleyan
Kraus Wilderness Preserve. The mesocosms were filled with a subset of
the local pond community including bacteria, phytoplankton, zooplankton,
and a subset of the macroinvertebrate community. After 6 weeks, Sevin
was added to the mesocosms to obtain carbaryl concentrations (the active
ingredient in Sevin) of 0.0 mg/ml 0.1 mg/ml, 1.0 mg/ml, and 20 mg/ml. To

determine the response of the pond community, we sampled zooplankton,

periphyton, phytoplankton, bacterial and macroinvertebrate communities
for 6 weeks following the carbaryl addition. We also monitored ecosystem
productivity and respiration rates. HPLC analysis revealed that our initial
target treatment levels were reached, but by day 60 of the experiment
carbaryl levels in the water had decreased to near zero for all treatment
levels. Macroinvertebrate diversity was not affected by the carbaryl disturbance, while the zooplankton diversity declined with increasing carbaryl
concentration. After the addition of carbaryl, oxygen levels in the high
mesocosms decreased while the bacteria and periphyton biomass increased.
In summary, our results suggest that Sevin reduced the diversity of pond
communities, drastically increased bacterial abundance, and altered ecosystem productivity and respiration. Future work should explore how longterm, consistent exposure to aquatic pollutants affects aquatic biodiversity,
and how this may translate into long-term changes in ecosystem processing.
DEWALT, SAARA J.,1,* JULIE S. DENSLOW2 and JAMES L. HAMRICK.3 1 Louisiana State University, Baton Rouge, LA; 2 USDA Forest
Service, Hilo, HI; 3 University of Georgia, Athens, GA. Biomass allocation, growth, and photosynthesis of native and introduced genotypes
of an invasive tropical shrub.
Exotic pest plants often grow to greater stature, become more abundant,
and display increased shade tolerance in their introduced ranges than in
their native ranges. We tested the hypothesis that genetic differences in
growth, biomass allocation, or photosynthetic parameters between native
and introduced genotypes of Clidemia hirta (Melastomataceae) contribute
to differences in abundance and shade tolerance between its native and
introduced ranges. We conducted a common garden experiment in a greenhouse with C. hirta grown from seed collected from four populations in
part of its native range (Costa Rica) and four populations in part of its
introduced range (Hawaiian Islands). Six-month old seedlings were placed
in high (10.3-13.9 mol m-2 day1) or low (1.4-4.5 mol m-2 day-1) light treatments and grown for an additional six months. Our study provided little
evidence that Hawaiian genotypes of C. hirta differed genetically from
Costa Rican genotypes in ways that would contribute to the observed differences in habitat distribution and abundance. Hawaiian and Costa Rican
genotypes displayed no differences in relative growth rates, maximal photosynthetic rates, or specific leaf areas in either light treatment. Some of
the genetic differences that were apparent, such as greater allocation to
stems and leaf area relative to whole plant biomass in Costa Rican genotypes and greater allocation to roots in Hawaiian genotypes, were contrary
to predictions that genotypes from the introduced range would allocate
more biomass to growth and less to storage than those from the native
range. As predicted, however, Hawaiian genotypes allocated more biomass
to reproductive parts than Costa Rican genotypes in the high light environment. We conclude that environmental differences, such as pest loads,
between the native and introduced ranges likely play a greater role than
genetic shifts in resource use or allocation in determining habitat distribution and abundance for this species.
DIAZ, SANDRA.* Functional biodiversity and environmental filtering
in different floras: a conceptual framework.
Functional diversity (FD) is defined as the value and range of plant functional traits present in a given community. Despite FD being considered a
key factor influencing ecosystem functioning, most authors still use species
diversity (SD) as the only measure of diversity. As well as practical reasons, this is due to the fact that FD is often assumed to vary in direct
proportion to SD. Such an assumption, however, is inappropriate under
most field situations, where community assemblage occurs as the result of
the filtering action of environmental factors on a regional species pool. In
real ecosystems, FD and SD, although not strictly independent, are seldom
linearly related. I propose a conceptual model to predict the shape of the
relationship between FD and SD under different scenarios of species-pool
limitation and environmental filtering by climate and large-scale disturbance. On the basis of links between plant traits, environmental factors,
and ecosystem processes, the framework can also predict what combinations of key functional traits would be expected under different filtering
scenarios. I empirically test some of the predictions of the model by using
plant-trait databases from six regional floras from continents and islands,

representing different combinations of species-pool limitations and filtering

factors, and showing evolutionary histories of disturbance.
DICKMAN, ELIZABETH,* MICHAEL J. VANNI and WILLIAM H.
RENWICK. Miami University, Oxford, Ohio. Phytoplankton photosnthesis, primary production and nutrient limitation in reservoirs along a
gradient of watershed land use.
Land use can have many effects on aquatic ecosystems. However, little is
known about how lake phytoplankton respond to changes in watershed land
use. We investigated how reservoir phytoplankton photosynthetic parameters, primary production, and nutrient limitation status vary along a gradient
of watershed land use. Our seven reservoirs reside in watersheds that range
from mostly forested (80% forest, 13% agriculture) to highly agricultural
(89% agriculture, 9% forest). Urban areas represent only a small fraction
of land in all watersheds. We found that late summer primary production
(mg C/m2/d) was highly correlated with the percentage of watershed land
comprised of agriculture (r2.0.8). Maximal (light-saturated) photosynthetic
rate of phytoplankton was also correlated with percent agriculture, but the
relationship was much weaker than that between primary production and
percent agriculture. Phytoplankton were nutrient limited in all reservoirs.
Nitrogen (N) was the limiting nutrient is some lakes, while phosphorus (P)
was limiting in others; co-limitation by N and P was common. There was
no relationship between land use (percent agriculture in watersheds) and
either the identity of the limiting nutrient (N versus P), or the severity of
nutrient limitation. Our results suggest that agricultural activities in watersheds increase primary production, most likely by increasing nutrient subsidies. However, across the land use gradient, phytoplankton apparently
attain a biomass at which they exhaust resources, and thus become nutrient
limited.
DICKSON, TIMOTHY L.* and BRYAN L. FOSTER. University of Kansas, dicksont@ku.edu, Lawrence, KS. The control of plant species richness in relation to productivity: Seed vs. microsite limitation.
It has been hypothesized that plant species richness is controlled primarily
by microsite availability under highly productive conditions, but by seed
availability under less productive conditions. We tested this hypothesis with
a field experiment in northeastern Kansas grassland. In a 2 x 2 x 2 factorial
arrangement of 48, 2m x 2m plots we manipulated productivity (no fertilizer; 16g nitrogen/m2 year), seed availability (no seed addition; seeds of
40 species added), and water availability (no irrigation; plots irrigated every
4th dry day). We also clipped half of each plot to 3cm height in mid-June,
to yield a split plot design. Percent cover of all species was recorded in
the spring and fall, and results varied between the spring and fall. In the
spring, species richness was limited by microsite availability in the high
productivity plots and by seed availability in the low productivity plots
(added species had no significant effect on richness in high productivity
plots). In the fall, for both low and high productivity plots, species richness
was limited by microsite availability in the non-clipped plots (no effect of
seed addition) and by seed and microsite availability in the clipped plots
(seed addition, productivity, and irrigation all significantly affected species
richness). These results suggest that the relative importance of seed and
microsite limitation in controlling plant species richness may vary along a
productivity gradient and through time.
DIEHL, SEBASTIAN.* Ludwig-Maximilians-Universitaet, diehl@zi.
biologie.uni-muenchen.de, Munich, Germany. Enrichment with light: Effects of mixing depth on herbivore-producer dynamics in a planktonic
system.
Enrichment is defined as an increase in supply with production-limiting
resources. Studies of enrichment have almost exclusively focussed on enrichment with nutrients, whereas enrichment with light (increased light supply) has received little consideration. In aquatic systems, increased light
supply may shift the elemental composition of algal biomass towards low
nutritional quality [high carbon to phosphorous (C:P) ratio] for herbivores.
Under laboratory conditions, stimulation of algal production with increased
light may even reduce the growth rate of herbivores, if the light-induced
drop in algal quality outweighs the increase in algal biomass. Does this
seemingly paradoxical phenomenon occur under natural conditions? In

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planktonic systems, the majority of primary production occurs in the mixed

surface layer, and differences in the depth of the mixed layer create substantial variation in light supply among lakes. To explore the responses of
planktonic herbivore-producer dynamics to changes in the relative supply
of light and nutrients, we created an experimental gradient of mixing depths
(1-12m) in continuously mixed enclosures filled with nutrient-poor lake
water and stocked with an inoculum of herbivorous Daphnia. Average water temperature was around 138C and was very similar within and among
enclosures of different mixing depth. Throughout the 55-day experiment,
primary production decreased with mixing depth and was efficiently converted into Daphnia biomass at all depths. Consequently, Daphnia grazing
controled algal biomass at low levels (preventing the build-up of high algal
C:P ratios) and responded positively to increasing light supply (decreasing
mixing depth). Because algal food quality would be expected to be lowest
(C:P ratios highest) in shallow, clear, nutrient-poor mixed layers (as used
in the experiment), the data suggest that natural levels of enrichment with
light may not have negative effects on grazers in a fully dynamic phytoplankton-zooplankton system.
DIEKMANN, LUCY O.* and DEBORAH LAWRENCE. University of
Virginia, Charlottesville, VA, USA. Changes in soil phosphorus following shifting cultivation in the Yucatan Peninsula.
Phosphorus is a limiting nutrient in the dry tropical forests of the Americas.
Following forest conversion, phosphorus impoverishment is a leading cause
of declining soil fertility. Typically reduced phosphorus availability results
from the transformation of available to unavailable phosphorus rather than
net losses from the system. To determine the cumulative effect of shifting
cultivation on soil phosphorus dynamics in a repeatedly disturbed ecosystem, we studied 24 sites that represent a cultivation history gradient around
El Refugio, an ejido in the southern Yucatan Peninsula. These sites ranged
in age from recently abandoned agricultural fields to mature forest (undisturbed for .50 yrs.) and had undergone from 0-6 cultivation-fallow cycles.
The cultivation history of each site was found through interviews with local
farmers. Using a sequential chemical extraction, we determined the size
and availability of soil phosphorus pools during shifting cultivation. Preliminary results in an ongoing study showed no significant difference in
total phosphorus between stands that had undergone one or two cultivation
cycles and were less than 6 yrs. old. However, the range of soil phosphorus
in these stands (0.13-0.15 mg P/g) is only 69% of total phosphorus in
mature forests. Our results suggest that repeated anthropogenic disturbance
in these forests has a significant effect on soil phosphorus. Phosphorus
transformations influence forest resilience following disturbance, making
an understanding of soil phosphorus dynamics in a habitually disturbed
ecosystem essential to developing sustainable agricultural and forestry
practices in the dry tropics.
DIEMER, KATHERINA E.1,* and JOHN E. FAUTH.2 1 College of Charleston, Charleston, SC, US; 2 College of Charleston, Charleston, SC, US. Impoundments on headwater streams: Do many small ponds have a large
cumulative effect on water quality?
Landowners often dam headwater streams to create ornamental or fishing
ponds on their property. These impoundments are unregulated in many
states, and their effects on water chemistry and the composition of the
stream biota are largely unknown. I am comparing water chemistry parameters (pH and nutrients) and distribution and abundance of amphibians, fish
macroinvertebrates in impounded and unimpounded streams, to determine
if if small ponds affect watershed dynamics. My sampling design permits
comparisons between both impounded and unimpounded streams, and between sections above and below ponds. My study addresses concerns of
private stakeholders about the effects of their ponds on stream quality, and
provides data for environmental scientists interested in the cumulative effects of small perturbations on larger watersheds. This study will assist
stakeholders in making educated decisions regarding watershed conservation, including headwater sections that are important for refugia for many
threatened and endangered species.
DIETZE, MICHAEL C.* Duke University, University Program in Ecology,
Durham, NC. North Atlantic hurricane disturbance: Current patterns
and climatic phases.
Hurricane force winds are a significant disturbance factor for much of eastern North America. Since the structure and composition of ecological com-

88

Abstracts

munities can be strongly impacted by changes in disturbance frequency

and magnitude, it is important to understand the spatial variability of hurricane impact, as well as how the pattern of disturbance varies with changes
in climate. To address this problem, hurricane return intervals were estimated at a 1/10th degree resolution for eastern North America based on
the historical hurricane record (NOAA HURDAT database, 1851-2002) and
a simple model of hurricane wind fields derived from the HURRECON
model. Return intervals were statified by El Nino-Southern Oscillation
(ENSO) phase, tropical Atlantic sea surface temperature (SST), and season.
El Nino years are associated with reduced hurricane activity (increased
return intervals) while La Nina years show increased hurricanes. Decreased
sea surface temperatures in the tropical North Atlantic are associated with
decreased hurricane activity. Neither ENSO nor SST are associated with
substantial shifts in the spatial pattern of hurricane impact. This suggests
that estimates of global changes in hurricane activity should scale well to
changes in regional activity. However, storm impacts do shift spatially
throughout the hurricane season. Early season storms have the broadest
spatial range. However, as the hurricane season progresses storm activity
is reduced in the western Gulf of Mexico and eastern U.S., becoming more
concentrated in Florida and the Caribbean. These seasonal shifts suggest
that most hurricane impacts in temperate deciduous forest will be before
leaf-fall, when disturbance will be largest. Finally, regional patterns of forest damage, based on the USFS Forest Inventory and Analysis, were compared to recent hurricane paths. There is a broad agreement between storm
intensity (wind speed) and severity (loss of basal area and biomass). Tree
damage also varies by species and increases with tree diameter.
DIEZ, JEFF M.* and H. RONALD PULLIAM. Institute of Ecology, University of Georgia, Athens, GA. The distribution and demography of a
terrestrial orchid across environmental gradients.
The geographic ranges of many species are characterized by strong gradients in abiotic conditions over multiple spatial and temporal scales. Understanding and accounting for the variability in ecological processes at
these various scales is important for making realistic predictions of species
dynamics and issues such as responses to climatic change. I am attempting
to understand how understory plant species respond to climatic and soil
gradients by studying the distribution and demography of a terrestrial orchid, Goodyera pubescens. Long-term study plots have been established
along a gradient from the southeastern Piedmont to the southern Appalachian mountains, encompassing a wide range of environmental conditions,
of which temperature, soil moisture, light levels, and soil nutrients are
monitored on each plot. I describe here a hierarchical modeling approach
for estimating the parameters of demographic matrix models aimed at exploring the spatiotemporal variability in growth and fecundity. This modeling approach is used to explore links to measurable abiotic variables and
demographic impacts on observed patterns of distribution.
DIJKSTRA, FEIKE A.,* SARAH E. HOBBIE and PETER B. REICH.
University of Minnesota, Saint Paul, Minnesota, USA. Nitrogen losses
and cycling affected by fire and vegetation in an oak savanna.
Long-term prescribed fires have increased woody canopy openness and
reduced nitrogen (N) cycling (i.e. net N mineralization) in an oak savanna
in Minnesota, USA. It is unclear if the reduced N cycling is caused by
greater N losses with fire that reduce mineralizable N pools or by the
conversion in vegetation from oak to C4 grass dominance, which reduces
N mineralization per gram of soil N. We established oak-dominated plots
in unburned sites, and oak- and grass-dominated plots in frequently burned
sites (burned 2 out of 3 years on average during the last 40 years) that
allowed us to make contrasts across fire history and between vegetation
types. We measured in situ and potential net N mineralization, total soil N,
and N losses through volatilization and leaching. In frequently burned sites,
we also measured in situ and potential net N mineralization and total soil
N underneath large single oaks or cluster of oaks that died 5, 10, or 15
years ago and that are now dominated by C4 grasses. Net N mineralization
in the oak-dominated unburned plots were two times higher than in oakdominated burned plots, which in turn were four times higher than in the
grass-dominated burned plots. Nitrogen losses in the frequently burned
plots were much higher than in the unburned plots and mainly occurred
through volatilization that most likely reduced total soil N and thus net N

mineralization in the frequently burned plots. Nitrogen losses were not

different between the oak- and grass-dominated burned plots. Therefore,
the conversion from oak to grass dominance must have further reduced net
N mineralization in the burned plots. Net N mineralization in soils near
dead trees, irrespective of the time of death, was as low as in the grassdominated plots. The rapid reduction in net N mineralization after tree
death suggests that this may be caused by the disappearance of living trees
rather than by the increased abundance of grasses. Our results show that
long-term prescribed fires directly reduce N cycling because of increased
N losses and that the reduction in N cycling is exacerbated by a change in
vegetation from oak trees to C4 grasses.
DILUSTRO, JOHN J.,* BEVERLY S. COLLINS and LISA K. DUNCAN.
Savannah River Ecology Laboratory, Aiken, SC. Soil nitrogen cycling in
mixed forests of varying soil texture at Fort Benning, Georgia.
Soil nitrogen cycling is influenced by soil texture, nitrogen inputs, vegetation, and land use. We analyzed soil nitrogen mineralization, nitrification,
and leaching in mixed pine hardwood forests as part of a study examining
land use and soil texture effects on vegetation and nitrogen dynamics at
Fort Benning, Georgia. These forests are managed by prescribed fire (three
year rotation), and burns were conducted on all sites prior to our research.
The 32 forest stands sampled range from sandy to clayey soil texture and
lighter (infantry) to heavier (mechanized) military training intensity. Pooled
soil organic layers were collected from all sites in 2001 and 2002, and
laboratory incubations were used to measure soil mineralization and nitrification during the growing seasons. In 2002, field soil incubations were
conducted in both the dormant and growing seasons, and tension lysimeters
were used to measure soil nitrogen leaching in eight stands. 2001 laboratory results indicate greater initial extractable mineral soil nitrogen in clayey sites with lighter land use intensity. After 84 days, heavier use areas
with sandy soils had the greatest nitrate production and overall mineral
nitrogen pool. Organic layer dry mass (1166 g/m2) and nitrogen pool (8.7
g N/m2) were greatest in clayey sites with lighter military training. However, nitrate production was low in all sites. Evaluating nitrogen transformations can supplement management practices to enhance productivity in
these southeastern mixed pine hardwood forests.
DIRZO, RODOLFO,1,* AMY WHIPPLE,2 NEIL COBB2 and THOMAS
WITHAM.2 1 Instituto de Ecologia, Unam, AP Post 70-275, Mexico, DF;
2
Merriam-Powell Center, Dept. Biological Sciences, Northern Arizona
University, Flagstaff, AZ. Long-term effects of herbivory on male and
female reproductive function of a dominant tree.
We present results derived from long-term experiments (fourteen yrs) and
observations (three yrs) that quantify how a keystone insect herbivore that
attackes juvenile trees and has long-term fitness consequences that greatly
exceed its short-term effects. At our study sites, the scale insect, Matsucoccus acalyptus, preferentially attacks juvenile pinyon. In our observations we measured the variation in scale herbivory and pinyon reproduction
within a natural population in three years (1997, 1998 and 2000). The longterm experiment consisted of the comparison of co-occurring resistant, susceptible, and experimental susceptible trees, which have had scales removed since 1986. Three major major patterns were detected. First, in the
unmanipulated pinyon population, we found the strongest negative relationship between pinyon male reproductive function and herbivory with a
measure of scale herbivory that integrates over six years of cumulative
impact. Second, in the experimental population susceptible trees suffer a
dramatic 13- to 20-times reduction in male strobili and female conelet
production, respectively, as compared to resistant trees. Third, to differentiate between potential alternative hypotheses, when scales were experimentally removed for 14 years, their reproduction rebounded to resemble
resistant trees. A total accounting of the negative effects of herbivory (e.g.,
scale herbivory) on trees (e.g. pinyon trees) requires long-term experiments
and observations that include the analysis of both male and female reproductive functions.
DITTBRENNER, BENJAMIN J.,* REBECCA R. SHIRER, ANNA L.
HARTWELL and G. S. KLEPPEL. Dept of Biological Sciences, University
at Albany, SUNY, 1400 Washington Avenue, Albany, New York, United
States. High-resolution, hyperspectral assessment of wetland ecosystems: Classification from a remote platform.
Efforts to ascertain the integrity and functionality of wetland ecosystems
are constrained by their diversity and abundance in certain parts of the

country, and by the frequency with which access to wetlands is limited by

private property issues. Wetland monitoring from remote platforms is considered by many as a solution to several of these problems. We are studying
the feasibility of using an aircraft-flown 37-channel, hyperspectral scanner
(resolution51 sq m) for wetland assessment in New York State (which has
ca. 10 million ha of wetlands). We are testing the hypotheses that: (i)
hyperspectral imagery (HI) can be used to classify wetlands on the basis
of the biological and hydrogeomorphic (HGM) attributes and (ii) HI can
be used to detect disturbance in wetland ecosystems. In this contribution
we address hypothesis (i). The study area consists of 80 sites, 40 in each
of the Upper and Lower Hudson River valley basins, targeting emergent
marsh or forested wetlands. Data collected on the ground at each site includes plant community structure, hydrologic regime and a suite of standard
physical and chemical attributes. High resolution, remotely sensed hyperspectral data are compared with spectral reflectance signatures of the dominant wetland macrophytes obtained on the ground to characterize each
wetland floristically. Additional detail for the classification is obtained from
hydrologic and geologic features that can be resolved both on the ground
and remotely.
DIVITTORIO, CHRISTOPHER T.* University of California, Berkeley,
Berkeley, CA. Local dispersal and seed limitation promotes native
grass persistence.
Disturbance colonization and native grass persistence are related to patterns
of local dispersal in a species rich, highly invaded California grassland.
Disturbances were created along a gradient of native to exotic grassland.
Species composition and abundance of colonists over two years were related to the abundance and composition of species in seed rain, seed banks,
and the local (1 m2) community. This allows resolution to the species level
of the importance of dispersal, dormancy, and local community control.
The results show that the best predictor of the composition and abundance
of species on disturbances was the species composition and abundance of
the local community, indicating a high degree of local control and a high
prevalence of seed limitation. Contrary to popular theory, native grass colonists dominated disturbances where they dominated the local community.
After two years, native species continued to dominate these sites, and to
coexist with exotic species in those sites whose local community contained
a combination of native and exotic species to begin with. Seed trapping
revealed that local dispersal was the primary mechanism of colonization
for the dominant native and exotic species. There was, however, considerable variation between common and rare species, and between different
functional groups. Rare grass and forb species showed different patterns,
with a greater proportion of species colonizing from dormant seeds in the
soil. Some exotic forb species possessing superior dispersal capabilities
colonized all sites regardless of presence in the local community. This study
demonstrates that grassland communities can be characterized by high degrees of seed limitation and that local dispersal which leads to the persistence of native species, even within a matrix of exotics. Additionally, functional classification of species is essential to our predictive capability, and
to understanding the mechanism of past invasion.
DIXON, JEREMY,1,* WALTER MCCOWN,2 THOMAS EASON3 and
MADAN OLI.1 1 University of Florida, Gainesville, FL; 2 Florida Fish and
Wildlife Conservation Commission, Bear Management Section, Gainesville, FL; 3 Florida Fish and Wildlife Conservation Commission, Bear Management Section, Tallahassee, FL. Gene flow and dispersal among Florida black bear populations: Evaluating the effectiveness of a regional
corridor.
Florida black bears (Ursus americanus floridanus) once occurred throughout Florida, and southern portions of Georgia, Alabama, and Mississippi.
Currently, the subspecies primarily occurs in eight distinct and mostly isolated populations throughout Florida. The long-term isolation of these populations could lead to a loss of genetic variation and evolutionary potential
and may reduce population viability. The connection of populations with
regional corridors has been proposed as a way to mitigate the effects of
isolation by increasing dispersal and gene flow. One such potential corridor,
located between the Ocala and Osceola bear populations, was evaluated for
effectiveness using non-invasive hair snares and molecular analysis. Within
the corridor, grids with cells measuring 20 km2 were overlaid on potential

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89

areas, and at least one hair snare was placed within each cell. From May
to November of 2002, 295 black bear hair samples at 33 hair snare sites
were collected. Additionally, samples were collected opportunistically from
existing fences and bears killed on roads. Hair samples were analyzed using
microsatellite analysis and provided 12-loci genotypes. Using population
assignment tests, preliminary analyses revealed that black bears sampled
from the Ocala-Osceola corridor were genetically consistent with bears
from the Ocala population, indicating a total one-way movement of bears
from the Ocala population. However, more samples and analyses will be
needed to fully characterize the role of the corridor in facilitating dispersal
and gene flow between the two populations. Conservation implications of
preliminary findings will be discussed.
DIXON, KEN R.* and STEPHEN B. COX. The Institute of Environmental
and Human Health, Texas Tech University, Lubbock, Texas. Metapopulation models of bubonic plague in black tailed prairie dogs: Individual
vs. population level approaches.
At the regional scale, black-tailed prairie dog colonies are a classic metapopulation: a spatially subdivided population whose subunits (colonies)
experience periodic extinction events and are linked by migration. A traditional metapopulation modeling approach would determine the relative
influences of landscape attributes on plague prevalence and outbreak frequency. However, this approach would ignore many of the demographic
and individual level attributes (that may themselves be functions of surrounding landscape attributes) about prairie dogs and fleas that may determine potential plague threat to humans. We compared results from individual level and population level models of plague dynamics within a prairie dog metapopulation. The population model was a modification of a SIR
type model initially developed for rats. The individual based model incorporated aspects of the SIR model, but with the spatial component being
based upon a cellular automata model developed for an epidemiological
study of rabies in raccoons. Simulation results suggest that variation in
individual movement patterns, as determined by surrounding landscape features, have a significant impact on the probability of disease outbreak. This
variation in individual movement patterns is not reflected in population
level models.
DIXON, ROBERT M.* and ANN B. CARR. The Imprinting Foundation,
Tucson, AZ. Land imprinting specifications for ecological restoration
and sustainable agriculture.
The no-till method for seeding called land imprinting has been under development in Tucson, Arizona since 1976. Ranchers have applied imprinting to restoring perennial grasses on their degraded rangeland for erosion
control and forage production. Since 1980 some 20,000 hectares have been
interseeded with grasses in southern Arizona. Early imprinters were massive machines with large diameter rollers that were designed to operate on
rocky, brushy terrain of the southwestern deserts. Imprinters are now being
designed for ecological restoration and the establishment of cover crops.
They have smaller diameter rollers and are easier to transport. Some can
work on 2:1 slopes and even steeper. Simple seeders, directly driven from
the imprinting roller, can deliver complex mixes of native seeds to the roller
top where they are carried forward, dropped on the soil surface and then
imbedded in the imprint surfaces. V-shaped imprints funnel resources
downward where they can work in concert to germinate seeds and establish
seedlings. Based on more than 2 decades of field experience, land imprinting specifications have been developed for ecological restoration and sustainable agriculture. These include general imprinter and seeder design
specifications that will help to insure success of revegetation projects. Also
experienced fabricators can use these specifications as a guide for constructing state-of-the-art seeding imprinters.

terranean resources as larvae but are ground surface dwellers as adults. To

investigate the connections between belowground and aboveground food
webs, we are conducting a study in a desert region of SE Spain. Our first
goal has been to describe the composition and distribution of the soil macroinvertebrate fauna. From April 2001 to March 2002, we have conducted
a preliminary sampling using soil cores to extract the macrofauna up to 80
cm depth. Samples were taken monthly under the dominant shrub species,
on bare soil areas and under ant-nests. A total of 1847 individuals belonging to 18 orders were collected. The assemblage was dominated by arthropods, which comprised 99.8% of the total number of macroinvertebrates.
The most abundant groups were Hymenoptera (mostly Formicidae) Coleoptera (principally Tenebrionidae, Carabidae, Cebrionidae and Scarabaeoidea), Hemiptera (Margarodidae) and Embioptera (Oligotomidae). The fauna showed significant differences in vertical distribution, most families
(85%) occurring in the more superficial level (0-13.5 cm). In general, abundance decreased with soil depth, except for Formicidae and Scarabaeoidea
which showed higher number of individuals at deeper levels. Macroinvertebrates also differed in microhabitat distribution. Results show that diversity and abundance were significantly lower in bare ground areas and differed among shrub species. Holometabolous insects comprised most of the
species and abundance in this desert region, contrasting with soil assemblages reported for other ecosystems. These results suggest an important
role of these arthropods as ecosystem engineers and decomposers connecting below and above ground processes in these Mediterranean arid
habitats.
DOBROWOLSKI, WENDY M.,* PATRICK MINGUS and NANCY
GRULKE. Pacific SW Research Station, USDA Forest Service, Riverside,
CA. Tree-level nitrogen limitation at a nitrogen-saturated site: An example from California black oak.
Monthly measurements of leaf and branch growth, foliar mortality, and gas
exchange of Quercus kelloggii were conducted at an N-saturated mixed
conifer stand in the San Bernardino Mountains for the duration of the 2002
growing season. Half of the trees had been fertilized for 5 years with 50
kg ha-1 yr-1 slow release urea, doubling the annual N deposition already
high from atmospheric sources swept in from Lost Angeles, CA. Seasonal
patterns of gas exchange rates of fertilized and background N trees were
nearly identical, except that late summer net assimilation was 35% lower
for fertilized trees. The depressed gas exchange appeared to be correlated
with higher foliar chlorosis in fertilized trees, an unexpected result. In the
European literature, medium term fertilization of Betula resulted in a mitigation of foliar ozone injury. Stem and leaf elongation growth was greater
in fertilized vs. background N deposition trees. However, leaf production
was lower, and leaves were lost more quickly in background N trees, an
unexpected result from published data on litterfall on a whole tree basis.
The number of leaf primordia stimulated to grow was similar in the two
treatments. It is well-established that high N (and ozone) deposition sites
have greater litterfall, contrary to our branch-based data. At a highly polluted site, N deposition may significantly affect foliar ozone injury expression. Elevated N deposition may also affect canopy carbon allocation
and retention in unexpected ways.
DOBSON, ANDREW P. Princeton University, Princeton, NJ. Infectious
Diseases in Multi-Host Systems.
Most emerging diseases are ones that have transferred between host species. Most important in this respect are pathogens that move between wild
hosts and domestic hosts, or from domestic (and wild) hosts to humans.
Understanding the ecology of these diseases requires us to develop a framework in which to examine the dynamics of pathogens that infect two or
more host species. In this talk I will describe and illustrate some mathematical models that allow us to explore this problem. I will then provide
examples from pathogens of carnivores in the Serengeti and of birds in
Hawaii that illustrate some of the patterns predicted by the models.

NCHEZ-PIN
ERO*
DOBLAS-MIRANDA, ENRIQUE, FRANCISCO SA
LEZ-MEGIAS. Dpto. Biolog
and ADELA GONZA
a Animal y Ecologa,
Facultad de Ciencias, Univ. Granada, Granada, Granada, Spain. Composition and distribution of belowground macroinvertebrates in a desert
region of southern Spain.

DOBSON, F. STEPHEN,1,* DAVID BROUSSARD1 and JAN O. MURIE.2

1
Department of Biological Sciecnes, Auburn University, Auburn, Alabama,
USA; 2 Department of Biological Sciences, University of Alberta, Edmonton, Alberta, CANADA. Capital and income breeding in Columbian
ground squirrels.

Belowground detrital food-webs are connected with ecological processes

aboveground via the complex life cycles of many insects which use sub-

Reproductive females may use stored resources (capital) or daily intake of

food (income) to produce and nurture offspring. We studied capital influ-

90

Abstracts

ences on reproduction in an income-breeding small mammal, Columbian

ground squirrels, in two geographically separated populations. In both populations, reproductive investments were significantly correlated with body
mass of mothers when they emerged from hibernation in the spring, suggesting an influence of capital on their reproductive tactics. On the other
hand, mothers gained personal mass during reproduction, suggesting that
reproduction was supported by daily resource acquisition. Differences in
spring body mass were associated with different tactics of reproductive
investment. Mothers in poor spring condition gained more personal body
mass during reproduction, at a cost of producing fewer young. Mothers in
good spring condition invested less in themselves, but maintained superior
body condition while producing more young. These different tactics had
fitness consequences, as reflected by the latter mothers producing more
offspring that survived to a subadult age class. Thus, we documented a
capital influence on reproduction in an income breeding species.
DOMINGUES, TOMAS F.,1,* LAWRENCE B. FLANAGAN,2 LUIZ A.
MARTINELLI,3 JEAN P. OMETTO1,3 and JAMES R. EHLERINGER.1
1
University of Utah, Salt Lake City, Utah, USA; 2 University of Lethbridge,
Lethbridge, Alberta, Canada; 3 University of Sao Paulo, Piracicaba, Sao
Paulo, Brazil. Relationships between carbon assimilation and foliar nitrogen concentration in tropical rain forest functional groups.
Significant correlations have been observed between maximum assimilation
rate and foliar nitrogen concentration in leaves of many temperate species,
largely because the nitrogen is allocated to photosynthetic activities. This
assimilation-nitrogen relationship has been used to simplify a number of
ecosystem-scale carbon balance models. In the present work we tested the
hypothesis that carbon assimilation rates can be predicted based on leaf
nitrogen content for a tropical rain forest ecosystem in Santarem (PA),
Brazil. Four distinct plant functional groups were evaluated: top canopy
lianas, top canopy trees, mid canopy trees and understory trees. As a functional group, trees did follow the expected linear assimilation-nitrogen relationship. The slope of the relationship shifted with season, becoming
steeper during the dry season. The lianas functional group showed high
foliar nitrogen concentrations, similar to those present on top canopy trees.
However, high nitrogen levels in lianas did not translate into high photosynthetic capacity, placing this group below the observed assimilation-nitrogen trend line. We observed that lianas operated with significantly lower
stomatal conductances, resulting in a greater stomatal limitation to carbon
assimilation. Current work is being done to understand why lianas maintain
high nitrogen levels on their leaves without benefiting from high assimilation levels.
DONAHUE, MEGAN J.* University of California, Davis, mjdonahue@
ucdavis.edu, Davis, CA. Gregarious settlement and size-dependent
trade-offs: Better to be small than dead?
Gregarious settlement is a common phenomenon among marine organisms,
in which na
ve larvae settle preferentially with conspecifics. Researchers
in conspecific attraction have proposed several benefits and costs of this
behavior; benefits include (i) information about habitat quality, (ii) reduced
risk of predation, (iii) facilitation, and (iv) future mating opportunities;
costs include (i) enhanced competition, (ii) increased predation if predators
are attracted to prey aggregations, and (iii) increased sensitivity to disturbance. When predation and competition are size-dependent, the tradeoffs
inherent in gregarious settlement become more severe: predation risk is
higher for small individuals, but they suffer more from competitive interactions. These tradeoffs between growth and mortality may be important
in the life history of Petrolisthes cinctipes. Jensen (1989, 1991) demonstrated that P. cinctipes settles preferentially with conspecifics over congeners and showed that settlers accrued two benefits from doing so: information on habitat quality and decreased risk of predation by intertidal fishes. In another study, I have documented the costs of gregarious settlement:
growth rates of individual P. cinctipes decline with conspecific density and
this decline is more pronounced for smaller individuals. Here, I draw on
data from competition and predation experiments, and work by Jensen
(1989, 1991), to model the fitness costs and benefits of gregarious settlement. Using this model, I consider what balance of size-dependent costs
and benefits can result in the observed functional form of gregarious settlement in P. cinctipes.

DONALDSON, JACK R.,* ERIC L. KRUGER and RICHARD L. LINDROTH. University of Wisconsin - Madison, Madison, Wisconsin. Genotype and environment effects on defensive chemistry and growth in
aspen.
Aspen (Populus tremuloides) exhibits considerable genetic variation in allocation to defensive chemistry and in inherent growth rate. Understanding
how environment interacts with genotype to influence growth and defense
in aspen can provide important insight into the ecological interactions of
this widespread tree species. This study examined the effects of genotype,
soil fertility, and below ground competition on defensive chemistry and
growth in aspen. Four aspen clones were micropropagated and grown in a
common garden. Resource availability was manipulated artificially by soil
amendment, and half of the trees were grown in competition with marsh
reed grass (Calamagrostis canadensis). Phenolic glycoside (PG) and condensed tannin concentrations differed among clones and responded markedly to both fertility and root competition treatments. Aspen relative growth
rate (RGR) varied 3-fold in response to fertility and by 20% in response
to grass root competition, but was only marginally different among the four
aspen clones used in this study. We found significant genotype by fertilization and genotype by competition treatment effects on PG and tannin
concentrations, but there were no significant interactive effects on RGR.
Leaf nitrogen was strongly correlated to photosynthesis, RGR, and tannin
concentrations. In fact, leaf tannin concentration was the best predictor of
RGR explaining 73.9% of the among treatment variation in RGR. In this
short-term study we observed strong main effects and interactive effects of
treatments on chemical allocation, however these effects did not lead to
genotype by environment effects on RGR.
DORN, NATHAN J.* and JEREMY M. WOJDAK. W. K. Kellogg Biological Station, Michigan State University, Hickory Corners, MI. Crayfish
alter succession of pond communities.
Succession studies in freshwater ecosystems have commonly focused on
the seasonal-succession of plankton in pelagic zones of lakes and relatively
less is known about succession of littoral habitats. In this study, we documented the establishment and abundance of plants and animals in replicated ponds (2 m deep, 29 m diam.) with and without crayfish from June
2001 to June 2002. The ponds were constructed in November 2000, and
in May 2001 bluegill sunfish (Lepomis macrochirus) were added to all 6
of the ponds and crayfish (Orconectes virilis) were added to 3 (density
within natural range). Zooplankton biomass was higher in crayfish ponds
in both years (due to crayfish effects on fish recruitment in 2001), however
phytoplankton biomass was also higher in crayfish ponds during 2001.
High phytoplankton biomass in crayfish ponds was probably the result of
bioturbation and resuspended nutrients (crayfish ponds had higher levels
of suspended inorganic matter in 2001). In control ponds, Chara and other
macrophytes covered 34% of the bottom by June 2002, while crayfish
ponds completely lacked Chara and other macrophytes. In 2001, mats of
metaphytic green algae (Cladophora and Zygnema) were abundant in the
control ponds but were absent from the crayfish ponds. Selective grazing
by crayfish incited a bloom of filamentous blue-green algae (Gleotrichia),
which dominated the metaphyton community in crayfish ponds late into
the summer, but was never found in the control ponds. Bullfrog tadpoles
were absent from crayfish ponds due to indirect effects (breeding site destruction). Snails were also less abundant, and the negative effects on these
grazers cascaded to periphyton which was more abundant in the crayfish
ponds at the end of 2001. Total benthic invertebrate biomass did not differ
between treatments, but chironomids and mayflies had altered size-distributions. These results indicate that omnivorous crayfish can have strong
impacts on successional pathways of shallow freshwater ecosystems.
DORNBUSH, MATHEW E.* and JAMES W. RAICH. Iowa State University, Ames, IA. Evaluating the influence of aboveground production on
grassland soil respiration in Central Iowa, U.S.A.
Globally, soil respiration rates correlate positively with mean annual temperature and precipitation. Under similar climates, differences in vegetation
and land use produce notable differences in soil respiration. While consideration of these factors is important for predicting soil respiration, unexplained site-to-site variation within systems still remains. To improve our

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91

understanding of the factors contributing to this variation, we tested the

hypothesis that within-system, site-to-site differences in soil respiration
rates result from differences in aboveground plant production. We addressed this hypothesis by measuring soil temperature, soil moisture,
aboveground net primary production (ANPP), and end of year root biomass
at four Central Iowa riparian grasslands growing on the same soil series,
from May 2001 through May 2002. Annual soil respiration rates ranged
from 730 to 1,300 gC m22, while ANPP ranged from 760 to 1800 gOM
m22. Annual soil respiration rates were poorly correlated with ANPP, however a stronger relationship between annual soil respiration and end-ofseason root biomass was found. Neither live aboveground biomass nor net
primary production rate were significant terms in seasonal models. When
growing season data (April to September) was considered alone, the importance of live aboveground biomass increased, accounting for roughly
5% of model-explained variation. Our results suggest that while aboveground production may be an important predictor of annual soil respiration
rates, this effect does not appear to be independent of belowground biomass
at the spatial scale of this project. Furthermore, correlations between changes in soil temperature and aboveground biomass, and the prolonged winter
dormant season appear to limit aboveground productions explanative potential for seasonal changes in soil respiration.
DORNELAS, MARIA,* SEAN R. CONNOLLY and TERRENCE P.
HUGHES. Centre for Coral Reef Biodiversity, Department of Marine Biology, James Cook University, Townsville, QLD, AUSTRALIA. Indo-Pacific coral reefs and the neutral theory: A multi-scale comparison.
The neutral theory aims to explain patterns of biodiversity and community
structure, and makes specific predictions about how species abundance distributions should vary across spatial scales. It has been proposed that coral
assemblages are well described by the neutral theory. This study compares
species abundance distributions generated by the neutral model to coral
data over three spatial scales (site, island and region), for three reef habitats
(flat, crest and slope), and five regions across the Indo-Pacific biodiversity
gradient. The model9s parameters (the fundamental biodiversity number
and immigration rate) are estimated by maximum likelihood fits to coral
species abundance distributions. In contrast to the theory9s predictions, the
shape of the species abundance distribution is almost invariant across spatial scales. At the reef crest and slope the distribution is log-series like, but
at the reef flat it is frequently bimodal. Estimates of the fundamental biodiversity number, for the five metacommunities, vary predictably across the
biodiversity gradient. Estimates of immigration rate for the local communities are orders of magnitude greater than published estimates from population genetic studies. These data suggest that the neutral theory, at this
stage, is inconsistent with observed patterns in coral assemblages.
DOTT, CYNTHIA E.* FORT LEWIS COLLEGE, 1000 Rim Drive, Durango, CO, USA. Debris flow impacts on riparian vegetation following
the Missionary Ridge Fire, southwest Colorado.
Extensive fires in the American southwest during summer 2002 have
caused significant changes in landscape dynamics. In particular, increased
rates of erosion in burn areas can lead to major re-working of stream drainages. The Missionary Ridge Fire in southwestern Colorado, USA burned
through steep terrain on the flanks of the San Juan Mountains. This study
marks the beginning of a long-term monitoring project to document the
process of fire-induced sedimentation, vegetation loss, and subsequent recovery. Hydrophobic soils created on intensely burned sites triggered major
debris flows in many drainage basins with the onset of monsoon season
rains. Unconsolidated sediments in burn areas have caused a shift in the
location of sediment storage: sediments are removed from the upper reaches of drainage basins, by-pass the high gradient stream channels, and are
deposited in the low-gradient reaches at the base of slopes, where they
inundate riparian habitats. The impact on riparian vegetation is markedly
different in different portions of the drainage basin, since it is disrupted by
erosion and down-cutting on high gradient stretches and by inundation on
lower gradient stream reaches. These impacts, however, are similar in different drainages, regardless of pre-burn vegetation on the uplands. Thus,
in the arid southwest, where periods of extended drought are linked to
increases in fire frequency and intensity, there is an unexpected but predictable effect on riparian vegetation. Drought leads to fire and increased

92

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rates of sedimentation, which leads to inundation and disruption of riparian

system dynamics. In this manner, fire impacts are carried well beyond the
perimeters of the burn area, often impacting people and property thought
to be safe from the flames.
DRAKE, JAMES A. University of Tennessee-Knoxville, Knoxville, TN,
USA. The emergence of structure and control in ecological systems.
Complicated structures and systems are common in nature. They are complicated because the observer (you and me) has difficulty integrating the
entities and processes upon which the system is based, not because of
anything inherently difficult in such systems. This is simply a consequence
of the human neural net and our collective reliance on equilibrium-based
physics of centuries past. Analytical innovation typically resolves complication by providing a new way of observing the system with some sort of
operational adequacy. Complex systems on the other hand need not be
complicated. For example, a very simple cellular automaton can produce
structure that simply cannot be predicted even with a perfect knowledge
of the rules of the game. A hallmark of systems that are truely complex is
emergent behavior and self-organizing mechanics. Here, I outline the nature
of complexity by focusing on the results of experimental studies in system
development. Using a laboratory-based model system I generate, or selforganize, ecological communities which exhibit striking differences in an
emergent propertyvulnerability and resistence to species invasion. This
occurs despite the fact that all systems contain the same species. While
species composition is identical, the systems "phenotypes" differ. To understand complexity one must first be able to create it.
DRAKE, JASON B.,1,* GEORGE C. HURTT,2 RALPH O. DUBAYAH,3
MATTHEW FEARON2 and PAUL MOORCROFT.4 1 D.B. Warnell School
of Forest Resources, University of Georgia, Athens, GA; 2 Department of
Natural Resources, University of New Hampshire, Durham, NH; 3 Department of Geography, University of Maryland, College Park, MD; 4 Department of Organismic and Evolutionary Biology, Harvard University, Cambridge, MA. Lidar detected canopy structure for improved model estimates of carbon stocks and fluxes.
Carbon estimates from terrestrial ecosystem models are currently limited
by large uncertainties in the current condition of the land surface. In this
study we combine recent developments in remote sensing and ecological
modeling in an attempt to improve carbon stock and flux estimates. We
used airborne lidar remote sensing, to measure fine-scale heterogeneity in
the vertical structure of vegetation. This vertical structure is then used to
initialize a new height-structured terrestrial ecosystem model (the Ecosystem Demography model, ED), which is capable of calculating the consequences of fine-scale heterogeneity in vegetation structure in broad-scale
analyses of carbon stocks and fluxes. We initially used a simple lidar statistic, mean canopy height within each 1 hectare grid cell, to initialize the
ED model. This combined approach produced reliable model estimates of
above-ground biomass and provided substantial constraints on model estimates of above-ground carbon fluxes at several study sites in tropical and
temperate forests. At our tropical forest study site (La Selva Biological
Station) we then evaluated the utility of other lidar statistics (e.g., standard
deviation of canopy heights within each grid cell) and lidar waveforms
(profiles of lidar energy reflected from the top of the canopy to the ground).
We found that these metrics resulted in improvements in model estimates
of above-ground biomass (compared to canopy height alone) and further
constrained model estimates of above-ground carbon fluxes. The continued
development and combination of these two technologies is shown to be a
promising approach for improving broad-scale carbon stock and flux estimates.
DRAKE, SARA J.,1,* REBECCA R. SHARITZ,1,2 JOHN J. DILUSTRO2
and BEVERLY S. COLLINS.2 1 Department of Plant Biology, The University of Georgia, Athens, Georgia; 2 Savannah River Ecology Laboratory,
Aiken, South Carolina. A model for predicting C and N transformations
and annual net primary productivity under differing burn frequencies
in a southeastern mixed pine-hardwood forest.
Fire can regulate nutrient cycling and pine dominance in southeastern forests, and the ability to predict effects of differing fire frequencies can help

land managers select an optimal burn regime. Several nutrient cycling models have been developed for southeastern forests, and some of them incorporate fire; however, to our knowledge, no model has considered the effects
of fire frequency on nutrient cycling and annual net primary productivity
(ANPP) of groundlayer plants. Our objective was to develop a model using
data from field studies at Fort Benning, GA, to predict C and N transformations and ANPP of groundcover components under differing burn frequencies. ANPP (biomass) of groundlayer plants is a key model component
because, for example, a large biomass of legumes may help to replenish
nitrogen lost during fire. Preliminary model runs reveal a trend for higher
levels of soil organic layer nitrogen and NH3 and NH4 + in mineral soil
with a 10-year versus a 1-year burn interval. In contrast, NO3 - in mineral
soil and CO2 appear greater with a 1-year burn interval. These results are
expected and suggest the model accurately predicts C and N transformations. Field data show that different abundances of grasses and legumes
under different burn frequencies can be used to refine the model and aid
land managers in choosing a burn frequency that best promotes long-term
productivity of southeastern mixed pine hardwood forests.
DRENOVSKY, REBECCA E.* and KATE M. SCOW. University of California, Davis, Davis, CA. Microbial community succession in desert
basin soils.
In the arid western United States natural and anthropogenic changes in
basin lake levels have exposed saline, alkaline substrates. At Mono Lake,
CA lake level recession has created a dune chronosequence varying in
exposure age, salinity, alkalinity, and plant community composition. We
investigated how exposure age and soil chemistry influence soil microbial
communities, hypothesizing more recently exposed soils would have lower
microbial biomass, lower diversity, and decreased fungal:bacterial ratios.
Microbial community composition was assessed using phospholipid fatty
acid analysis. We controlled for plant effects on rhizosphere communities
by sampling soil near the same shrub species, Sarcobatus vermiculatus, at
three sites along the chronosequence (exposed ,10, 60, and .500 years
before present, respectively). Correspondence analysis of PLFA data indicated microbial community composition varied by site. Older sites had
higher fungal:bacterial ratios and higher diversity (measured as numbers of
detected fatty acids). Relationships between exposure age and microbial
biomass were more complex, although older sites tended to have higher
biomass than the younger site. Canonical correspondence analysis indicated
soil salinity, alone, significantly influenced microbial community composition. However, soil salinity was not correlated with total microbial biomass. Although soil C:N and pH did not appear to influence overall microbial community composition, both were negatively correlated with total
microbial biomass (P50.005 and P50.03, respectively). These results suggest different abiotic factors influence microbial community composition
and size at these three sites along the chronosequence.
DRESS, WILLIAM J.,1,* STEFANIE L. WHITMIRE1,2 and MARK R.
WALBRIDGE.1 1 West Virginia University, Department of Biology, Morgantown, WV; 2 Kellogg Biological Station, Hickory Corners, MI. Chemical characteristics of river and floodwaters of forested floodplains in
the SE United States.
Forested floodplains are common wetland ecosystems bordering rivers and
streams that receive water from the adjacent river during periods of high
streamflow (bankfull discharge). Floodplain sites were established adjacent
to two types of riverine systems (alluvial, blackwater), with underlying
parent materials of two ages (young Pliocene-Miocene 2-24 mya, old
Cretaceous 66-144 mya) (n 5 3 per river type - parent material combination). River water samples were collected to characterize baseflow stream
chemistry in anticipation of sampling flooding events. Water samples were
also collected from 2 sites that experienced flood events in October 2002:
Chicod Creek (young-blackwater system) and Cape Fear River (old-alluvial
system). Blackwater rivers had greater dissolved organic carbon (DOC)
concentration than alluvial rivers (14.3 vs. 5.2 ppm, respectively); concentrations of inorganic ions (i.e. NH4+, NO3-, PO43-) were highly variable
among the river systems. In the Chicod Creek floodplain, concentrations
of NH4+, NO3-, and PO43- were significantly lower within the floodplain
than in river water collected both above and below the floodplain, but there
were no differences in pH, conductivity or DOC between floodplain and

river water samples. In the Cape Fear floodplain, pH (5.65 vs. 6.62, respectively) and conductivity (67.1 vs. 122.3, respectively) were significantly lower within the floodplain, and DOC concentrations (24.6 vs. 12.9
ppm, respectively) were significantly higher. Concentrations of both NH4+
and NO3- were reduced by 95-96% within the floodplain, but there were
no differences in the concentration of PO43-. Differing results from these
flooding events are compared with soil and baseflow water characteristics
from the two forested floodplain ecosystems.
DREWA, PAUL B.1,* and JARROD THAXTON.2 1 Case Western Reserve
University, Cleveland, OH; 2 Louisiana State University, Baton Rouge, LA.
Fire regimes and groundcover vegetation responses in longleaf pine
savannas: testing old-growth hypotheses in second-growth systems.
Prior to European settlement, southeastern longleaf pine savannas were
frequented by lightning-initiated fires that burned large areas during the
early growing season. Such natural fires have been postulated to have a
negative effect on resprouting woody vegetation. It has also been proposed
that anthropogenic alteration of natural fire regimes has resulted in increased abundances and expanded distributions of shrubs, as are commonly
observed in fragmented old-growth savannas. We have experimentally examined the effects of fire season and intensity on shrub demography in
second-growth longleaf pine savannas across the southeastern Gulf coastal
plain. Our research demonstrates that anthropogenic dormant season fires
result in increased stem densities of root crown-bearing shrubs. Following
biennial dormant season fires in bog habitats of Florida savannas, stem
densities of Hypericum spp. were seven times greater than densities prior
to these fires. By contrast, densities did not change appreciably after the
re-introduction of biennial, early growing season fires. Relationships between stem densities of root crown-bearing shrubs and fire intensity are
likely contingent on species and fuel type. Stem densities of H. microsepalum, but not H. brachyphyllum, decreased with increases in intensity of
fires. These fine fuel fires were ineffective in completely killing genets of
these species. Instead, heavier fuels are needed to create fires intense
enough to reduce shrub densities. In longleaf pine savannas of Louisiana,
experimental additions of woody fuels to growing season fires not only
reduced stem densities, but also induced genet mortality of root-crown
bearing species, including Quercus spp. and Ilex vomitoria. Returning natural fire regimes to old-growth savannas will likely maintain present stem
densities of root-crown bearing shrubs. Genet mortality and reductions in
densities appear likely only at localized scales where woody fuels create
hotspots with fire intensities comparable to those associated with treefalls
in old-growth longleaf pine savannas.
DRINKWATER, LAURIE E. Managing biodiversity to restore ecosystem function in intensive agricultural systems.
Reliance on external nutrient sources and biocides has enabled intensive
agricultural systems to maintain productivity with plants selected primarily
for their capacity to produce needed food, fiber and forage. As a result,
most agroecosystems lack the diverse array of plants with heterogeneous
functions that are essential in most unmanaged terrestrial ecosystems. The
extreme simplification of the primary producers in these managed ecosystems has led to changes in belowground biodiversity and a diminished
capacity of key biogeochemical functions. Sustainable management of agricultural systems will depend on our ability to better manage primary
productions to restore internal processes that regulate nutrient use efficiency, plant competition, food web dynamics and net primary productivity.
Because plants do not exist as single species but rather as autotrophicheterotrophic assemblages the impact of primary producers on belowground and aboveground ecosystem processes reflects both the direct and
indirect mechanisms. For example, some plants may have an inordinate
amount of influence on ecosystem processes due to characteristics such as
root architecture, phenology or nutrient acquisition characteristics. Others
may exert influence through associated species, i.e. bacteria or fungi of the
rhizosphere community. Strategic management of plant biodiversity will
require a sophisticated understanding of the direct role of plant characteristics themselves as well as how these plant characteristics shape belowground community structure and function. This presentation will examine
the potential for increased biodiversity of the primary producers to restore
and sustain internally-driven processes through plant- and microbially-mediated mechanisms in agroecosystems.

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93

DUBERSTEIN, JAMIE A.* and WILEY M. KITCHENS. University of

Florida, Gainesville, FL. Tidal forest communities on the lower Savannah River floodplain.
The tidal forests of the deltaic floodplain of the Lower Savannah River
(GA and SC), are forested wetlands subjected to semidiurnal tidal pulses
of freshwater inundation. Development activities have radically influenced
the hydrology of the Lower Savannah (remote impoundments, upstream
and harbor dredging and distributary manipulations, locally). We are investigating forest community structure as an expression of the hydrologic
history of selected sites and indicators of these development activities.
Typical species encountered are of the cypress-tupelo habitat type with
inter- and intra-stand differences detected by cumulating relative frequency,
relative density, and relative dominance. Species-area curves were used to
select plot quadrat types and sizes in two selected areas. Data were collected in 32 plots in 2 stands to quantify community composition, microtopography, soil properties, as well as distance to nearest channel. Tree
cores were also gathered, and dendrochronological techniques were used
to age individuals of known size. DBH was then correlated to age using
regression analysis. Hydrological data was used with the Indicators of Hydrologic Alteration computer software (Smythe Scientific Software 2001)
to discern trends and timeframes (as markers) in the hydrological record
as a basis for age-specific classes for analysis. Multivariate exploratory data
analyses were used to identify species groups, age structure characteristics,
and factors affecting the community compositions.
DUFFIE, CAROLINE V.* The University of Georgia, Athens, GA, USA.
Clay Lick Use by Parrots in Eastern Ecuador: Factors Affecting Daily
Abundance and Distribution.
Geophagy, or clay eating, is a known behavior of many tropical parrots.
One widely held explanation for this behavior is that the clay binds up
toxins present in green fruit seeds, which compose much of their diet. This
study was conducted along the Napo River in Eastern Ecuador where many
species congregate on exposed riverbanks, locally called salados, to take
clay. At one such site, four parrot species in the family Psittacidae are
regularly observed. These species are the Mealy Amazon (Amazona farinosa), the Yellow-crowned Amazon (Amazona ochrocephala), the Blueheaded Parrot (Pionus menstruus), and the Dusky-headed Parakeet (Aratinga weddellii). The first objective of this study was to determine what
effect, if any, that weather and noise disturbance have on daily abundances.
The second was to determine if competition plays a role in determining
how each species utilizes the clay lick. The wall was divided into four
sections and the birds were observed from a nearby blind. Each section
was described in terms of quality as a function of slope steepness and
presence of perching ledges. The following observations were recorded at
fifteen minute intervals: weather conditions, number of parrots, clay lick
sections occupied, presence or absence of parrots in surrounding vegetation,
behavior (fighting, eating, and perching), and number of people inside the
blind. Study results show that light intensity is positively correlated with
abundance for all species except the Blue-headed Parrot. More studies are
needed to determine the effect of noise disturbance on abundances. Results
also show that competition between species plays a role in determining
species distribution on the clay lick. The Mealy Amazon can out-compete
smaller species for superior sections. Smaller species may also utilize the
clay lick at different times of the day or under less favorable weather
conditions as a way of minimizing competition.
DUKES, JEFFREY S.,1,* M. R. SHAW,1,2 N. R. CHIARIELLO3 and C. B.
FIELD.1 1 Carnegie Institution of Washington, Department of Global Ecology, 260 Panama St., Stanford, CA, USA; 2 The Nature Conservancy of
California, 201 Mission St., San Francisco, CA, USA; 3 Department of
Biological Sciences, Stanford University, Stanford, CA, USA. Grassland
productivity responses to individual and combined global changes:
four years at Jasper Ridge.
While many studies have examined ecosystem responses to individual
global environmental changes, few have examined responses to multiple
changes. Because many global changes are happening simultaneously, our
ability to predict ecosystem functioning depends on an understanding of
responses to interacting changes. Here we show how net primary produc-

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Abstracts

tivity (NPP) of a California grassland responded to four individual and

interacting global changes over four years of exposure. While simulated
nitrogen deposition consistently increased NPP, warming and increased precipitation generally had little effect. Elevated atmospheric carbon dioxide
concentrations had increasingly negative effects on NPP over the course
of four years. Each of the four factors had more consistent and usually
more positive effects when applied individually than when applied in combination with other factors. Collectively, results from the Jasper Ridge
Global Change Experiment suggest that productivity responses to global
changes are less than additive, and are harder to discern when factors interact.
DUNCAN, LISA K.,* JOHN J. DILUSTRO and BEVERLY S. COLLINS.
Savannah River Ecology Laboratory, Aiken, SC. Avian response to forest
management and military training at Fort Benning, Georgia.
Evaluating intensity and effects of land use disturbance is difficult, especially in sites with multiple land use, and reliable measures of disturbance
are needed. Many bird species have been shown to respond to habitat
heterogeneity and edge effects created by disturbance. We conducted select
species point counts to determine if abundance of bird species could be
used to assess military training and forestry management practices at Fort
Benning. We selected resident and Neotropical migratory species that might
be expected to respond to disturbance [Bachmans Sparrow (Aimophila
aestivalis), Brown-headed Cowbird (Molothrus ater), Eastern Wood Pewee
(Contopus virens), Indigo Bunting (Passerina cyanea), Northern Bobwhite
(Colinus virginianus), Northern Cardinal (Cardinalis cardinalis), Prairie
Warbler (Dendroica discolor), and Yellow-breasted Chat (Icteria virens)]
and evaluated four disturbance combinations: 1) current burn year, heavy
use (2002H); 2) current burn year, light use (2002L); 3) 2 years post burn,
heavy use (2000H); 4) 2 years post burn, light use (2000L). Point counts
were conducted in 16 upland forest stands, at the center of a 100 x 100 m
plot and 50 m from both ends of two 300 m transects in each stand. Indigo
Bunting and Northern Bobwhite abundance differed between the treatment
extremes; i.e., 2002H and 2000L. Abundance did not differ among treatments for any other species. These results suggest it may be difficult to
use bird species as indicators of differing levels of land use disturbance
within upland forests.
DUNKER, KRISSY J.,1,* JOHN C. VOLIN1 and WILLIAM F. LOFTUS.2
Florida Atlantic University, Davie, FL; 2 U.S.G.S. Center for Water and
Restoration Studies, Miami, FL. Non-indigenous fishes in restored and
natural wetlands in the Everglades Big Cypress Swamp.
1

Over the last century, wetland systems in South Florida have been dramatically altered by the construction of canals and levees and the conversion of natural wetlands. The impact of these changes on native fish populations and invasion by non-indigenous fish populations is poorly understood. This is particularly true for the Big Cypress Swamp, which has been
studied less intensively than other portions of the Greater Everglades ecosystem. This study conducted within the Big Cypress Swamp employed a
variety of sampling techniques to examine fish assemblages within a newly
restored wetland and adjacent canals as well as in relatively intact natural
wetlands. Sampling was done during both the wet and dry seasons at sites
located across a range of habitat types in the restored and natural wetlands.
These included cypress domes and strands, wet prairies, and deepwater
refugia. Initial data showed that canals, which provide water for the restored wetland, have the highest richness of both native and non-native fish
species as compared to restored or natural wetland sites. Overall native
species diversity was significantly higher in natural wetlands than in the
restored wetlands, but there was no significant difference in non-native
diversity. This study provides baseline data for fish communities in the Big
Cypress region. In addition, we have documented the presence of two new
non-native fish species in the Big Cypress Swamp, Hoplosternum littorale
and Belonesox belizanus.
DURBIN, KATIE,* PAUL BRUNKOW and SARAH BURNS. Southern
Illinois University Edwardsville, Edwardsville, IL. Food web manipulation of a hypereutrophic lake.
Cougar Lake on the SIUE campus is a degraded reservoir similar to many
lakes throughout the Midwest. These eutrophic reservoirs are characterized

by poor water quality as defined by high primary production, high sediment

deposition and lowered oxygen levels. This study examined effectiveness
of manipulating food web elements on improving water quality as defined
by a lake that is more environmentally sound, recreationally useful, and
aesthetically pleasant. Experimental pools, 1.75m in diameter, were floated
in the lake, and each contained sediment and lake water. Zooplankton and
native mussels were added to pools to create a fully-crossed two-way design with unstocked pools as controls. Pools remained in the lake for ten
weeks. Neither zooplankton nor mussel grazing/filtering had significant effects on chlorophyll concentration throughout the study; dissolved oxygen
was also not affected by treatment combinations. Temperature and oxygen
concentrations in pools paralleled those in the surrounding lake; chlorophyll concentrations in pools did not experience the same dramatic decline
as observed in the lake. Lack of treatment effects were likely due to very
high levels of primary production in this ecosystem as well as logistical
difficulties of maintaining food webs.
DUURSMA, REMKO A.,1,* JESSE B. NIPPERT2 and JOHN D. MARSHALL.1 1 University of Idaho, Dept. Forest Resources, Moscow, ID, USA;
2
Kansas State University, Div. of Biology, Manhattan, KS, USA. Eight
months of chlorophyll fluorescence measurements on six conifer species
on an altitudinal gradient in northern Idaho.
Light-use efficiency models often make crude assumptions about the time
course of photosynthesis in the fall (when cooling down) and in the spring
(when warming up). One candidate measurement to overcome this problem
is the ratio Fv/Fm, which indicates the efficiency of the photosynthetic
apparatus. Fv/Fm decreases following photoinhibition, damage of the photosynthetic apparatus caused by excess solar energy, especially under low
temperatures. To assess the seasonal time course of Fv/Fm, we measured
Fv/Fm 31 times from September to May on 2 samples of 39 trees of 6
species on 8 sites. The sites were located on an altitudinal gradient between
400 and 1400m asl in northern Idaho; the differences in minimum and
maximum temperatures between the top and bottom of the gradient were
on average 1 and 10 8C, respectively. During the measurement period,
minimum temperature varied between -15 and 3 8C, and maximum temperature between -3 and 40 8C. We analyzed the data with mixed models
to account for the nested structure of the design. A large proportion of trees
retained photosynthetic potential all winter long. Fv/Fm varied significantly
between species (p , 0.001), with maximum and minimum temperature
(p , 0.001), and between fall and spring at any given temperature (p ,
0.001). Species also differed in their response to temperature (p , 0.001).
Ponderosa pine and grand fir were least sensitive to decrease in temperature, whereas Western red-cedar and Engelmann spruce were most sensitive. Using a moving average of minimum temperature with variable window size, we concluded that Fv/Fm was best correlated with the average
minimum temperature over the previous 2-3 weeks.
DWIRE, KATHLEEN A.,1,* CATHERINE G. PARKS,2 DANA L. PERKINS,3 MICHAEL L. MCINNIS4 and BRIDGETT READ.2 1 USDA Forest
Service, Laramie, WY, USA; 2 USDA Forest Service, LaGrande, OR, USA;
3
USDI Bureau of Land Management, Challis, ID, USA; 4 Oregon State
University, Corvallis, OR, USA. Seed production, dispersal, and age
determination of Potentilla recta L., an invasive non-native species in
northeast Oregon.
Sulfur cinquefoil (Potentilla recta L., family Rosaceae) is an invasive, herbaceous perennial, native to Eurasia. It has a wide ecological amplitude
and has become established throughout North America in numerous habitat
types. We suspect that the range of sulfur cinquefoil, although rapidly expanding, is underestimated because the plants frequently co-occur with native Potentilla species, and are misidentified as native. Sulfur cinquefoil
reproduces only by seed (achenes). To characterize its demography and
seed ecology, we quantified seed production, measured seed dispersal, and
aged plants from sulfur cinquefoil infestations in different habitats in northeast Oregon. Seed production for 2 years was 3-4 times higher than previously reported (mean 5 5600 seeds per plant; range 5 2770 to 8870
seeds per plant). Seed dispersal was measured using sticky traps (30 x 100
cm, replaced every two weeks) radiating in four cardinal directions from
individual plants. Seeds fell from early July through September (2001 and
2002), and were mostly dispersed within 1 m of the parent plant. Distri-

bution patterns of sulfur cinquefoil also strongly suggest that human and
animal-mediated dispersal contribute to satellite expansion at multiple spatial scales. Plant age was determined by counting annual rings in the primary taproot, which were cross-sectioned just below the root crown and
stained. Preliminary results indicate that most plants are 4-6 years old
(maximum 12 years), and that stand age is related to local disturbance
events. Ongoing research on sulfur cinquefoil in the Pacific Northwest includes studies on seed germination, seed bank viability, population genetics, construction of population models, pollination biology, and response
to herbicide and controlled burn treatments.
DYER, SUSAN A.* and TRACY MCLANE.* Westinghouse Savannah
River Company, Savannah River Site, Aiken, South Carolina, USA. Geographic Information System tools for watershed-scale ecological risk
evaluations.
A set of Geographic Information System (GIS) tools have been developed
and are successfully being used to conduct ecological risk evaluations on
large-scale watersheds. The GIS tools automate (1) contaminant source
identification and transport modeling, (2) contaminant concentration
screening against ecologically relevant benchmarks, (3) statistical summarization and graphical display of analytical data, (4) media-specific background evaluations in addition to (5) frequency and pattern of exceedance
evaluations. The watershed-scale project also includes geo-referenced multidisciplined biological data; protected species and habitat locations, and
serves as an archive system for thousands of site-specific research abstracts,
species photographs, and reports.
DZIALOWSKI, ANDREW R.1,* and VAL H. SMITH.2 1 University of
Kansas, dzial@ku.edu, Lawrence, Kansas; 2 University of Kansas, Lawrence, Kansas. Effects of trophic complexity on invasion success of the
exotic cladoceran Daphnia lumholtzi.
We manipulated trophic structure in a series of mesocosms in order to
determine whether differences in food web composition could provide biotic resistance to invasion attempts by the exotic cladoceran Daphnia lumholtzi. We created a gradient of trophic complexity consisting of 1-3 trophic
levels: algae-only; algae + zooplankton; and algae + zooplankton + invertebrate predators (Notonecta). This gradient was also crossed with two
levels of nutrient supply levels to determine whether invasion success was
further affected by productivity. We hypothesized that invasion success
would be highest in the algae-only treatment due to high food resource
availability, and lowest in the algae + zooplankton treatment due to strong
food resource competition between the herbivores. Furthermore, we hypothesized that the addition of an invertebrate predator would influence
invasion success due to D. lumholtzis anti-predator adaptations. Daphnia
lumholtzi possess extravagant predator-deterrent head and tail spines that
are longer than those found on native competitors. Predators therefore could
selectively feed upon native zooplankton, potentially leading to lower zooplankton biomass, greater food resource availability, and reduced biotic
resistance to invasion. Our initial results show that: (1) Invasion success
was high in algae-only mesocosms, and D. lumholtzis total biomass was
positively correlated with water column nutrient availability; (2) D. lumholtzi was unable to invade when other zooplankton competitors were present, regardless of the nutrient supply level; and (3) the presence of an
invertebrate predator did not influence invasion success. Our results suggest
that the native zooplankton communities of U.S. lakes and reservoirs may
exhibit natural biotic resistance to invasion attempts by D. lumholtzi, and
that further research is needed to identify the mechanism(s) that are responsible for the successful invasion and range expansion of D. lumholtzi.
EATON, JAMES M.* and DEBORAH LAWRENCE. University of Virginia, Charlottesville, VA. Woody detritus in a shifting agricultural ecosystem.
Woody detritus is an often overlooked, but potentially significant, carbon
source in older secondary forests and an important sink in younger secondary forests. The southern Yucatan peninsular region (SYPR) is characterized by a mosaic of agricultural lands and secondary forests. Shifting
cultivation, in the form of slash and burn agriculture, degrades this dry
tropical forest, yet leaves much woody detritus due to incomplete burns.

Abstracts

95

These inputs are important in an early successional ecosystem because

woody detritus rebuilds soils, restores soil nutrients, and limits erosion. We
explored the major environmental constraints on the transition from sink
to source, by sampling along a chronosequence of sites and a regional
precipitation gradient. To determine the heterogeneity, structure, and dynamics of woody detritus during succession, we inventoried woody detritus
stocks at two locations in the southern Yucatan peninsular region, while
monitoring inputs and decomposition of woody detritus over a two year
period at one of these locations. Stocks of coarse ($10 cm diameter) woody
debris were inventoried in two 16 m fixed radius plots, while a sub-sample
of these plots using 1m2 quadrats yielded estimates of fine ($1.8 cm to #
10 cm diameter) woody debris for each site. Every 6 months, we measured
inputs of coarse woody debris (CWD) to these plots. Allometric equations,
based on diameter at breast height of living trees, were used to estimate
inputs of fine woody debris (FWD). The decomposition rates for the 4
most common secondary forest species are being monitored for 2 size
classes over a two year period. Examining the impact of land-use and landuse change on the heterogeneity, structure, and dynamics of woody detritus
in an ecosystem will allow us to understand the function of woody detritus
in secondary forests.
EATOUGH JONES, MICHELE,* TIMOTHY D. PAINE and J. DANIEL
HARE. University of California, Riverside, Riverside, CA. A comparison
of the effects of ecosystem management and unintended anthropogenic
impacts on community richness, diversity and abundance.
Human activities are widely reported to impact ecosystems on both local
and global scales. These alterations can negatively affect ecosystem goods
and services that we rely on. Species richness is frequently used as an
indicator of ecosystem health. Loss of species is assumed to be one of the
driving mechanisms for changes in ecosystem function. We compared effects on communities for two categories; intended alterations through management activities and human activities with unintended ecosystem impacts. Management activities included logging, fire, pesticides and species
introductions for biological control. Unintended impacts included pollution,
habitat fragmentation and invasive species. We surveyed ca. 120 studies
examining community richness, diversity and abundance responses from a
variety of trophic levels and taxonomic groups. Study outcomes were
scored as increased, decreased or no change for each community index.
We assumed the direction of community change would be random and x2
tests were performed to see if all outcomes were equally likely. Abundance
and diversity had non-random outcomes for both types of alterations. Effects on species richness were non-random for unintended impacts but not
for managed systems. Community abundance was affected similarly by
both impacts (decreased in 60% of studies), but species richness and diversity more often decreased due to unintended impacts (60%, 70%) than
for management activities (40%). Vertebrate communities showed the largest difference in response to management activities and unintended impacts.
They were unlikely to be altered in response to management activities, but
showed decreased richness with unintended impacts. Plant, arthropod and
nematode communities were more sensitive to unintended impacts than
management activities. Across pooled trophic levels, we found differences
in the way communities respond to managed and unintended impacts. Community abundance decreased in response to both types of alterations, but
richness and diversity were more often maintained under management activities. Within taxonomic groups, unintended impacts most often resulted
in negative community responses, but management activities had variable
effects.
EBERT-MAY, DIANE1 and CELIA EVANS.2 1 Michigan State University,
E. Lansing, MI; 2 Paul Smiths College, Paul Smiths, NY. Assessing the
Impact of Integrating Scientific Research and Education: What Evidence Will We Accept?
Scientists interested in improving ecological education by integrating scientific research and educational programs need to design appropriate methods for assessment and analysis of data to determine the effectiveness of
these approaches on promoting student learning. We developed systematic
strategies, parallel to the methods of discipline-based science research, for
designing questions and problems to assess student understanding about
ecological concepts in several research projects. Also assessed were faculty

96

Abstracts

and departments involved in educational reform projects. Data from assessment guided decisions about innovative courses, curriculum, and project design. Assessment methods, both quantitative and qualitative, will be
discussed in relation to helping to understand student thinking about the
content and concepts of the discipline. Examples include long-term research about students understanding of the carbon cycle and evolution,
and changes in science faculty teaching.
ECONOMO, EVAN P.,* ANDREW J. KERKHOFF and BRIAN J. ENQUIST. Department of Ecology & Evolutionary Biology, University of
Arizona, Tucson, AZ, USA. Allometric life histories and population energetics.
Establishing connections between individual physiology and ecosystem
level processes is of critical importance to ecology. An important step towards that goal is the linking of individual and population level energetics.
Ecologists have long recognized that individual growth curves and stage
structured mortality rates can be used to quantify the production and efficiency of stationary populations. However, these models have remained
phenomenological with respect to species specific growth and mortality
functions. Recently it has been shown that organisms across diverse taxonomic groups obey a universal growth model. In addition, advances in life
history theory have demonstrated links between growth physiology and life
history parameters, including mortality. Here we combine traditional theoretical approaches to modeling population energetics with allometric models of growth and life history to construct a general model of energetics in
stationary populations. The model approximates the scaling of production
and efficiency in mammalian populations.
EDWARDS, J,1,* P REICH,2 J OLEKSYN2,3 and D EISSENSTAT.1 1 Penn
State University, University Park, PA; 2 University of Minnesota, St. Paul,
MN; 3 Polish Academy of Sciences, Poznan, PL, Poland. Causes for differential calcium accumulation among diverse forest trees: The role of
root distribution.
While carbon and nitrogen cycling have been the primary focus in studies
of biogeochemical processes in ecosystems, recently there has been increased recognition of the importance of calcium as well. Trees vary widely
in the concentration of calcium in their leaves. Differences among species
in the acquisition of calcium and the cycling of calcium via leaf-litter may
profoundly affect ecosystem processes. In a common garden of 14 hardwood and conifer trees planted on relatively homogeneous soil we found
that in just 30 years pH ranges from about 4.0 under Larix deciduas,and
Pinus sylvestris to about 6.0 under Fagus sylvatica and Tilia cordata. The
concentration of Ca in leaf litter ranged from 0.39% to 2.37% and correlated positively with soil Ca, soil pH, soil organic matter and soil C:N.
Root distribution to a depth of 1 meter was determined using a novel
scanning approach. Analysis of preliminary data showed a correlation of
Ca in leaf litter with density of roots, total number of roots and % of total
roots in the C horizon (R50.7). There were no correlations with the O, A
or B horizons. Our data supports the hypothesis that trees that accumulate
more Ca in their leaves deploy more roots in the C horizon than non Caaccumulating trees, contributing to greater Ca acquisition.
EDWARDS, KATRINA J.* Woods Hole Oceanographic Institution, Department of Marine Chemistry & Geochemistry, McLean Lab, MS#8,
Woods Hole, MA, USA. Education and Training in the Biogeosciences:
Challenges, Considerations, and Opportunities for the Next Decade.
In recent years, scientists in the Earth and Life Sciences have embraced
and promoted growth of interdisciplinary science in the various forms that
we collectively term the Biogeosciences. As interest and enthusiasm has
grown, however, so have the challenges that face researchers, students,
educators, and administrators, as we attempt to meet the growing education
and training needs unique to this field. In order to meet the complex educational and training challenges in the Biogeosciences, and other growing
interdisciplinary fields, many specialized initiatives (training grants, educational centers and programs, etc.) have developed in order to help these
communities effectively meet these needs, that otherwise may not be met
by more traditional means. In this presentation I will discuss the specific
challenges unique to interdisciplinary education and training in the Bio-

geosciences. I will discuss ways in which various communities have approached bridging the Earth and Life Science disciplines and infrastructure
needs, and present overviews of some opportunities that presently exist for
development or strengthening of education and training. Finally, I will summarize strides made over the past decade, and the discuss potential obstacles we may face in the next.
EGAN, SCOTT P.,* GREG T. CRYER and JAMES R. OTT. Southwest
Texas State University, San Marcos, Texas, 78666. Host plant quality and
demic adaptation as determinants of the distribution of a gall-forming
herbivore.
Hypotheses correlating variation in host plant quality and herbivore fitness,
when linked with the hypothesis of local (demic) adaptation, provide a
synthetic explanation for the patchy distribution of insect herbivores within
host plant populations. We tested both the among-plant fitness variation
and demic adaptation hypotheses for the host-specific leaf-galling insect,
Belonocnema treatae (Hymenoptera:Cynipidae), on its host, plateau live
oak, Quercus fusiformis. A manipulative transplant experiment was performed where mated females from populations of each of six high gall
density trees (focal trees) were bagged onto branches of (a) their respective
four nearest-neighbor hosts (1 per cardinal direction), (b) the five alternative focal trees, and (c) their natal trees. Each treatmentnearest-neighbor,
alternative focal, and natal was replicated three times at the level of the
individual tree. For each replicate, we recorded: (a) oviposition intensity
(# ovipositor insertion scars/leaf), (b) oviposition success (# galls established/# oviposition scars), (c) individual gall size (diameter), and (d) emergence success (# of gall producing a B. treatae/total # of galls). Herein we
report on gall size, an index of host plant quality. Mixed model ANOVA
showed gall size to be greater (p , 0.001) on natal trees (x 5 5.10mm 6
0.28; N 5 2109 galls) than on neighboring trees (x 5 3.16mm 6 0.28; N
5 2734 galls). Repeated measures ANOVA was used to test for demic
adaptation. Gall size was significantly greater (p , 0.05) on natal trees (x
5 4.53mm 6 0.38; N 5 2116 galls) than alternative focal trees (x 5
3.24mm 6 0.19; N 5 7337). These results suggest that (a) heavily galled
focal trees are surrounded by host trees of lower quality to the herbivore
and (b) the herbivore has undergone local adaptation at the level of the
individual host tree (i.e. deme formation).
EGERTON-WARBURTON, LOUISE M.,1,2 JOSE IGNACIO QUEREJETA,2 AMY E. LINDAHL2 and MICHAEL F. ALLEN.2 1 Chicago Botanic
Garden, Glencoe, IL; 2 University of California, Riverside, CA. Resource
availability and co-existence of mycorrhizal fungi in an oak savanna.
Resource partitioning is an important determinant of species diversity in
many plant and animal communities, but its importance in structuring mycorrhizal fungal communities has been difficult to resolve. This is due
mainly to the difficulties in studying in situ the distribution of fungal hyphae, and how certain species might proliferate in response to resource
availability. We used molecular approaches to study the distribution and
abundance of arbuscular (AM) and ectomycorrhizal (EM) fungal root tips
and hyphae, the structures involved in the uptake of water and nutrients,
in coast live oak woodlands during a severe drought. Comparing the distribution of mycorrhizal types and species with the pattern of available
water across the site (mesic valley to xeric hill top) and with vertical depth
suggested that mycorrhizal diversity was related to a fungal species ability
to exploit a scarce resource (soil water) and tolerate drought or moist conditions. That is, species diversity and dominance, and community composition closely matched soil water potentials. High levels of soil moisture
corresponded with greater EM species diversity, whereas dry soils contained an abundance of AM and certain EM species (e.g., Cenococcum).
These findings are consistent with the idea that shifts in mycorrhizal community composition and species dominance with resource availability
might facilitate the co-existence of mycorrhizal fungi.
EHRENFELD, JOAN G.,1,* JEAN-CHRISTOPHE CLEMENT,1 JUNO
SHRESTHA2 and PETER R. JAFFE.2 1 Rutgers University, New Brunswick, NJ, USA; 2 Princeton University, Princeton, NJ, USA. Effects of
roots and nitrates on redox chemistry of wetland soils.
Wetlands are known for their ability to remove nitrogen from influent waters, largely through denitrification but also through plant uptake. While

the biogeochemical controls on these transformations are reasonably well

known, there has been little investigation of the interactions between plant
roots and the biogeochemistry of the sediments. In wetlands, the distribution of roots may have an especially important role in controlling biogeochemical processes in the sediments because there are numerous potential
feedback processes that connect the spatial distribution of root biomass to
the function of the soil microbial community. Moreover, because denitrification is part of a complex chain of redox reactions, variations in denitrification rate may affect the overall redox status of the soil. We report
initial results from a study designed to determine if and how the presence
of plant roots affects denitrification rates, and, by so doing, alters the redox
chemistry of the soil, and whether these relationships vary with inputs of
nitrate and the quantity of roots. In a field experiment in a riparian hardwood swamp, all aboveground vegetation was removed from experimental
plots, and slow-release N fertilizer was added to half the plots and unmodified control plots. Laboratory microcosms have been established in
parallel to determine the determine the kinetics of iron reduction with and
without nitrates present. The initial samples from the field study showed
that removing the vegetation resulted in increased concentrations of oxidized iron (Fe(III)), and that this effect was accentuated by the addition of
N. Reduction of Fe(III) to Fe(II) is slow, and increases in reduced iron are
only observed after 28 days of soil incubation. These initial results provide
support for the hypothesis that added nitrogen and the density of roots
affect the redox chemistry of wetland soils.
EILTS, ALEX and TRAVIS E. HUXMAN. University of Arizona, Ecology
& Evolutionary Biology Department, Tucson, Arizona. Invasive grass influence over paloverde performance in the arid Southwest.
In the Sonoran Desert, buffelgrass (Pennistemum ciliare) has been systematically converting landscapes, primarily through mortality induced by alteration of the fire cycle. This study addresses the effects the bufflegrass
on a suite of leaf level physiological traits and whole-plant survival in
paloverde (Cercidium microphyllum) using a natural invasion gradient that
has not experienced fire. Contrary to the expectation that buffelgrass would
negatively affect the paloverde, pre-dawn water potential values were not
significantly influenced by the presence of non-native species within the
paloverde canopies as compared to plants without bufflegrass. Leaf-level
photosynthetic gas exchange and chlorophyll fluorescence exhibited a similar pattern. In contrast the ecophysiological data, there was greater mortality for trees in bufflegrass patches. The difference in the data types is
likely due to the habit in the paloverdes known as 9self pruning9; where
by the root to shoot ratio is altered in the tree, maintaining water relations
in the remainder of the tree. This study shows that in addition to the patterns of vegetation change associated with bufflegrass invasion and its influence on the fire cycle, native species can additionally be competitively
displaced in the Sonoran Desert.
EISSENSTAT, DAVID1,* and LIQIN WANG.1,2 1 Penn State University,
University Park, PA, USA; 2 Shandong Agricultural University, Tian, Shandong, China. Physiological responses of roots to pulses of nutrients.
Plant roots commonly proliferate in nutrient-rich patches, often increasing
nutrient acquisition and general plant performance. How do plant roots
respond when the nutrient-rich patch disappears? In a greenhouse study,
apple seedlings were grown with their roots split into two pots containing
unsterilized soil collected from an orchard that had not been fertilized for
at least four years. Roots in the high-nutrient pot received 4 mM nitrate
while those in the low-nutrient pot received 1.6 mM nitrate twice weekly.
Other seedlings had uniform low- or uniform high-nitrate addition to the
two pots. Roots in the high-nitrate pot were then supplied with only lownitrate concentrations. Both nitrate (15N) acquisition and root respiration
declined within 10 d after the shift in nitrate supply. Because both costs
and benefits declined in parallel, the daily root efficiency or cost of nitrate
acquisition was only moderately diminished by changes in nutrient supply.
Root longevity was also diminished in accordance with the diminished root
efficiency. Implications of the costs and benefits of root foraging in ephemeral patches will be discussed.

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ELLIOTT, KATHERINE J.* and JENNIFER D. KNOEPP. Coweeta Hydrologic Laboratory, Southern Research Station, USDA Forest Service,
Otto, NC, USA. Effects of regeneration harvest methods of floral diversity and soil properties.
Forest practices that alter site conditions, either as a consequence of a
timber harvest or a practice intended to improve tree regeneration, may
change biological diversity. We examined how regeneration harvest methods (two-aged shelterwood (2A), shelterwood (SW), and group selection
(GS) cutting) affected floristic diversity, species composition, and soil characteristics of high elevation, mixed-hardwood ecosystems in the southern
Appalachians. Within each treatment area and two uncut areas (UC), four
0.08 ha permanent plots were located to inventory overstory, understory
and herbaceous layer species before (1994) and one year (1998)and three
years after (2000)the harvest treatments. Within each plot, we collected soil
samples at two depths (0-5 cm and 5-15 cm) to determine soil pH, exchangeable calcium (Ca), potassium (K), and magnesium (Mg), percent soil
carbon (C) and nitrogen (N), cation exchange capacity (CEC), and bulk
density. Species diversity was evaluated using species richness (S), Shannon-Wieners index (H), and Pielous evenness index (E). In the overstory,
H and E were significantly reduced in the SW and 2A treatments after
harvest, but no changes in diversity were detected for the GS or UC treatments. Understory density and basal area increased in the 2A and SW
treatments after harvest. H and E were higher on the 2A, SW, and GS
treatments than the UC sites, but there were no significant differences
among the three harvest treatments. In the herbaceous layer, differences
among treatments after the harvest were found for density and percent
cover in 1998 and S in 1998 and 2000. No differences were detected for
H, E, or D among treatments. In 1998, H and E were higher on all
regeneration harvest treatments than before harvest (1994). By 2000, H
and E were no longer different than before the harvest. However, we did
find changes in species composition following the harvest treatments. Significant differences among the treatments varied depending on soil depth
and year of post-harvest measurement. For the 0-5 cm depth, soil Ca and
C were higher for the 2A treatment in 1998 and CEC was lower on the
SW treatment in 2000 compared to the other treatments. For the 5-15 cm
depth, Ca and K were higher on the 2A than on the UC treatment. In 2000,
herbaceous layer diversity was significantly related to soil K, Ca, Mg, CEC
and N.
ELLIS, ALICIA M.1,2 and ERIC POST.1 1 Penn State University, University
Park, PA; 2 Dartmouth College, Hanover, NH. Predicting population response to climate change: A non-linear modeling approach.
The rapidly growing body of research on the ecological consequences of
global climate change has elicited a growing interest in the use of time
series analysis of long-term datasets to investigate population dynamics and
stability in a changing climate. Here, we use non-linear self-excitatory
threshold autoregressive (SETAR) models to investigate the potential influence of climate change on the stability and persistence of an isolated,
undisturbed wolf population on Isle Royale, Michigan, USA. The SETAR
model revealed that the population was influenced by climate only at low
densities, possibly reflecting the overriding influence of density dependence
on dynamics at high densities. Stability analysis suggests that if the population remains above a certain threshold density, it may maintain equilibrium densities despite climatic change. Conversely, if the population decreases below this threshold, climate change is likely to lead to population
decline, possibly to extinction, in the next 100 years. In the absence of
climate change, the population may exhibit multiple stable states, switching
between them when environmental perturbations are strong. Our results
indicate that non-linearity in the strength of density dependence can have
substantial implications for the stability and persistence of populations in
a changing climate. Further investigations into the implications of climate
change for population stability and changes in mean density may facilitate
the formation of appropriate conservation and management policies.
ELLISON, AARON M.1,* and NICHOLAS J. GOTELLI.2 1 Harvard Forest, Petersham, MA; 2 University of Vermont, Burlington, VT. Using
Bayesian and frequentist statistics to develop individual trait-based
models of population dynamics.
We used frequentist and Bayesian methods to estimate parameters for a
population growth model of Sarracenia purpurea, a perennial carnivorous

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plant. Individual, trait-based models typically use fixed parameters estimated from data with frequentist statistics. Variability (stochasticity) in
these models is applied uniformly across individuals by using a common
stochatic error term. In a Bayesian analysis, the parameter estimates are
not assumed to be fixed, but instead vary among individuals and across
time. We compared two classical models (without and with the stochastic
error term) with two models whose parameters were estimated with Bayesian analysis. These models are hierarchical: the classical models are nested
within the Bayesian models. Bayesian models outperformed frequentist
models in recovering observed variability in plant traits and population size
structure. Although individual plants in Bayesian models had lower forecasted survivorship than those in frequentist models, population growth
rates were higher and more variable in the Bayesian models; this led to a
lower probability of extinction predicted by the Bayesian models relative
to the frequentist models. Our results illustrate that population projections
are sensitive to the underlying assumptions regarding parameter estimation
and the sources of measurement and process error. Bayesian modeling
makes these assumptions explicit and could be especially useful in guiding
data collection and model development for the management of threatened
and endangered species.
ELLNER, STEPHEN P.* Cornell University, spe2@cornell.edu, Ithaca,
NY. Understanding simple population dynamics.
Hundreds of populations exhibit cyclic dynamics; in perhaps half a dozen
cases we think we know why. One promising approach to narrowing this
gap is based on fitting and comparing models that represent each plausible
candidate for the causal mechanism. Noise makes this difficult, in particular
the combination of measurement and process noise on top of data that
typically represent a low-dimensional projection of a high-dimensional
nonlinear system. This formerly intractable problem now suffers from an
overabundance of solutions. I will describe simulation-based methods that
can be applied to a wide class of models and data, and applications to field
and laboratory data. The field application is the pine looper moth Bupalus
piniarius, a classic forest insect cycle whose cause remained unresolved
despite many decades of intensive empirical study; for once it isnt trophic
interactions. The laboratory application is a plankton microcosm where
model-based conclusions about underlying mechanism were tested experimentally and confirmed. Many other methods work (in the hands of their
proponents, on computer-generated data); objective, comparative evaluation
of these is now a pressing need.
EMANUEL, RYAN,1 JOHN ALBERTSON,2 HOWARD EPSTEIN,1,* PAOLO DODORICO1 and CHRISTOPHER WILLIAMS.2 1 University of
Virginia, Charlottesville, VA; 2 Duke University, Durham, NC. Carbon
dioxide exchange over an evolving successional landscape.
Successional ecosystems are believed to be significant sinks of atmospheric
carbon dioxide. In order to explore the mechanisms of carbon sequestration
over successional timescales, an eddy covariance system was installed
above a crop field at the Blandy Experimental Farm in Virginia, USA in
January 2001. Half-hourly fluxes of carbon dioxide and water vapor, and
state variables such as soil temperature and soil moisture were measured
continuously through one season of crop growth and the period of succession following harvest of that crop. Here we report the results of the first
year of successional growth in the absence of cultivation. Despite primary
production on the order of 1 kg m-2 during the first full year of succession,
this ecosystem remains a net source of carbon dioxide to the atmosphere.
To investigate this phenomenon, a process-based model of ecosystem respiration was developed using nighttime eddy covariance measurements partitioned into bins based on soil moisture and vegetative structure, and evaluating nighttime flux in each bin as an exponential function of soil temperature. Output from the model was compared with concurrent groundbased measurements of ANPP and soil respiration. Our model implies that
ecosystem respiration exceeds photosynthetic uptake in this early stage of
succession, and demonstrates mechanistically the means by which carbon
flux is controlled by soil moisture, soil temperature and vegetative structure
at hourly to seasonal scales.

EMERY, SARAH M.,1,* SANDY J. ANDELMAN, CHRISTY BOWLES,2

KAREN CARNEY, ELSA E. CLELAND,3 JOHN DRAKE,4 JOEL
GRAMLING,5 M. CLAIRE HORNER-DEVINE,3 MELINDA D. SMITH2
and DAVID B. VANDERMAST.5 1 W.K. Kellogg Biological Station, Michigan State University, Hickory Corners, MI; 2 National Center for Ecological Analysis and Synthesis, University of California, Santa Barbara, Santa
Barbara, CA; 3 Stanford University, Stanford, CA; 4 University of Notre
Dame, Notre Dame, IN; 5 University of North Carolina, Chapel Hill, NC.
Relationships between plant diversity, productivity, and invasion
across six North American grasslands.
Non-native species can threaten native biodiversity and ecosystem function,
making it important to predict where species invasions are most likely. In
this study, we investigated how aboveground net primary productivity
(ANPP) and total species richness influence the likelihood of invasion by
non-native species across multiple community types and years, using data
from six North American grasslands (including five LTER sites.) Native
and non-native species richness within years were positively correlated at
all sites. Total species richness was most often negatively correlated with
both ANPP and the proportion of non-native biomass, but relationships and
their significance varied among sites and among years. These correlations
suggested that while high diversity sites were more likely to be invaded,
low diversity sites were more likely to be dominated by non-natives. We
also hypothesized that the success of non-natives was related to variation
in resource availability in space and time. To separate cause and effect in
these observational datasets, we used logistic regression analysis to ask
whether variation in ANPP, species richness, or precipitation in one year
influenced the abundance or richness of non-natives in the following year.
Interannual variation in these parameters differed in their ability to explain
invader success in the following year, depending on each sites method for
determining species composition, and whether non-native species richness
or abundance was used as the response variable. ANPP was consistently
negatively related to both the abundance and richness of non-natives in the
following year. Precipitation was positively related to the abundance of
non-natives in the following year at two sites, and negatively related to
non-native abundance at one. Interannual variation in total species richness
was significantly related to invader species richness in the following year,
but only in sites where permanent plots were used to measure species
composition.
ENDO, MASAHIRO* and KEIJI NITTA. Institute for Environmental Sciences, Rokkasho, Aomori, Japan. Examination on growing native wetland plants in a geo-sphere module of the Closed Geo-Hydrosphere
Experiment Facility.
The Closed Geo-Hydrosphere Experiment Facility (CGHEF) has been
studying the circulation and accumulation of gaseous radionuclides in the
ecosystem. The CGHEF consists of a geo-sphere module, a hydrosphere
module, and a material circulation system. The objective of this study is
to examine the differences between the geo-sphere module and a wetland
in Rokkasho, and to determine requirements to maintain an artificial geoecosystem in the module. Native plants at the border of the Obuchi-marsh
in Rokkasho were transplanted to containers in the module in June 2002.
Temperature in the module and water supply to the plants were set to the
average temperature and the average amount of rainfall, respectively, for
the previous 2 years in Rokkasho. Almost all of the native plants grew
well in the module, but the times of flowering, fruition, withering, and
dying were delayed from two to four weeks. It appears that temperature
stress was relatively minor in the module. Concentrations of Na+, Ca2+,
Mg2+, Cl- and SO42- in the ground water of the containers became higher
than those of the wetland.
ENGEL, E. CAYENNE* and JAKE F. WELTZIN. University of Tennessee,
Knoxville, TN. How does plant competition mediate the interactive effects of CO2, temperature, and soil water availability?
Global climate change (due to increasing atmospheric CO2 concentration,
increased average global temperature and changing precipitation regimes)
may drastically affect plant community and ecosystem functions. However,
investigations of the interacting factors are rarely attempted. A long term
ecological experiment at Oak Ridge National Environmental Research Park

was assembled to investigate the interactive effects of predicted global

climate change on an old-field plant community. This project, OCCAM
(old-field community climate and atmospheric manipulation,) utilizes opentop chambers with experimental plant communities constructed of seven
common old-field species, including C3 and C4 grasses, forbs, and legumes
to assess the implications of global climate change factors. We assessed
plant community level effects of CO2, temperature, and soil water availability as measured through changes in productivity and species composition. In addition we examined how species interactions may control the
response of the community. Pairwise competition experiments determined
the effects of whole-plant interactions and were used to separate and identify the role of interactions between aboveground components (i.e. light
interference) and belowground interactions (e.g. root competition.) Results
from the competitive assessments are compared with treatments in OCCAM and are used to inform the interpretation of species and community
level response within the experiment. Investigations of the species specific
interactions within these communities allow us to evaluate the ramifications
of abiotic effects on this system and make inferences toward expected
patterns of community development.
ENGEL, VICTOR C.,1,* ESTEBAN G. JOBBAGY2 and ROBERT B.
JACKSON.1 1 Biology Department, Duke University, Durham, NC, USA;
2
Catedra de Forrajes, Facultad de Agronomia, Buenos Aires, BA, Argentina. Canopy water use, groundwater fluctuations and hydraulic gradients beneath adjacent plantation and grassland areas.
Heat dissipation techniques were used to measure sap flow in E. camaldulensis (n518) in a 50 yr. old plantation in the Pampas region of central
Argentina. Piezometer readings of water table depths along a transect from
the plantation interior to the surrounding grassland were also recorded.
Transpiration rates derived from sap flow measurements ranged from 2 to
4 mm d-1 during the summer of 2002-3 and correlate well (r250.65) with
values derived from water table fluctuations beneath the plantation. Residual analysis shows the variance in transpiration estimates between these
two techniques is most closely related to moisture content in the vadose
zone. These results suggest root uptake of soil water by E. camaldulensis
alternates from deep groundwater to more shallow sources depending on
availability and evaporative demand. During dry periods between precipitation events, piezometer readings showed a sharp decline (.1 m) in water
table depth at the plantationgrassland boundary. Mass balance calculations suggest this local hydraulic gradient caused a net flux of groundwater
(;2 to 3 mm d-1) from the topographically low grassland areas to the more
upland plantation. To test our assumptions regarding plantation root activity, source water uptake, and groundwater contributions from the surrounding grassland, we employed a simulation model of the soil-plant atmosphere continuum that includes hydraulic constraints on transpiration and
groundwater transport. Model results compare favorably with sap flow and
water table observations, while on-going sensitivity analyses allow investigations of the influence of stand structure (e.g. root depths, leaf area, tree
height) on annual water balance. The results show the importance of changes in vegetation structure on local hydrologic budgets.
RAN V.1,2,* 1 Department of Ecology and Environmental
ENGLUND, GO
Science, Umea University, E-mail:goran.englund@eg.umu.se, Umea
, Swe, Sweden, Sweden.
den, Sweden; 2 Umea Marine Sciences Centre, Umea
Scale dependent impact of predatory fish on stream benthos: A test of
theory.
In open predation experiments the impact of predators on prey densities
can be influenced both by consumption and by prey movements. A model
predicts that the predator impacts observed in such experiments are scale
dependent over the scale range where there is a transition from movement
control (of prey densities) to consumption control (Englund 1997). To test
this model I examined the impact of predatory sculpins on densities of
stream invertebrates in instream channels of different length (0.5 m, 2 m,
and 8 m). The predator treatment affected the densities of six taxa. Per
capita emigration rates was scale independent for these taxa and comparisons prey emigration and predator consumption rates showed that predator
impacts were due to prey movements rather than predation. Thus, the model
predicts scale independent predator impacts. In contrast to expectations
predator impact on prey density was scale dependent for four taxa. For

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99

three large predatory invertebrates the predator impact increased with experimental scale, and for the fourth taxa, a small invertebrate predator, the
opposite pattern was observed. These scale effects occurred because prey
movement responses to predators were modified by the experimental scale,
a mechanism not included in the original theory.
ENQUIST, BRIAN J.,1,2,* EVAN P. ECONOMO,1 TRAVIS E. HUXMAN,1
ANDREW ALLEN,3 DANIELLE D. IGNACE1 and JAMES F. GILLOOLY.3 1 Department of Ecology and Evolutionary Biology, University of Arizona, Tucson, AZ, USA; 2 Center for Applied Biodiversity Science, Conservation International, Washington, D.C., USA; 3 Department of Biology,
University of New Mexico, Albuquerque, NM, USA. A General Model
for Scaling Temperature and Biochemical Kinetics from Cells to Ecosystems.
Understanding energy and material fluxes through ecosystems is central to
many questions in global change biology and ecology. Our ability to predict
variation in ecosystem processes is currently limited by our ability to mechanistically link biological processes across both spatial and temporal scales.
One promising approach is to focus on how biotic and abiotic factors regulate metabolic rates of individuals, which combine to determine ecosystem
flux rates. Metabolism is the fundamental process dictating material and
energy fluxes through organisms. Here we derive a general biophysical
model of ecosystem respiration based on the kinetics of metabolic reactions
and the scaling of resource use by individual organisms. The model predicts
that CO2 and energy flux is not constrained by ecosystem biomass, but is
strongly influenced by temperature, variation in cellular metabolism, and
rates of supply of limiting resources (water and / or nutrients). Analysis of
intra-site variation in ecosystem respiration, as calculated using a global
network of CO2 flux towers, provides robust support for the models predictions. However, data indicate that inter-site variation in annual flux is
not strongly dependent on average site temperature or latitude. This presents an interesting paradox with respect to the expected temperature dependency. Nevertheless, our model provides a framework by which to
quantitatively assess and integrate such additional biotic and abiotic influences on CO2 flux. Thus, a focus on the fundamental importance of metabolism offers a basis by which to integrate cellular, physiological and
physical/ecological attributes of ecological systems.
ENTCHEVA CAMPBELL, PETYA K.,1,* ELIZABETH M. MIDDLETON,1,* LAWRENCE A. CORP,1,* JAMES E. MCMURTREY,2 LADINE
M. BUTCHER1,* and EMMETT W. CHAPPELLE.1 1 Biospheric Sciences
Branch, Laboratory for Terrestrial Physics, NASA/GSFC, Greenbelt, MD
20771, USA; 2 Hydrology & Remote Sensing Laboratory, Agricultural Research Service, USDA, Beltsville, MD 20705, USA. Detection of changes
in vegetation properties induced by military range land-contaminants
containing TNT.
Due to the environmental impacts of toluene, EPA has mandated that land
mines on US military ranges must be located and removed. Furthermore,
pending international agreements require the removal of land mines placed
on foreign lands. Currently, there are no satisfactory methods for locating
military range land-contaminants, and for delineation of the ecological impacts of leaked trinitrotoluene (TNT). This research determines the changes
in vegetation morphology, bio-physiology, spectral fluorescence (F) and
reflectance (R), for vegetation growing on TNT contaminated soils. The
study aims at the establishment of indicators (biological, spectral F and R)
of the presence of TNT-related compounds sequestered in vegetation. During the summer of 2002, corn (C4), soybean (C3) and pigweed (C4) were
grown in a 3x4 factorial study: 3 species, 4 TNT levels (0% 5 water for
control, 25% 5 20 ppm TNT, 50% 5 40 ppm TNT, 100% 5 80 ppm
TNT), 12 replicates per treatment. Fully expanded sun leaves from mature
plants were used for R and actively induced F measurements. Contemporaneously, foliar photosynthetic function and plant constituent levels (chlorophyll, N and C) were determined. The TNT treatments induced significant
changes in vegetation morphology, spectral F and R properties, which were
more pronounced for the C4 plants (pigweed and corn). The TNT-treated
plants accumulated significantly less biomass, held lower amounts of tissue
water, were significantly lighter (DWcorn r250.82, p,0.01; DWpigweed
r250.88, p,0.01), shorter (Hcorn r250.72, p,0.05; Hpigweed r250.87,
p,0.01) and had thinner stems (BDcorn r250.75, p,0.01: Hpigweed

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Abstracts

r250.86, p,0.01). Foliar parameters (area, length, dry weight, pigment

levels and photosynthetic efficiency) were significantly reduced and the C/
N balance altered. This study documents that in the above-ground tissues
of vegetation, TNT induces significant morphological and metabolic changes, leading to alterations of vegetation spectral F and R properties that are
detectable via remote sensing. Monitoring of vegetation vigor based on
fluorescence and reflectance measurements could provide the necessary
means for detecting contamination from (TNT), a frequent military range
land-contaminant.
EPPERSON, DEBORAH M.1,* and CRAIG R. ALLEN.2 1 The Nature
Conservancy, Camp Shelby Field Office, Camp Shelby, MS; 2 South Carolina Cooperative Fish and Wildlife Research Unit, Clemson University,
Clemson, SC. Impacts of red imported fire ants on gopher tortoises
and their associated burrow commensal fauna.
We examined the potential impacts of red imported fire ants (Solenopsis
invicta) on gopher tortoise (Gopherus polyphemus) hatchling survivorship
and invertebrate burrow commensals. Red imported fire ants were introduced into the United States at the Port of Mobile in the 1930s and have
spread throughout the southeastern United States. Throughout the range of
the gopher tortoise red imported fire ants are now widespread. We experimentally manipulated fire ant densities using a fire ant bait (Fenoxycarb)
on one randomly chosen member of each of five paired sites at Camp
Shelby Training Site in southern Mississippi. Fire ant bait was applied in
the spring of 1998, spring and fall of 1999, and in the spring of 2000. Sites
were paired to reduce heterogeneity based on tortoise populations, S. invicta densities and overall habitat similarities. From 1997 - 2000, gopher
tortoise hatchlings were fitted with radio-transmitters and followed until no
longer extant. Cause of death was determined, and the number of days
alive calculated for each individual. Of forty-one individuals confirmed
dead, 27% of these mortalities were attributed to S. invicta. Comparison of
survivorship between treated and untreated sites using the Kaplan-Meier
method revealed higher survivorship on treated sites (Fishers Exact Two
Sided test P 5 0.087). Invertebrate burrow commensals were sampled twice
per year from 1998-2000 using a D-Vac suction apparatus. Ten burrows in
each site were vacuumed and invertebrates identified to species when possible. Burrow species richness, Shannon Diversity indices, and abundance
were compared between treated and untreated sites using a repeated measures randomized block ANOVA. We found significant treatment effects
for species richness (P 5 0.0016), diversity (P 5 0.0186), and abundance
(P 5 0.0852).
EPSTEIN, HOWARD E.* and WILLIAM M. YEATMAN. University of
Virginia, Charlottesville, VA, USA. Nitrogen resorption from senescing
plant tissue in arctic tundra and its effects on ecosystem properites.
Nitrogen resorption from senescing plant tissue for use in future growth is
a ubiquitous process and can represent a substantial portion of the nitrogen
used in net primary productivity. For vascular plants in arctic tundra ecosystems, the proportion of N retranslocated from senescing tissue has been
observed to range from 10-80%, largely differing based on plant functional
type and species. Values for N retranslocation efficiency are quite variable
within plant species and also across studies; standard deviations of leaf N
concentrations range from 0.02 to .0.50% for means on the order of 0.30
to 2.50% for dead and live tissue, respectively. In low arctic tundra, dominated by dwarf shrubs and tussock graminoids, resorbed N can account
for 40-50% of the N used in annual net primary productivity. Given the
importance of this process in terms of plant N availability, variance in N
retranslocation could lead to substantial changes in whole-ecosystem properties in arctic tundra. In addition to collecting a dataset on N resorption
in low arctic tundra from Ivotuk, Alaska, we used a dynamic tundra vegetation model (ArcVeg) to assess the sensitivity of tundra systems to variability in N retranslocation. Since shrubs represent a substantial portion of
the total resorbed N in the system (due to relatively high leaf biomass and
high resorption), we conducted a sensitivity analysis of N resorption rates
for Betula nana, a dominant deciduous dwarf shrub in low arctic tundra,
by altering resorption rates from 40% to 70%. Variation in N resorption
rates for a single, dominant species, led to substantial changes in simulated
plant community composition and total community biomass. Increasing N
resorption rates for B. nana (from 40 to 70%) increased its own biomass

by a factor of six and increased total community biomass by 25%. Other

community responses to increased N resorption in B. nana were a decline
in evergreen shrub biomass and an increase in non-vascular plants (mosses
and lichens). This study suggests that uncertainties in N resorption rates
by different plant species and functional types have a substantial impact
on our general understanding of the nitrogen cycle and vegetation dynamics
in arctic tundra.
ERICKSON, JOHN E.,* ERIC L. KRUGER and GLEN R. STANOSZ.
University of Wisconsin-Madison, Madison, WI, USA. Consequences of
a fungal leaf spot on photosynthesis of two hybrid poplar cultivars.
Leaf spot induced by Marssonina brunnea (Ell. et Ev.) is an important
foliar disease of many tree species in the Populus genus, yet the consequences of this disease on growth-related physiological processes are not
well known. To explore the effects of disease severity on photosynthesis,
leaf gas exchange and nitrogen concentration were measured on approximately 48 leaves from two hybrid poplar clones, each grown in 4 replicate
plots in the field. The selected leaves represented a wide range of disease
severities and canopy positions. The equation Px/Po 5 (1 - x)b was used to
relate relative net photosynthesis (Px/Po) and disease severity (x) in the two
clones, where b represents the ratio between virtual and visual lesion areas.
Leaf photosynthesis was strongly and inversely related with severity of
Marssonina infection in both clones (P , 0.001). However, b values differed significantly between clones (P , 0.001) with b 5 1.49 in the NM6 clone and b 5 6.84 in DN-34, and both differed significantly from 1 (P
, 0.001). These results were combined with canopy light and disease profiles to estimate the effects on instantaneous canopy assimilation, which
suggested declines of approximately 30% in NM-6 and 45% in DN-34.
Trends in leaf stomatal conductance (gs) were similar to observed patterns
in photosynthesis. In contrast, Leaf N concentration (Nmass) declined only
slightly if at all across the range of disease severities. Similarly leaf dark
respiration (Rd) and the ratio of intercellular to ambient CO2 remained
largely unaffected by disease severity. Thus, we concluded that declines in
photosynthesis were likely a result of disruption of the photosynthetic apparatus or metabolic pathways by the invading pathogen. These results
possess implications for modeling photosynthesis and productivity of terrestrial vegetation infected with foliar pathogens.
ERNEST, MORGAN.* Department of Biology, University of New Mexico,
Albuquerque, NM, 87131. Energy flow, body size, and community
structure in small mammal communities.
Energy is one currency that can link species with the function of ecosystems. Individual energetic requirements, as defined by metabolism, can link
individuals with ecosystems by elucidating the paths and rates of energy
flow through species and communities. Because of its relationship with
metabolism, body size is potentially of great importance in linking individual-level traits with community and ecosystem structure and function.
Indeed, there are a variety of studies that suggest body size may be important not only for understanding energy flow but also for understanding
community structure which has been shown to have some impact on ecosystem dynamics. Here, I examine the relationship between energy flow,
body size, and community structure in nine small mammal communities in
western North America. In all nine communities, energy flow was multimodal with respect to body size. Each energy mode was dominated by a
few species with the rest of the community relegated to body sizes fluxing
very little energy. When body size and energy flow distributions are compared across sites, I found that the body size distribution changes little
between habitats but that the energy flow distribution differs dramatically
between sites. These results are mirrored in long-term data that show
through time the body size distribution is less sensitive to changes in species composition than energy flow. In general, this study indicates that
small mammals appear to perceive the environment as providing resources
in a discontinuous manner with regards to body size but that the overall
body size distribution of the community does not appear to reflect patterns
of resource availability.

ERSKINE, JENNIFER A.* Unviersity of California, Davis, Section of

Evolution and Ecology, Plant Biology Graduate Group, Davis, CA, 95616.
Genotype by environment effects on an invasive species, Foeniculum
vulgare.
Foeniculum vulgare, fennel, is an invasive weed native to the southern
Mediterranean region. It is a prolific invader in California, found in coastal
sage and grassland communities. Fennel displays environmental variation
in invasiveness, and seed production. This study uses four geographically
distinct fennel populations from California to evaluate genotype by environment effects on fennel growth. Initial seed size was measured, and germination rates were calculated under a variety of temperatures. Significant
differences were shown between populations with ANOVA (F5253.945,
p,0.0001) and ( Scheffe post hoc for all comparisons, p,0.0001). Germination rates were significantly different for all four populations at the
temperatures ranging from 16-28 degrees Celsius (p,0.05). A common
garden experiment was used to evaluate genotype by environment effects.
Seedlings were planted in an abandoned agricultural field in Davis, CA, in
February 2002 after field vegetation was removed with herbicide. Plants
were monitored monthly for survival, growth, time to reproduction, and
reproductive output (number of umbels). Ten umbels were collected from
each plant during seed set for estimates of reproductive output, and above
ground biomass was harvested at the end of the growing season. Initial
data analyses indicate differential growth for the four populations, but survivorship at the end of the growing season was not different. Final data
analyses will help imply whether fennel has differential growth throughout
California is because of genetic variability between populations, or effects
of environment.
ERWIN, PATRICK M.* and ROBERT W. THACKER. University of Alabama at Birmingham, Birmingham, AL. Determining the specificity of
associations between marine sponges and symbiotic cyanobacteria.
Symbioses observed between marine sponges and cyanobacteria have
sparked both ecological and biotechnological interest. Associated cyanobacteria may benefit their host by providing additional nutrients (e.g. fixed
nitrogen) and synthesizing anti-predatory and anti-fouling compounds.
Sponges may benefit their cyanobacterial symbionts by providing nutrients
through excreted waste products and a refuge from predation within the
sponge mesohyl matrix. We examined the specificity of association between
host sponges and their cyanobacterial populations for 15 Verongid sponges,
including 7 species, 4 genera, and 2 families. Cyanobacterial 16S ribosomal
DNA (rDNA) was amplified from whole-sponge genomic extracts. For each
sponge specimen, 10 clones of cyanobacterial rDNA were isolated, PCRscreened using cyanobacteria-specific primers, and digested using 2 restriction enzymes. The resulting banding patterns were used as indicators of
the diversity of cyanobacteria associated with each sponge specimen.
Sponges in the genera Aplysina and Verongula hosted multiple genotypes
of cyanobacteria. Of the 10 different observed banding patterns, 1 was
exclusive to the genus Ianthella, 3 were exclusive to the genus Aplysina,
and 4 were exclusive to the genus Verongula, suggesting genus-specific
associations between eight cyanobacterial strains and their host sponges.
The remaining 2 banding patterns were common to sponge specimens from
different genera, suggesting a transient or generalist relationship between
these two cyanobacterial strains and their host sponges.
EUBANKS, MICKY D.,1,* DAVID E. CARR2 and JOHN F. MURPHY.1
1
Department of Entomology and Plant Pathology, Auburn University, Auburn, Alabama, United States of America; 2 Blandy Experimental Farm,
University of Virginia, Boyce, Virginia, United States of America. Complex effects of inbreeding on complex interactions: Self-fertilization alters the interactive effects of natural enemies on plant fitness.
We conducted a greenhouse experiment to determine the effects of inbreeding (self-pollination) on the ability of yellow monkeyflower, Mimulus guttatus (Scrophulariaceae) to tolerate attack by spittlebugs and a plant virus.
We conducted a 2 x 2 x 2 factorial experiment to quantify the effects of
pollination (self or outcross), spittlebug infestation (Philaenus spumarius)
and virus inoculation (Cucumber mosaic virus, CMV) on the fitness of 26
full-sib families of yellow monkeyflower. We found strong evidence that
spittlebugs and CMV interacted to affect plant fitness, but in several cases

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101

inbreeding altered the strength of this interaction and in a few cases inbreeding altered the direction of this interaction. In addition, the effect of
inbreeding on the interactive effects of spittlebugs and CMV varied among
monkeyflower families. Our study suggests that inbreeding can have complicated effects on interactions between inbred plants and their environment, but that these effects will vary among genotypes. This will make it
difficult to make predictions about the ecological effects of inbreeding on
small, isolated populations that are forced to self-fertilize.
EVANS, CYNTHIA A.* and JULIE A. ROBINSON. Lockheed Martin
Space Operations, Earth Observations Lab, NASA Johnson Space Center,
Houston, TX. Human-induced changes of western Pacific coastal zones
documented by remotely-sensed imagery.
Even as we struggle to understand the fundamental physical, geological
and biological processes important in coastline development, the rates and
magnitudes of some key agents are changing due to human interactions
with the Earth. Perhaps the best examples of this are the coastal landscapes
of East Asia, which have experienced unprecedented changes in the past
50 years due to human activities. Abundant natural resources (for example,
rich volcanic soils supporting hardwood forests and agriculture; economically viable mineral deposits, and natural harbors) attract large human populations and development. Increasing demands for water and electricity
drive developments such as large dam construction that become key agents
of geomorphologic change. For example, all major rivers in East Asia are
controlled by large dams, and hundreds of new dams are in stages of planning or construction. These developments dramatically change the water
and sediment fluxes along the coast. Impacts include changing rates of
deposition, shifting salinity balances in coastal waters, coastal erosion and
subsidence, changing ecosystems and biogeochemical balances and controls. Another important change agent is coastline modification through
land reclamation and development using dikes, levees, channel construction
and dredge and fill activities. Using remotely sensed imageryastronaut
images of Earthwe provide examples of human activities currently modifying landscapes along East Asian coasts. Here, natural and human factors
are interactive, and combine to create some of the largest rates of change
along coasts anywhere in the world. In a sense, these events provide a
laboratory for viewing natural processes at accelerated rates.
EVANS, DANAE A.,* R. B. PRATT and STEPHEN D. DAVIS. Pepperdine University, Malibu, California. Reddening and regreening: The role
of anthocyanin in water stressed leaves of a sclerophyllous shrub.
Malosma laurina is a dominant species in chaparral shrub communities of
Southern California. We observed reddening of leaves in this species during
a severe drought in winter 2002 where rainfall was the lowest in recorded
history. We hypothesized that severe water stress was contributing to increased anthocyanin production as a photo-protective mechanism for
leaves. To test this hypothesis, we measured water potential (Cx), dark
adapted leaf fluorescence (Fv/Fm), and recovery of fluorescence quantum
yield (Fv/Fm) after exposure to a 120 minute high light treatment (2200
mmol m-2 s-1). Measurements were made under laboratory conditions and
across three experimental treatments in the field over 8 months: red plants
experiencing drought, green plants experiencing drought, and green irrigated plants as a control. Water potential of red and green plants under
stress were not different (p 5 0.11-0.75), but irrigated control plants were
(p , 0.05). Dark adapted leaf fluorescence (Fv/Fm) at predawn and dusk
was not different for red plants compared to green plants (p 5 0.065-0.61).
Exposure of leaves to 2200 mmol m-2 s-1 for two hours caused a greater
decline in Fv/Fm for red leaves compared to green leaves. Red leaves
displayed slower recovery of Fv/Fm compared to green leaves. Red leaves
in situ regreened after the onset of fall rains and demonstrated Fv/Fm recovery patterns similar to those of irrigated plants. Our findings suggest
that leaves which turn red under water stress behave differently than red
and green leaves of senescent plants as previously reported in the literature.
EVANS, R.D.* Stable Isotope Laboratory, Biological Sciences, University
of Arkansas, Fayetteville, AR, USA. Disturbance and invasive species
in arid ecosystems.
Surface disturbance and subsequent invasion by annual grasses and woody
vegetation are dramatically altering biogeochemical cycles in arid ecosys-

102

Abstracts

tems. The primary effect of surface disturbance is removal of the biological

soil crust. This crust can be the primary source of nitrogen input into some,
but not all, arid ecosystems. Disturbance of the crust facilitates loss of
nutrients even in those ecosystems where nitrogen fixation is not important
by increasing erosion. The effects of invasion by grasses and woody plants
on soil carbon and nitrogen are contradictory and seem related to the
amount and seasonal pattern of precipitation. Invasion by annual grasses
may initially enhance retention of nitrogen in disturbed ecosystems by increasing microbial immobilization and decreasing gaseous loss, but in the
long-term an increase in fire frequency results in net loss of nitrogen. Disturbance and invasion also change taxonomic and functional diversity of
microbial communities altering patterns of substrate use causing changes
soil respiration and nitrogen availability.
EVINER, VALERIE T.* Institute of Ecosystem Studies, Millbrook, NY.
Genetic to landscape diversity and ecosystem function: critical insights
from agroecology.
Many experiments in natural ecosystems have investigated the role of diversity in ecosystems. However, we still know relatively little about the
effects of biodiversity on ecosystems because these experiments largely
focus on the effects of plant species richness on a subset of ecosystem
dynamics. Plant diversity and composition have been manipulated in agricultural systems for millennia in order to provide multiple ecosystem
functions. This talk will highlight the critical and unique insights to the
diversity-ecosystem function debate provided by agroecology. Agroecological studies have investigated the roles of multiple levels of diversity,
including genetic, species, functional-type and landscape diversity. These
studies do not simply focus on richness, but examine the effects of the
relative abundance of different components of diversity, as well as their
configuration in space and time. Agroecological experiments provide strong
evidence of the relative importance of diversity versus composition in maximizing different ecosystem processes. They also indicate the length of time
required for diversity and composition to impact different processes, and
the duration of these impacts even after the plant community has changed.
The processes investigated in agricultural systems extend beyond those
typically studied in other diversity experiments, and include productivity,
biogeochemical cycling, nutrient retention, disease and pest regulation, trophic interactions, weed invasion, evolutionary processes, and the fluctuations of these processes over time and changing environmental conditions.
In particular, these studies look beyond plant diversity, and consider the
overall ecosystem impacts of the interactions among a number of different
organisms. The investigation of the role of diversity in ecosystems can gain
a lot from agroecology, particularly because of its focus on the overall
ecosystem- accounting for the simultaneous effects of diversity and composition on multiple ecosystem functions and how these effects may change
under shifting environmental conditions.
EWE, SHARON M.,1,2,* CATHERINE E. LOVELOCK2 and ILKA C.
FELLER.2 1 Smithsonian Marine Station, 701 Seaway Drive, Fort Pierce,
Florida, USA; 2 Smithsonian Environmental Research Center, 647 Contees
Wharf Road, Edgewater, Maryland, USA. Ecological and physiological
responses of mangroves to the interaction between nutrients and salinity.
This study compared the ecological and physiological responses of three
mangrove species (Rhizophora mangle, Avicennia germinans, Laguncularia racemosa) under differing nutrient and salinity regimes to gain a better
understanding of the ecological and physiological tolerances of each species. Field data (from the Indian River Lagoon, Florida) indicated that plant
responses to nitrogen enrichment differed with forest type and among species. Therefore, it was hypothesized that each mangrove species was physiologically adapted to different salinity and nutrient levels and that interspecific differences would affect their distribution and overall abundance
within the community. This study was conducted in a shadehouse with two
daily tidal cycles. Plants were grown in sand for eight months at either 0,
17, 35, 52, 70 parts per thousand (ppt) salinity in combination with a
nutrient regime (either control, nitrogen (N) or phosphorus (P)). We found
that: 1) all three mangrove species grew fastest at 17 ppt salinity (0.5 times
seawater strength) with N-enrichment, 2) salinity tolerance was lowest in
the red mangrove (R. mangle) and highest in the black mangrove (A. ger-

minans), 3) at high salinities (1.5 and 2 times seawater strength), N additions increased plant survivorship and 4) enrichment with P did not increase
plant growth or gas exchange. Significant interspecific N-enrichment differences were observed in the ecological and physiological responses of
the three mangrove species. The results imply that areas of high N and
salinity will be dominated by A. germinans while high N and low salinity
will contribute to L. racemosa being the dominant species. This research
is significant as it provides empirical evidence of the: 1) differences in
ecological and physiological responses among keystone plant species within the mangrove ecosystem in response to nutrient enrichment and 2) mechanisms by which increases in nutrient availability potentially contribute to
patterns of long-term forest structure.
EWERS, BRENT E.,1,* STITH T. GOWER,2 BEN P. BOND-LAMBERTY2
and CHUAKUAN K. WANG.3 1 Department of Botany, University of Wyoming, Laramie, Wyoming, USA; 2 Department of Forest Ecology and
Management, University of Wisconsin, Madison, Wisconsin, USA; 3 Department of Biology, University of Winnipeg, Winnipeg, Manitoba, Canada. Effect of time since fire on growing season length and annual transpiration from boreal black spruce forests.
The microclimatic and successional changes during recovery from stand
killing wildfires impacts growing season length and annual transpiration.
We quantified growing season length and annual transpiration in 2001 and
2002 across a chronosequence of regenerating stands of boreal black spruce
originating from wildfires in 1850, 1930, 1964, 1981 and 1989. In each
stand, we used either Granier-type sap flux sensors (trees greater than 4 cm
DBH) or Kucera-type sap flux sensors (trees less than 4 cm DBH) to quantify sap flux per unit sapwood area and scaled these up to the whole tree
and stand level using site specific allometric equations for leaf area and
sapwood area. The dominant species across the chronosequence consist of
paper birch (Betula papyerifera), trembling aspen (Populus tremuloides),
jack pine (Pinus banksiana), and black spruce (Picea mariana). We measured sap flux in six to 12 trees of each species present in the five stands
for a total of 84 trees across the chronosequence. Analysis of the 2001 data
showed that stand transpiration was not proportional to leaf area index
across the five stands. This was explained by differences in transpiration
per unit leaf area for the four species and changing species compositions
across the chronosequence. In addition, black spruce, which dominated the
two oldest stands, had a declining sensitivity of stomatal conductance to
vapor pressure deficit with increasing age. We hypothesize that growing
season length will decrease with increasing stand age but that species level
differences across the chronosequence will have more of an impact on
annual transpiration than growing season length.
FADEN, MIKE and V.T. PARKER.* San Francisco State University, San
Francisco, CA. Seed dispersal of California bay-laurel (Umbellularia
californica) by western gray squirrels (Sciurus griseus).
Western gray squirrels (Sciurus griseus) act as seed dispersers in mixedevergreen forest in coastal California by scatterhoarding seeds of dominant
tree species including California bay-laurel (Umbellularia californica) and
oaks. Factors influencing dispersal distance and survival of cached Umbellularia seeds were investigated in fall 2001 and 2002, by following the
fate of thread-marked seeds placed under Umbellularia canopies. Factors
measured or estimated in one or both years included seed mass, fruit mass,
cache pilferage, and local abundance of fruits eaten by squirrels including
Umbellularia fruits and acorns. In fall 2001, numbers of seeds cached,
cache survival, and dispersal distance were correlated with the local ambient abundance of fallen Umbellularia fruits. Mean dispersal distance was
greater (11.2-15m) at sites where fruits were scarce or absent, compared
with a mean distance of 1.7-3.7m at sites where fallen fruits were abundant.
Overall, 26% of caches survived intact for more than six months; of these,
84% were at sites where fallen fruit was abundant in fall 2001. Field experiments suggested that burial at depths typical of rodent caches is essential for Umbellularia germination and seedling establishment. In fall 2002,
trends in caching rates and dispersal distance were less clear. It is suggested
that caching behavior may have been influenced by the fact that in contrast
to 2001, oaks (Quercus chrysolepis, Lithocarpus densiflorus) at study sites
produced a large acorn crop that coincided with Umbellularia fruitfall. The
squirrels cached thread-marked acorns as well as Umbellularia fruits.

FAGAN, MATTHEW E.* and DAVID R. PEART. Dartmouth College,

Hanover, NH. Effects of understory invasion by the exotic shrub glossy
buckthorn (Rhamnus frangula) on the regeneration of canopy tree species.
Non-native shrubs are invading many forest understories in the Northeast
and world-wide, yet their effects on canopy tree recruitment remain poorly
understood. In a field survey and experiment, we investigated the impact
of a non-native, tree-like shrub, glossy buckthorn, on the performance of
juvenile tree species differing in shade tolerance. At the seedling stage,
shade intolerant species had much lower growth and survival under buckthorn, while shade tolerant species were affected less. At the sapling stage,
all canopy species grew much less under buckthorn cover, but the effect
was significant only in the last 8 years, implying that buckthorn out-grew
and suppressed neighboring saplings following a recent invasion event.
Buckthorn invasion apparently increases the relative abundance of shadetolerant seedlings while reducing the absolute abundance of the understory
sapling bank. In forests whose understory becomes dominated by buckthorn, the rate of gap-filling by canopy trees is likely to be reduced, and
the species composition of the canopy is likely to be shifted to more shadetolerant species over the long term.
FAGAN, WILLIAM F.1,* and PETER J. UNMACK.2 1 University of Maryland, College Park, MD; 2 Arizona State University, Tempe, AZ. Rarity,
fragmentation, and the scale-dependence of extinction risk in desert
fishes.
Theoretical efforts and small-scale experiments have given rise to the widespread belief that the fewer occurrences a species has or the more fragmented its distribution is, the more vulnerable that species should be to
extinction. Lacking, however, are large-scale studies exploring the connection between these aspects of spatial rarity and local extinction risk across
many species. We present a landscape-level, biogeographic test of this
widely assumed linkage. Using a unique dataset detailing the occurrence
patterns of native freshwater fishes of the Sonoran Desert, we obtained
several measures of spatial rarity for each of 25 species. Some of these
rarity measures were scale-dependent, and one, the "scale-area slope" was
independent of spatial scale. This slope statistic, which characterized the
degree to which species ranges were historically fragmented, proved a
consistently strong predictor of extinction risk, and reached a maximum of
predictability at intermediate scales. At the 100km scale, historic range
fragmentation explained over 90% of the among-species variance in realized extinction risk, and desert fish species with the most fragmented historic distributions were more than nine times more likely to be currently
absent from a given stream reach than were species with the most continuous distributions. In contrast, the number of reaches occupied (as defined
on a series of hierarchical scales) was a significant predictor of extinction
risk only if fragmentation had not already been accounted for and decreased
in importance as scale increased. These findings have three major implications. First, they underscore what a strong link exists between spatial
distribution and vulnerability to extinction. Second, they clarify that the
link exists even at the landscape-level and across an entire biogeographic
fauna. Last, they demonstrate how extintion risk can be a scale-dependent
phenomenon that is affected by aspects of species distributions operating
at both finer and coarser scales.
FALK, DONALD A.* Ecology & Evolutionary Biology, Laboratory of
Tree-Ring Research, University of Arizona, Tucson, AZ, USA. The eventarea relationship: Scale dependence in the fire regime of a New Mexico
ponderosa pine forest.
The fire regime is a multivariate characterization of events in space and
time. Scale dependence of the fire regime is observed by examining the
behavior of metrics of central tendency and higher moments in the distribution of fire intervals. We apply analytical tests adapted from the speciesarea framework to evaluate the form and underlying mechanisms of scale
dependence in the fire regime, substituting fire years for species in a study
of an old-growth ponderosa pine forest in the Jemez Mountains of northern
New Mexico, USA. Accumulation functions and fire- (species-) richness
functions followed the predicted forms, indicating sensitivity to both sample size and area; the latter is the event-area relationship for the number

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103

of disturbance events as a function of area. The inverse of frequency, fire

interval, is correspondingly scale dependent; mean fire intervals varied by
an order of magnitude (2.5-25 yr) for sample areas of 0.5-250 ha. The
interval-area function (IAF) is linear in logarithmic space, and the slope of
the IAF provides an index of spatio-temporal synchrony. Differences in
synchrony of fires are attributable to climatic phases; thus, the slope of the
IAF offers an index of the strength of climate entrainment. Probability
models of the observed fire interval frequency distributions are positively
skewed at small scales (1-50 ha). From first principles in fire ecology, we
justify the lognormal distribution as the default model for fire intervals in
a surface fire regime, considering fires as multiplicative processes involving
a large number of contingent factors. Each of these contributory factors
has its own area-scaling function, producing scale-dependence in the fire
interval probability distribution as a whole. Scale dependence in the parameters of probability distributions generates the observed sensitivity in
metrics of the fire regime to area, creating a synthesis between empirical
observations and underlying ecological and statistical theory.
FALKOWSKI, PAUL G. Rutgers, The State University of New Jersey,
New Brunswick, NJ, US. The coevolution of elemental stoichiometry in
the biogeosphere.
The elemental stoichiometry of C,N,P,S and 13 trace elements varies in
relation to phylogenetic affiliation in 16 species of marine phytoplankton
representing extant taxa that sequentially evolved over 3 Ga, from the Archean through the Cenozic. Our analysis suggests that, whereas average
elemental compositions emerge as a function of scale in aquatic ecosystems, taxon-specific compositions are readily inferred that correspond to
evolutionary trends in the fossil record. The taxon-specific signatures cluster with plastid inheritance, but the genes responsible for the inherited bulk
characteristics were transferred to the host-cells genomes early in the symbiotic association. The light elements have left a geochemical imprint in
the soluble phase of the oceans, while the trace element composition of the
organisms is reflected in black shales and sapropels. This evolutionary
trajectory was further modified by terrestrial ecosystems through the emergence and radiation of grazing mammals and the rise of grasses in the
Cenozoic; processes that strongly influenced the weathering supply of essential elements such as P, Si, and Fe. Our results suggest that the mobilization, acquisition, and transformation of trace elements by marine photoautotrophs has imprinted lithospheric composition since the evolution of
life on Earth.
FANG, WEI.* State University of New York at Stony Brook, weifang@
life.bio.sunysb.edu, Stony Brook, New York. The effects of Acer platanoides on the community and ecosystem dynamics of a natural forest.
The role of a single species, Norway maple (Acer platanoides), on the
community and ecosystem dynamics of a natural forest was evaluated in a
field manipulative experiment. Norway maple is an exotic tree species that
is aggressively invading the interior of natural mesic forests in the northeastern US and is negatively associated with the diversity of native species
in forest understory. I hypothesized that Norway maple changed the community structure either through its superior competitive abilities for resources aboveground and/or belowground or through certain novel ecosystem functions of litter or other manipulation of soil properties. Seedlings
of 3 native and 3 exotic woody species were transplanted under both Norway maple adults and its native congener A. rubrum. The performance of
these transplanted seedlings indicated competitive hierarchy among these
species under various combinations of experimental manipulations and how
such inter-specific interactions translated into community dynamics. A series of ecosystem measurements tested the alternative hypothesis and linked
the role of single species to the dynamics of the whole ecosystem. The
results of the experiment suggested that Norway maple was a superior
aboveground competitor for light, and was also associated with the alteration of soil ecosystem properties without being a superior belowground
competitor. The competitive hierarchy among the understory species was
altered by the invasion of Norway maple. The impacts of an invasive species at the community level and the ecosystem level were interrelated and
inextricable.

104

Abstracts

FANKHAUSER, CRYSTAL L.,* DARREN L. BADE and STEPHEN R.

CARPENTER. University of Wisconsin-Madison, Madison, Wisconsin,
USA. Carbon sources for invertebrate predators in a whole-lake experiment.
Chaoborus, the phantom midge, is an important invertebrate predator in
many lake ecosystems. The carbon in Chaoborus could be derived from
endogenous (autochthonous) primary production of the lake itself, or from
exogenous (terrestrial) sources. Using results from a whole-lake addition
of inorganic 13C, we estimated ecosystem level inflows and outflows of
carbon to Chaoborus spp. Using a simple dynamic model fitted to the data,
we estimated the proportion of Chaoborus carbon that derives from primary
production within the lake. Chaoborus d13C signature changed from -36 to
-11 per mil following the isotope addition. Carbon inputs to Chaoborus
were estimated at 20 mmol C m-2 d-1. Although this is less than half of the
mean gross primary production during the experiment (49 mmol C m-2 d1
), using our model, we found that 20 to 50% of Chaoborus carbon originated from outside the lake (exogenous sources). This represents a significant terrestrial subsidy to this pelagic invertebrate predator and to the
secondary production of this lake ecosystem.
FARGIONE, JOE,1,* RAY DYBZINSKI,1 CHRIS CLARK,1 JANNEKE
HILLE RIS LAMBERS,1 STAN HARPOLE,1 MICHEL LOREAU2 and
DAVID TILMAN.1 1 University of Minnesota, St. Paul, MN; 2 Ecole Normale Superieure, Paris, France. From selection to complementarity:
Temporal trends in a long-term biodiversity experiment.
A long term biodiversity experiment at Cedar Creek Natural History Area
demonstrates that productivity increases as a function of plant species richness. This positive effect of biodiversity on productivity has grown stronger
over the seven years that data have been collected, and may be due to the
increased likelihood of diverse mixtures containing productive species (ie
a selection or sampling effect) or positive interactions among species (ie a
complementarity effect). We partitioned the net biodiversity effect into selection and complementarity components using the statistical techniques of
Loreau and Hector (2001). We found a positive selection effect in the first
two years of data, and a positive complementarity effect in the remaining
five years of data. Thus, the mechanisms controlling the positive relationship between diversity and productivity shifted from selection to complementarity over time. We further partition these selection and complementarity effects by individual species. We show that the complementarity effect of 8 of 16 species in the experiment significantly increased over time.
Our results highlight the importance of long-term experiments in understanding biodiversity effects.
FARINA, JOSE M.,1,2,* JULIE C. ELLIS1 and JON D. WITMAN.1 1 Brown
University, Providence, Rhode Island, USA; 2 CASEB-Pontificia Universidad Catolica de Chile, Santiago, RM, CHILE. Gulls and cormorants
exerting nutrient exchanges between marine and terrestrial ecosystems
at the Isles of Shoals (Maine-USA).
Guano accumulation in seabird colonies often alters soils chemistry and,
through it, can change plant species composition and productivity. Because
the magnitude and extent of these alterations depend on nesting behavior,
major differences are expected between seabird species. We compared soil
nutrient concentrations and plant species composition and performance
from gull (Larus marinus) and cormorant (Phalacrocorax auritus) colonies
in the Isles of Shoals (Maine). Additionally, we assessed the contribution
of marine nutrients to terrestrial ecosystem (using stable isotopic analysis)
and tested plant responses (germination and growth) to soil nutrient concentrations recorded in the colonies. In the colonies, soils have significantly
higher concentrations of nutrients (ammonia, nitrate and phosphate) and
plant species assemblages were dominated by nitrophilic species. These
features were extreme in the cormorant colonies, which had the highest
concentrations of nutrients and where practically no plants occurred. A
greenhouse experiment showed that germination and growth could be inhibited at the highest soil nutrient concentrations (cormorant colonies). The
different effects of gulls and cormorants on terrestrial island ecosystems
were related to nesting densities. We discuss our results in reference other
examples of nutrient exchanges between marine and terrestrial ecosystems.

FARRIS, MARIANNE G.,1,* LISA J. SAMUELSON,1 THOMAS A.

STOKES,1 MARK COLEMAN2 and JOHN BLAKE.2 1 Auburn University,
Auburn, AL; 2 USDA Forest Service, Aiken, SC. Seasonal sap flux density in young versus old loblolly pine stands.
Sap flux density is being monitored over two years in young and old loblolly pine (Pinus taeda L.) stands receiving irrigation to better understand
tree physiological differences in response to increasing trees size and age,
and environmental factors controlling stand level water use. Thermal dissipation probes were installed in three-year-old loblolly pine stands of an
average diameter of 7 cm and winter LAI of 1.0 m2 m22, respectively.
Probes were also installed at different depths and aspects in dominant,
approximately 70-year-old pines ranging in diameter from 32 to 56 cm and
with a winter LAI of 2.0 m2 m22. Both plantations are located on the
Savannah River Site in New Ellenton, S.C. Irrigation is based on PET and
is similar between sites. During the winter months of 2003, average maximum sap flux density was 35 and 25 g m22 s21 in saplings and mature
trees, respectively. However, winter sap flux density was as high as 100 g
m22 s21 in saplings compared to a maximum of 30 g m22 s21 in mature
trees, indicating potentially lower hydraulic conductance in mature trees.
In mature trees, radial patterns in sap velocity were examined at sapwood
depths of 0-10, 0-20, 20-40, 40-60, 60-80 and 80-100 mm. Surprisingly,
winter sap velocity was highest at 40 mm but significant above the 40-60
mm depth. Sap flow was approximately six times higher in mature trees
compared to saplings. Environmental factors controlling stand level transpiration will be compared between seasons and stand ages.
FATH, BRIAN D. and BERNARD C. PATTEN.* Biology Department,
Towson University, Towson, MD, USA. Ecological network analysis:
Weak links and indirect effects in food webs.
A recent paper in Science (Neutel et al. 2002, Science 296:1120-23) uses
traditional food web analysis to demonstrate the importance of "weak links"
in web stability. We consider the described methodology to grossly underestimate the importance of weak links, and also the indirect effects these
carry. The cited paper, regarded by Rafaelli (2002, Science 296: 1035-37)
as a theoretical breakthrough to "modern formal analyses", made no reference to an already existing body of modern formal network analyses
based on input-output (I/O) methods. The two literatures, "food web" and
"network", coexist but overlap little even through they treat the same basic
kind of model structure (mathematically, directed graphs). We consider how
this arose from original conceptions of Darwin vs. Haeckel, and how conceptual dichotomy led to methodological dichotomy. Then we reinterpret
the food web approach in I/O terms to produce a clearer account of energymatter transactions in ecological networks and how indirect relationships
come to dominate through the weak links in transfer processes.
FAUTH, JOHN E. College of Charleston, Charleston, South Carolina,
USA. African dust: Causal agent of coral reef declines, alias of El Nino,
or a mirage?
Caribbean coral reefs declined precipitously in the past thirty years, but
causal agents remained elusive. Recently, trans-Atlantic African dust
storms were hypothesized to contribute to reef declines, but data were not
formally analyzed. I used stepwise logistic regression to show accumulations of African dust were more closely associated with benchmark events
than either the El Nino-Southern Oscillation Index or sea-surface temperature anomalies. Spores of the soil fungus Aspergillus, which kill sea fans,
have been found on African dust, and other pathogens and contaminants
also may be transported on it. However, African dust explained ,25% of
observed variation, suggesting it was not solely responsible for disease and
bleaching. Therefore, researchers should investigate African dust as one of
many plausible contributors to declining Caribbean coral reefs.
FEAGIN, R. A.* Texas A&M University, College Station, TX. Individual
versus community processes and the impact of sea-level rise on pattern
formation in a model of sand dune plant succession.
A cellular automata model of species within a sand dune plant community
on Galveston Island, Texas, USA was utilized to test ecological hypotheses
regarding individual plant interactions and their impact upon community

organization. Simulations demonstrated that both an environmental gradient

and facilitative succession resulted in the formation of characteristic sand
dune patterns. The results showed that the plant patterns were due to individual plant responses to their environment within their local neighborhood, yet these responses were constrained by the history of community
development. The local neighborhood was related to ecological field theory.
The ratio of the scale of environmental variability versus the scale of a
plants local neighborhood distance determined the relative importance of
individual and community level processes in forming pattern. Additional
hypotheses about sand dune pattern formation were tested, where alteration
of the gradient by rising sea-level and global climate change constrained
plants to a narrow habitat between hard structures and the forebeach, resulting in a breakdown of community organization.
FEI, SONGLIN,* KIM C. STEINER, JAMES C. FINLEY, MARC E.
MCDILL and PETER J. GOULD. The Pennsylvania State University, University Park, PA. Regeneration stage stand development in upland Appalachian Forests.
Field measurements of 61 upland forest stands in Pennsylvania were performed on 6370 subplots during 1996 - 2002. All the stands were measured
one year prior to harvest, 41 stands have been re-measured one year after
harvest, 13 stands have been again re-measured 4 years after harvest, and
5 stands have been re-measured 5 years after harvest. In addition, 15 stands
with the age of 6 - 10 years after clearcutting were measured in 2001 2002. Regeneration size and density were used in this study to analyze
stand development patterns and identify measures of regeneration success.
Compared to one year before harvest, 22 percent of the regeneration cohorts had increased in both density and height one year after harvest, 37
percent had increased in height but decreased in density, 12 percent had
increased in density but decreased in mean height, while 29 percent had
decrease both in density and height. At four years after harvest, not surprisingly, all cohorts had increased in height, 38 percent increased in density, and 62 percent had decreased in density compared to one year after
harvest. Stem mortality was observed in some 5-year-old stands, which
indicates that the stem exclusion stage can occur as early as 5 years after
harvest. Dead stems represented up to 51 percent of the total density in
some subplots. A seedling size-density relationship was statistically estimated for these stands. The frontier exhibits a curvilinear relationship between log-scaled maximum density and quadratic mean height. Minimum
seeding density for full site occupancy (B-level stocking) was also modeled
based upon best-fit estimates of tree area ratios. Knowledge of regeneration
development and potential biological frontiers are prerequisites for the development of ecologically sound silvicultural management prescriptions
and realistic predictions.
FELDMAN, DAVID L.* Ethics, Social Values and Commitment as
Conditions for Sustainable Water Management.
There are ethical alternatives for helping policymakers manage water resources in a sustainable manner. Sustainable development is an ethical concept predicated on the ability to encompass economic prosperity and social
capacity. The latter embraces the ability to live a fulfilled existence in
which basic needs are satisfied, and in which the opportunity for personal
growth and fellowship within a larger community that embraces nature is
present. To achieve sustainability, a water management system must: encompass the interests of all watershed stakeholders; protect future generations who are not in a position to affect present-day decisions; preserve
and restore the integrity of natural systems; and empower affected parties.
To achieve sustainability requires adopting a regional approach to managing water that transcends political jurisdictions and more faithfully connects
social institutions and watersheds. Such an approach would: recognize that
water problems in one part of a region affect the welfare of other parts;
acknowledge interrelationships among physical, ecological, social, economic, and institutional factors; provide a comprehensive, coordinated decisionmaking framework to enhance cooperation among jurisdictions, agencies,
and stakeholders; and anticipate sources of conflict before they lead to
impasse. I develop this argument in three ways. First, I show that U.S.
water resources policy has been based on a defective philosophy: a combination of utilitarianism and gratuitous favor. This philosophy has sought
to produce the greatest benefits for the largest number of beneficiaries at

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105

the lowest possible cost to the latter. Second, I analyze the features of three
alternative ethical approaches to water management: covenants, categorical
imperatives, and stewardship. Third, I contend that implementing these
alternatives requires embracing a social learning paradigm that utilizes
adaptive management so we can: 1) recognize previous mistakes; 2) monitor and measure change; 3) adopt mid-course corrections; and, 4) apply
what we have learned to larger, more complex contexts.
FELDMAN, TRACY S.* Biology Department, Duke University, Durham,
NC, USA. Can one plant species rescue another from low-density effects on pollinator visitation and fruit set?
Plants occurring at low densities may receive fewer visits by pollinators,
and may produce fewer seeds than plants occurring at higher densities.
These low density effects may be altered by co-flowering plants if these
plants increase or decrease attractiveness of a focal species to pollinators.
Even if pollinator visits increase, reproductive success of focal plants could
either decrease or increase if co-occurring plants compete for or facilitate
pollination, respectively. I conducted a large scale spatial experiment to
test whether one plant species can rescue another from low-density effects
on pollinator visitation and fruit set, using the common perennial plant
Piriqueta caroliniana as a focal species, and Coreopsis leavenworthii as a
co-flowering species. I set up eight artificial arrays of 10 P. caroliniana
plants each, at four different densities, either alone or evenly interspersed
with the co-flowering species. On each of five days, I recorded all pollinator
visits to each patch in 15-minute observation periods, and visitation sequences of all pollinators within each patch. After each of five temporal
replicates, I counted the number of fruits on each P. caroliniana plant. I
used these data to calculate visitation rate to patches, visitation rate to
individual plants, and percent fruit set, and conducted Analyses of Deviance to test for effects of density and co-flowering species on these variables. The visitation rate of pollinators to patches, the visitation rate of
pollinators to plants within patches, and fruit set all increased with increasing plant density. However, I detected no effect of the co-flowering species,
perhaps due to lower than predicted overlap among pollinator species, or
to high pollinator constancy. Low density effects may have serious consequences for populations (e.g., Allee effects) if plants at low densities do
not produce enough seeds to replace dying individuals.
FERAL, CHRISTIE* and HOWARD EPSTEIN. Environmental Sciences
Department, University of Virginia, P.O. Box 400123, 291 McCormick
Road, Charlottesville, Virginia, USA. Bare ground and bovines: Effects
of grazing history and intensity on savanna soils and vegetation.
Reductions in interspecific competition due to grass removal and shifts in
nutrient availability are key factors in plant community changes linked to
cattle grazing in savannas. We studied the effects of continual and discontinued cattle grazing on soil properties and vegetation composition along
grazing intensity gradients. Plots (100 m2) established at increasing distances (up to 3 km) from a central borehole ran along grazing intensity
gradients at two farms with nearly opposite grazing histories near Ghanzi,
Botswana: a cattle farm grazed until 1994 by wildlife only and a converted
cattle farm supporting only wildlife since 1995. Soil properties and vegetation composition were measured along three 10-meter lines in each plot.
1) Soil pH is significantly lower at the cattle farm (ANOVA, p,.00001),
possibly due to differences in soil parent materials and patterns of urine
and dung deposition. Soil pH decreases with distance at the game farm
(r250.18, p,.0001)and may be related surface calcrete near the borehole.
2) Mean soil crust strength increases with distance at both sites (r250.310.62, p,.0001) and is greater overall at the game farm (ANOVA, p,.0001)
except at a distance of 50m from the borehole. 3) At the cattle farm, a
negative relationship between distance from the borehole and the alongline proportional cover of bare soil (r2 5 0.77, p,0.001) appears correlated
with a similar relationship between distance and grazing intensity. 4) At
both sites, grass density is positively related to distance from the borehole
(r250.69-0.86; p,0.001), while forb density decreases with distance
(r250.23-0.43, p,0.01). Mean forb and woody plant density are greater at
the game farm (ANOVA, p,.05-.0001). 5) Acacia mellifera is a species
associated with bush encroachment. Juvenile A. mellifera density is higher
at the cattle farm (ANOVA, p,0.05). Seventy-two percent of juveniles
grew beneath Grewia flava canopies. These results suggest that grazing

106

Abstracts

intensity predominantly controls proportional cover by bare ground and

grass, as well as soil crust strength. Recovery from cattle grazing pressures
may be dominated by forbs and shrubs, while increases in A. mellifera may
be tied to the presence of facilitating shrubs.
FERNER, MATTHEW C.,* DELBERT L. SMEE and YIN CHANG.
School of Biology, Georgia Institute of Technology, 310 Ferst Drive, Atlanta, Georgia, USA. A field study of blue crab responses to conflicting
chemical cues.
Animals must often balance the conflicting demands of resource acquisition
and predator avoidance. In aquatic habitats where visual cues are limited,
dissolved chemicals can provide useful information about the location of
food and the proximity of predators or conspecifics. Chemical alarm signals
released by threatened or injured prey act to warn other animals of danger
and may permit alerted individuals to alter their morphology, physiology,
or behavior in order to reduce vulnerability to predation. Behavioral responses to alarm signals include fighting, fleeing, or concealment, all of
which could compromise foraging success. Our field study of blue crab
behavior in estuarine tidal creeks revealed a trade-off between responses
to feeding attractants and alarm pheromones. Baited traps containing an
injured crab caught significantly fewer foraging crabs than baited traps
alone. Aging the injured crabs for various periods of time before trap deployment confirmed that alarm signals discourage foraging for at least 8
hours but are no longer deterrent after 22 hours. Additional field tests
confirmed that the alarm signals are both chemical and species-specific.
These data demonstrate that blue crabs will sacrifice foraging opportunities
when confronted with the scent of freshly injured conspecifics. Not only
should such responses to alarm pheromones facilitate predator avoidance,
but associated changes in movement or habitat use could have more farreaching ecological consequences. For example, if crabs are conditioned to
spend less time foraging or to avoid high-risk habitats, these same areas
may provide a refuge for their own prey (e.g., bivalves). On the other hand,
crab predators (e.g., birds) may forage more successfully in areas where
the effectiveness of alarm pheromones is confounded by hydrodynamic
mixing or by elevated background stimuli. Manipulative field studies of
animal responses to multiple chemical cues should help to clarify the processes that regulate trophic interactions within natural communities.
FERRARI, JAMES B.* Wesleyan College, Macon, Georgia. Seasonal patterns of avian diversity in central Georgia.
Seasonal trends in bird species diversity in central Georgia were compared
at two spatial scales using different sampling techniques. At a 0.26 ha
wooded residential area, monthly bird checklists were kept for six years.
In addition, arrival and departure dates of seasonal residents and all dates
for rare or transient species were recorded. The second site, only 4.8 km
away, was a 36 ha portion of a college campus featuring a mixture of
forest, field, and aquatic habitats, where two years of weekly timed censuses were conducted. Weekly species richness was determined for each
site using the observed bird diversity and by interpolation. The timing of
spring and fall migration was virtually identical at the two sites, peaking
from mid-April to early May and then again from the middle of September
through the first week in October. Bird species diversity was lowest in
summer at the residential site, whereas summer diversity at least equaled
winter diversity at the campus site; interpolation indicates that species diversity was greater in summer than in winter at the college. Although the
timing of peak bird diversity during migration was identical at both spatial
scales, diversity of summer residents was highest at the site featuring the
greatest diversity of foraging and nesting habitats.
FERRARI, MATTHEW J.1,* and MARK L. TAPER.2 1 IGDP in Ecology,
The Pennsylvania State University, Universtiy Park, PA, USA; 2 Department of Ecology, Montana State University, Bozeman, MT, 59717. Using
auxiliary data on measurement error to improve estimates in population time series.
Few, if any, populations are monitored perfectly. Population monitoring
often involves a trade-off between precision of the counts and the expense
of the monitoring program. As a result, time series of population counts
incorporate two sources of variability: the process variance inherent in the

population dynamics, and variance due to stochastic measurement error. If

ignored, the variance component due to measurement error will result in
over-estimates of the biological process variance (a quantity of interest in
population viability analyses) and poor precision in estimates of the population trend. Often auxiliary data, in the form of double counts, can be
collected which provides information on the measurement error, independent of the time series and independent of the assumed population model.
These auxiliary data can be combined with the time series observations to
form a joint likelihood which can be maximized to obtain simultaneous
estimates of population trend and the process and measurement variance
components and associated standard errors. We present a justification for
the joint likelihood estimation using a simple example. We also evaluate
the performance of the joint likelihood estimation relative to the mixedmodel estimators for geometric population growth in the presence of measurement error developed by Staples et al. (2003). The incorporation of
auxiliary data through the joint likelihood resulted in a reduction of the
expected confidence interval for population trend as well as the variance
components without sacrificing the coverage rate. Thus, in future monitoring programs, less expensive, imperfect counts could be improved by collecting coincident data on the measurement process itself and calculating
simultaneous estimates of measurement and biological parameters.
FERTIG, WALTER F.,1,2,* ROBERT THURSTON2 and WILLIAM A. REINERS.2 1 Grand Staircase-Escalante National Monument, walt@
xpressweb.com, Kanab, UT, USA; 2 University of Wyoming, Laramie, WY.
Modeling the potential distribution of rare plants in Wyoming.
Predictive modeling of plant distributions rests on the assumption that correlations exist between the presence/absence of a species and selected climate, topographic, substrate, and land cover variables. Once these underlying patterns are determined, maps can be created in GIS that identify all
areas that meet the specific conditions for a given species. Such maps can
be used to prioritize areas for field surveys of rare plants or assist decision
makers in project clearance activities. Using classification tree analysis, we
developed correlational models for 44 Wyoming plant species listed as
BLM Sensitive or Threatened or Endangered under the Endangered Species
Act. Presence/absence of each species was the response variable in the
models and was derived from location records of the Wyoming Natural
Diversity Database and Rocky Mountain Herbarium. Environmental variables, including total monthly precipitation, average monthly air temperature, monthly shortwave radiation, number of wet days, growing degreedays, local topographic relief, bedrock and surficial geology, soils, elevation, and land cover, were used as predictors. Location data were randomly
subdivided into model-building and validation data sets to test the classification success of the final models. We found that the distribution of rare
species in Wyoming was most strongly correlated with specific bedrock
and soil types, but was also influenced by topographic relief, land cover,
and various monthly precipitation and temperature values. Overall, our
models were conservative in the area predicted for these species and typically had low commission error rates. The distribution maps produced by
correlational modeling did an excellent job of identifying areas where rare
species are unlikely to occur and did a good job of highlighting areas of
potential habitat that warrant additional on-the-ground survey.
FIELD, CHRISTOPHER B.* Carnegie Institution, Stanford, California.
Policy implications of carbon-nitrogen interactions.
Carbon-nitrogen interactions have important implications for several aspects of climate change and carbon management policy. Progress on at
least four issues is critically dependent on ecological information. These
are (1) the role of nitrogen in regulating future terrestrial carbon storage,
(2) potential carbon storage from purposeful nitrogen fertilization, (3) climate forcing from methane and nitrous oxide resulting from purposeful
nitrogen fertilization, and (4) prospects for quantifying effects of unintended fertilization with nitrogen and carbon dioxide in purposeful carbon management schemes. Some of the contributions of ecological information can
take the form of improved quantities for modeling and accounting systems.
Others should help structure policies to keep them consistent with a realistic
assessment of the information ecological research can and cannot provide.

FIGUEIRA, WILL F.* Duke University Marine Lab, Beaufort, NC, USA.
(Re?)structuring our approach to source-sink metapopulation dynamics in marine systems.
The use of source-sink metapopulation terminology in marine systems is
steadily increasing. Conservation and management actions have brought
about the need to more fully understand the spatial component of population dynamics in these systems towards the specific end of identifying the
relative "quality" of different areas for the purpose of protection or regulation. While the application of the theory is certainly appropriate and
potentially quite useful, some of the key conceptual features have been lost
or simply misapplied in the translation from terrestrial systems. Of key
importance here is the terrestrial bias towards habitat-level effects versus
the marine bias towards dispersal-level effects. In this study I develop an
analytical framework for conceptualizing and modeling source-sink dynamics in marine metapopulations. This theory is grounded in the work of
terrestrial systems but retooled to account for the important role dispersal
can play in marine systems. The end conceptual and analytical model tracks
patch-level effects on the overall metapopulation and in so doing allows
for us to evaluate the relative contributions of each patch within the metapopulation quantitativelythat is as either a source or a sink. I apply this
framework to a simple metapopulation to demonstrate how the overall connectedness of the system can affect source-sink characterizations, especially
when populations are regulated in a density-dependant manner. The framework I present here for considering source-sink metapopulations in marine
systems combines habitat and dispersal characteristics to present an overall
picture of patch-level contribution to the metapopulation. By linking these
factors together we can evaluate the effectiveness of metapopulation-level
management efforts and come to understand outcomes that would perhaps
not have been otherwise anticipated.
FINE, PAUL V.A.,1,* PHYLLIS D. COLEY1 and YTALO MESONES A.2
University of Utah, Salt Lake City, Utah; 2 Universidad Nacional de la
Amazonia Peruana, Iquitos, Peru, Peru. Herbivory enhances the evolution and maintenance of habitat specialization in Amazonian white
sand and clay forests.
1

Tropical forests include a diversity of habitats, which has lead to specialization in plants. Near Iquitos, in the Peruvian Amazon, nutrient-poor white
sand forests are found immediately adjacent to nutrient-rich clay forests,
each harboring a unique composition of habitat specialist trees. Why does
habitat specialization occur? Our hypothesis is that the combination of impoverished soils and herbivory creates strong selective pressure for plant
defenses in white sand forests. Species that have not evolved high levels
of antiherbivore defense are therefore at a disadvantage in white sand forests and excluded by herbivores. In clay forests, due to the tradeoff between
growth and defense, heavily defended white sand species should grow more
slowly than clay species and be out-competed. To test to what degree herbivores maintain the low overlap in species composition between the two
forest types, we conducted a reciprocal transplant experiment and manipulated the presence of herbivores. We transplanted 880 seedlings from 20
species from seven genera that included phylogenetically independent
white sand and clay specialist species into 22 herbivore exclosures and 22
controls in both white sand and clay forest. After 20 months, clay species
exhibited significantly higher growth rates (both height and leaf area) in
white sand forest than white sand species when protected against herbivores. When left unprotected, clay species average leaf area and height
was less than white sand species averages. In clay forest, white sand species had significantly higher growth rates than they did in their home forest,
but always grew significantly less than clay species. These results are the
first experimental evidence of the impact herbivores have on plant species
distribution in tropical forests and suggest a new way that herbivores influence plant evolution: by sharpening habitat boundaries due to abiotic
factors and thereby increasing the potential for habitat specialization and
speciation.
FINKE, DEBORAH L.* and ROBERT F. DENNO. University of Maryland, College Park, Maryland, USA. Trophic cascades dampened in
complex food webs with intraguild predation.
The prevalence of trophic cascades in terrestrial systems has been debated.
Food web structure is one factor that may mediate the ability of predators

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107

to suppress herbivores and thus indirectly enhance plant primary productivity. In complex food webs with reticulate interactions such as intraguild
predation, the strength of cascading predator effects may attenuate from
one trophic level to the next, precluding a trophic cascade. Thus, trophic
cascades may only occur largely in communities with low species diversity
and with only a few trophic links among species. The objective of our
study was to determine the impact of increasing food web complexity on
the strength of trophic cascades on a mid-Atlantic salt marsh. We manipulated food web complexity by increasing predator diversity, both the number of predator species and the number of intraguild predators present. Four
food webs of varying complexity were established: plants only (Spartina
cordgrass), plants + herbivores (Prokelisia planthoppers), plants + herbivores + single predator (the mirid bug Tytthus), and plants + herbivores +
a predator complex including both strict predators (Tytthus bugs and the
web-building spider Grammonota) and intraguild predators (the wolf spiders Pardosa and Hogna). After two months of interaction, the effects of
the food web complexity treatments on the size of the herbivore population
and plant biomass were assessed. In the simple food web with only one
predator, a trophic cascade occurred. Mirid bugs drastically reduced the
planthopper population and indirectly enhanced plant biomass. In contrast,
in the complex food web with a diverse predator assemblage, suppression
of the planthopper population was diminished precluding a trophic cascade.
This was due in part to the intraguild predation of mirid bugs by spiders.
Thus, reticulate interactions such as intraguild predation can dampen the
strong cascading effects of predation and potentially stabilize food-web
dynamics.
FINKELSTEIN, SARAH A.* and ANTHONY M. DAVIS. University of
Toronto, Toronto, Ontario, Canada. The interpretability of wetland pollen records.
The coastal wetlands of the Great Lakes region have been subject to heavy
anthropogenic stresses over the past two centuries. Protecting and restoring
these wetlands each require an understanding of natural rates and mechanisms of community change, including responses to shifts in climate and
water level. Pollen analysis provides a useful method for studying the longterm dynamics of these systems. Essential to this analysis is the identification of sensitive plant species with interpretable pollen signatures. Using
contemporary pollen rain studies, detailed morphological analysis of selected taxa, and the literature on species biology, we conclude that wetland
pollen records can be useful in detailing long-term changes in wetlands.
We emphasize here the representation in the pollen record of commonly
dominant plants in Great Lakes coastal wetlands. Typha latifolia, T. angustifolia, T. x glauca and Sparganium eurycarpum can be separated in
the pollen record on the basis of morphology. These have different sensitivities to water depth and different impacts on wetland communities. Typha pollen production, however, can be low relative to the abundance of
plants; pollen percentages as low as 7% can indicate monodominant Typha
stands. Zizania aquatica can be distinguished palynologically from other
stand-forming grasses on the bases of size and exine sculpturing. Its pollen
production is large relative to its plant abundance. Although Phragmites
pollen is distinctive, the habitat range of this species is broad enough that
its value as a paleoecological indicator is ambiguous.
FINZI, ADRIEN C.,1,* EVAN H. DELUCIA,2 JOHN LICHTER,3 ROBERT L. SINSABAUGH4 and WILLIAM H. SCHLESINGER.5 1 Boston
University, Boston, MA, USA; 2 University of Illinois, Urbana, IL, USA;
3
Bowdoin College, Brunswick, ME, USA; 4 University of New Mexico,
Albaquerque, NM, USA; 5 Duke University, Durham, NC, USA. Progressive nitrogen limitation to ecosystem function in the Duke Forest FACE
Experiment.
Experimental increases in the concentration of atmospheric carbon dioxide
increase plant growth and net primary production (NPP). In the absence of
a concomitant and stoichiometrically balanced increase in N inputs to an
ecosystem, the increase in NPP under elevated CO2 should increase the C:
N ratio of the entire ecosystem. This in turn, should lead to a progressive
limitation of ecosystem function (e.g. net ecosystem production, NEP) by
N. This suggests a short-term response of terrestrial ecosystems to increases
in atmospheric CO2 concentrations. We tested this simple model for ecosystem development using data collected from the first six years of forest

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Abstracts

growth under elevated CO2 in the Duke Forest FACE experiment. Initial
increases in NPP increased the C:N ratio of the O-horizon and top layers
of mineral soil. The increase in the C:N ratio of these horizons appears to
have increased microbial-N immobilization as indicated by the lower concentrations of inorganic N in soil extracts and a decrease in the rate of net
mineralization. Analyses of extracellular enzyme activities suggest significant microbial-N limitation in the O-horizon. Additionally, there has been
a large immobilization of N in woody biomass under elevated CO2. Ecosystem mass balance shows that the accumulation of N in woody biomass
and O-horizon exceeds the rate of N input to the soil system via atmospheric deposition. Collectively these data suggest that labile N pools in
soils are being depleted by the increase in the C flux into this ecosystem.
Consistent with the progressive N limitation hypothesis, these data predict
that down-regulation of NPP should be forthcoming.
FITZHUGH, ROSS D.,1,* GARY M. LOVETT,2 KATHLEEN C. WEATHERS2 and MARY A. ARTHUR.3 1 University of Illinois, Urbana, IL; 2 Institute of Ecosystem Studies, Millbrook, NY; 3 University of Kentucky,
Lexington, KY. Canopy tree species, nitrogen fertilization and soil solution chemistry in forest ecosystems of the Catskill Mountains, New
York.
The accumulation of N in soils from elevated rates of atmospheric N deposition may enhance rates of nitrate leaching and thus accelerate the loss
and depletion of base cations from the soil exchange complex. The objectives of this study were to elucidate the effects of N addition on the concentrations of N, Ca, Mg, Na and K in soil solutions at plots in the Catskill
Mountains of New York. Soil solutions were collected in the mineral soil
below the rooting zone at reference and treatment plots (addition of granular ammonium nitrate at a rate of 50 kg N/ha/yr). Plots were monospecific
stands of American beech, eastern hemlock, sugar maple, northern red oak
or yellow birch. Leaching of total dissolved N and nitrate below the rooting
zone increased by factors of approximately 4 and 9, respectively, in response to the fertilization treatment, and the strongest response was evident
for American beech and the weakest for red oak. Given that beech typically
has leaf litter chemistry associated with relatively low rates of N cycling,
the strong response of N loss to N addition at beech stands was surprising
and may have been related to beech bark disease. Although fertilization
increased the leaching of ammonium in the mineral soil by a factor of 6,
soil solution ammonium concentrations were an order of magnitude smaller
than nitrate in treatment plots. Dissolved organic N was unaffected by the
treatment in the mineral soil solutions. Leaching of Ca and Mg below the
rooting zone were increased by 190 and 160%, respectively, and exhibited
the greatest response under sugar maple and the least under hemlock and
oak. Soil solution K and Na were not affected by the fertilization treatment.
Our results suggest that the responses of N and nutrient base cation loss
to N deposition may be strongly influenced by the composition of canopy
tree species.
FLEMING, RICHARD A.1,* and JEAN-NOEL CANDAU.2 1 CANADIAN
FOREST SERVICE, SAULT STE. MARIE, ONTARIO, CANADA; 2 ONTARIO FOREST RESEARCH INSTITUTE, SAULT STE. MARIE, ONTARIO, CANADA. Spatio-temporal dynamics & landscape ecology of
spruce budworm disturbances & the implications for a changing climate.
The spruce budworm (SBW) represents a dominating disturbance factor in
North Americas boreal forests and during outbreaks trees are often killed
over vast areas. This extensive tree mortality constitutes an important fire
hazard and shifts the forest toward younger age-classes which contain less
biomass (and sequester less carbon). We developed spatially explicit regression tree models from Ontarios historical records to describe how climate and forest structure have influenced defoliation patterns. When climatic predictions corresponding to the next outbreak (2003-2038) were
input, the models forecast a general northward extension and intensification
of defoliation, with defoliation declining in southeastern, and particularly
in southwestern Ontario. To understand how climate affects post-outbreak
fire hazards, we studied the fire and defoliation records for the 19,950 km2
experiencing both wildfire and SBW defoliation. Analysis of the spectra of
time lags between the two disturbance types indicated that fires occurred
3-9 years after an outbreak disproportionately often. This "window of op-

portunity" for wildfire starts later after SBW outbreak and lasts longer in
western than in eastern Ontario. In addition, 7.5% of the areas containing
SBW killed trees were burnt in western compared to 4.8% in eastern Ontario. We suggest that these geographical differences result at least partly
from slower decomposition of dead fuels in the drier climates of western
compared to eastern Ontario.
FLETCHER, ROBERT J.1,2,* and ROLF R. KOFORD.2 1 University of
Montana, Missoula, MT; 2 Iowa Cooperative Fish and Wildlife Research
Unit, Ames, IA. Interannual climate variation in wetlands influences
distribution and demography of wetland songbirds.
Annual variability in abiotic factors can be pronounced, especially in systems relying on precipitation, such as arid regions and prairie potholes. We
report how annual variation in precipitation between 1999-2002 in the Prairie Pothole Region of Iowa affected both density and reproduction of two
interspecific competitors: yellow-headed blackbirds (Xanthocephalus xanthocephalus) and red-winged blackbirds (Agelaius phoeniceus). During dry
years yellow-headed blackbirds, an obligate wetland-breeding species,
showed a marked reduction in density and a complete reproductive failure,
in which none of the nests we monitored fledged young. This reproductive
failure was attributed primarily to predation, which was negatively correlated with water levels in wetlands. Conversely, red-winged blackbirds, a
facultative wetland-breeding species, showed little variation in density and
nest success. Both species exhibited similar patterns of reduced clutch size
and later nest initiation dates in dry years, measures often tied to bottomup effects of food availability and/or age of individuals. Yet top-down
effects of nest predation had stronger population implications, because lower clutch size did not result in fewer young fledged per successful nest.
Ultimately, these effects could be exacerbated with either directional trajectories of decreased precipitation or increased variability in precipitation,
owing to global climate change.
FOLLSTAD SHAH, JENNIFER J.* and CHELSEA L. CRENSHAW. University of New Mexico, Albuquerque, NM, USA. Allometric scaling of
whole stream metabolism.
Metabolic rate can be defined as the rate at which an organism utilizes
energy from the environment, converts it biologically, and allocates it to
growth, reproduction and maintenance. Recently derived allometric scaling
relationships for the metabolic rates of individual organisms have shown
that metabolic rates are determined by temperature and body size. Similar
scaling relationships appear to hold for ecosystem metabolism, as seen in
the relationship between temperature and carbon flux in forested ecosystems. We have explored whether the relationship between ecosystem metabolism and temperature can be seen in stream ecosystems. Whole stream
respiration rates are measured as the change of CO2 or O2 from dusk to
dawn over a known area. Data compiled from various published works on
stream respiration were transformed to a common currency measuring
nighttime stream respiration (mg C d-1) and plotted against the inverse of
temperature in degrees Kelvin multiplied by the Boltzmann constant. The
model predicts that the slope of this regression represents the activation
energy for metabolism across assemblages of a variety of organisms. Biological activation energies range between 0.2 1.2 eV, and averages 0.6
eV. Our data fall within this range both when taken together (N 5 44, slope
-0.599, r2 5 0.22) and when separated into autotrophic (P:R . 0.5) and
heterotrophic (P:R , 0.5) stream ecosystems (N 5 18, slope 5 -0.997, r2
5 0.66 and N 5 26, slope 5 -0.404, r2 5 0.17, respectively). We further
propose that this model can be expanded to explain rates of nitrogen uptake,
nitrification, and denitrification across varied stream ecosystems.
FORBES, ANDREW E.* and ANTHONY R. IVES. University of Wisconsin-Madison, Madison, WI. Contrasting effects of heterogeneous host
density on emerging wildlife diseases, such as chronic wasting disease.
Using computer simulations, we show that spatial variability in wildlife
host density may have dramatic effects on rates of disease spread through
a population. The nature of these effects varies greatly, depending on the
mechanism responsible for generating differences in local host densities.
In our model, diseases spread faster through systems in which spatial heterogeneity in host density is created by variability in levels of local re-

sources than in comparable systems with uniform resource distributions. In

contrast, systems in which spatial heterogeneity in host density is created
by variability in local migration rates are not prone to this increased rate
of disease spread. This model demonstrates the importance of considering
variation in local density when managing wildlife and, more importantly,
the need to understand the underlying processes responsible for variation
in local density. The broad conclusions of this study may be of use in
deriving management strategies for slowing the spread of emerging diseases of wildlife.
FORD, CHELCY R.,1,2,* MARY ANNE MCGUIRE,1,* ROBERT J.
MITCHELL2,* and ROBERT O. TESKEY.1,* 1 University of Georgia, Athens, GA; 2 Joseph W. Jones Ecological Research Center, Newton, GA. The
effects of evaporative demand on the radial distribution of sap flux in
tree stems.
Earlier work has shown that the radial pattern of sap flux in conifers with
deep sapwood can change from tree to tree and throughout the day as
evaporative demand changes. We evaluated whether the shape of the sap
flux radial profile could be predicted temporally as a function of vapor
pressure deficit (D) and photosynthetically active radiation (PAR) in a plantation of mature loblolly pine, Pinus taeda L., growing in Georgia, USA.
Granier-style heat dissipation probes spanned the entire sapwood radius
with measuring points located every 2 cm in ten trees. These measurements
were simultaneously validated with independent 2 cm probes. Temperature
differentials were recorded every 5 min at all measurement points, and
measurements were averaged into 15 min values. We found that the radial
pattern of sap flux varied predictably with evaporative demand. Under higher evaporative demands, the inner xylem conducted disproportionately
more water than the outer xylem compared to times when evaporative
demands were lower. By knowing D and PAR, stem sap flux at any radial
point in the xylem can be predicted, and the total radial flux profile can be
used to accurately scale single point-measurements to whole-tree water use.
These results indicate that the conducting xylem of pines, and perhaps other
conifers, should be considered a variable-area flow path that changes dynamically depending on the atmospheric evaporative demand.
FORD, JESSE,1,* SUSAN ALLEN-GIL,2 JOHN SEIGLE,1 JOSHUA
NASHAGNIK, JOEB WOODS and MAASAK AKPIK. 1 Oregon State
University, Corvallis, OR; 2 Ithaca College, Ithaca, NY. Listening to fish:
Science and Wisdom on the Alaskan Arctic Coastal Plain.
Our collaborative work on routes of contaminant entry into Arctic Alaskan
ecosystems began when funding was approved for Inupiat whitefish experts
and translators to work with project scientists and students. The first project
meeting was held several months before the first field season and affected
basic aspects of the study design, including site selection and target species.
Reciprocally, the scientific studies have revealed unexpected information
ancillary to project goals, such as the discovery of a surprisingly deep hole
in one commonly used subsistence lake, or the nature and timing of marine
experience (as revealed by otolith microchemistry) of whitefish caught in
inland lakes. Specifically, one whitefish species appears to be exploiting
brackish or marine environments only infrequently (least cisco, Coregonus
sardinella, iqalusaaq). By contrast, a larger species (broad whitefish, C.
nasus, aanaakliq) is a frequent and heavy user of such environments, and
interestingly has lower Hg concentrations than even the generally low levels found in least cisco. Field work in most cases was done collaboratively
at subsistence camps, where catches proved to be higher than in lakes
whose position was estimated from state files or chosen by professional
judgement. In field camp, differences between the scientific team and local
experts generally related to pace and timing of activities. At the request of
the Kuukpikmiut Subsistence Oversight Committee we did not use helicopter access in our last season but rather travelled by snowmachine, which
was both cost-effective and also more scientifically productive, as it returned control over field activities to the on-site team instead of linking
them to pre-set helicopter schedules.
FORD, MARY E.,* ALAN R. BERKOWITZ, KATHRYN E. LOWE and
DANIEL
B.
NOVAK.
Institute
of
Ecosystem
Studies,
FordM@ecostudies.org, Millbrook, NY. Ecology Field Programs for K12 school groups at the Institute of Ecosystem Studies.
The mission of the Education Program at the Institute of Ecosystem Studies
(IES) is to advance the theory and practice of ecology education, fostering

Abstracts

109

ecological literacy - an understanding of and ability to use the ideas and

methods of ecological science for oneself - in youth, teachers, young scientists, professionals, decision makers and the general public. As part of
this program, the Institutes Ecology Field Programs (EFPs) support local
science curricula for kindergarten through 12th grade and provide students
with opportunities to learn about ecological concepts and processes. During
the 2001-2002 school year EFP participants included 4,200 students in 134
school groups coming from 27 different towns (including rural areas) and
cities in New York and Connecticut. The EFPs use unique resources at IES,
such as the greenhouse, permanent plots along trails, study stations at a
pond, a maple sugaring system, and observation groundwater wells, and
include classroom and field work. Through the EFPs students participate
in experimental design, hypothesis testing and data collection. This poster
will share insights about balancing alignment with local and national standards with addressing concepts and processes from the cutting edge of
ecological research. Assessment of the effectiveness of the EFPs, measured
through teacher surveys and embedded assessment of student learning, will
be discussed. Teachers surveys indicate that the EFPs are successful in
communicating content, and in providing meaningful learning experiences
relevant to classroom objectives. Student learning, assessed through a variety of tools including the use of pre- and post-program drawings, suggests
that students consistently gained knowledge from the EFPs. For instance,
drawings from our plant ecology program show that students obtained enhanced understanding of a plants interactions with its environment. The
EFPs serve as a model of how an institution with a focus on ecological
research can contribute to science education within schools.
FORKNER, REBECCA E.,1 ROBERT J. MARQUIS1 and JOHN T. LILL.2
Department of Biology, University of Missouri - St. Louis, St. Louis, MO;
2
Department of Biological Sciences, George Washington University, Washington, D.C. Feeny revisited: Condensed tannins and leaf-chewing insect communities of Quercus.

Community level oak-tannin-insect patterns have been largely unexplored

since the ground-breaking work by Paul Feeny. For herbivores of black
and white oak (Quercus velutina and Q. alba, respectively) in the Missouri
Ozarks, we tested the hypothesis that abundance and richness of leaf-chewing herbivores would be negatively correlated with foliar condensed tannin
concentration (CTC). In 2001, we quantified foliar CTC and nitrogen in
the understory and canopy of these two species and simultaneously sampled their herbivores at three times: in May as leaves were expanding, and
in June and August when leaves were fully expanded. Of the more than
30 common species encountered, only two species, Acronicta increta (Noctuidae) and Attelabus sp. (Curculionidae), both oak specialists, were negatively correlated with CTC in the canopy of Q. alba. One additional species, Chionodes pereyra (Gelechiidae), also an oak specialist, was marginally negatively correlated with CTC in the understory of Q. velutina. In
the understory, species richness of spring-feeding herbivores was negatively correlated with CTC for Q. velutina. In the canopy, both total insect
density and richness of August species on Q. alba were negatively correlated with CTC. Species determined to be understory "specialists" (i.e.,
tannin avoiders, as CTC was lowest in the understory) were not negatively
correlated with CTC when feeding in the canopy on foliage with high CTC.
Overall, our results indicate that 1) specialists were more likely than generalists to show negative correlations; 2) both early and late season fauna
were impacted by tannins; 3) species were more likely to show responses
to CTC when feeding on Q. alba and when feeding in the canopy, and 4)
CTC contributed to differences in community structure between forest strata and between oak species. Our results support Feenys original view that
tannins play an important role in structuring the herbivore fauna of Quercus.
FORSGREN, TODD,* MICHAEL F. PALOPOLI and JOHN LICHTER.
Bowdoin College, 6500 College Station, Brunswick, Maine, USA. Ectomycorrhizal composition and diversity during primary succession on
coastal Lake Michigan sand dunes.
Coastal dune chronosequences provide useful systems in which to study
patterns of primary succession. We examined the concomitant development
of plant and ectomycorrhizal communities across a well-established dune
chronosequence bordering northern Lake Michigan. Ectomycorrhizal fungi

110

Abstracts

from white pine seedlings were sampled on seven dunes ranging in age
from 30 to 800 years in age. Fungi were sorted by morphotype, and the
primers ITS1-F, ITS4, and ITS4-B were used to amplify and sequence their
DNA. The DNA sequence data were compared with a database developed
from herbarium specimens (Bruns et al. 1998) to identify the ectomycorrhizae to species. We found that species diversity was low during the first
100 years of forest ecosystem development, but increased rapidly to near
peak diversity after this initial period of low diversity. These results are
consistent with an early and late stage model of ectomycorrhizal succession. We hypothesize that colonization constraints delay ectomycorrhizal
establishment of many fungal species.
FORSHAY, KENNETH J.* and EMILY H. STANLEY. University of Wisconsin Madison, Center for Limnology, Madison, WI, USA. Nitrogen removal in water bodies of the Wisconsin River floodplain following
flooding and experimental nitrate additions.
NO3--N pollution in large river systems is a significant problem in the
agriculturally dominated landscape of the Midwestern United States. Numerous works suggest that floodplains are potential sinks for riverine N.
However, descriptions of the rate and extent of N removal in temperate
floodplain systems are scarce. Our goal was to identify patterns of N change
in floodplain water bodies during a flood pulse and to identify the processes
that drive N change. We monitored four floodplain water bodies of the
Wisconsin River floodplain in Sauk County Wisconsin for changes in NO3-N following a large flood event in April 2001. Water body NO3--N fell
from 1.09 mg/l (sd50.24) at peak flooding to less than 0.001 mg/l
(sd50.002)seven days later. Two NO3--N addition experiments in 2002 corroborated this pattern of rapid NO3--N loss. Denitrification potential ranged
from 57.60 to 8348 nmols N m-2hr-1 in acetylene block bioassays. Combining the results of NO3--N additions with denitrification bioassays, a mass
balance approach demonstrated that denitrification was responsible for 30
- 100% of the total NO3--N loss observed in floodplain water bodies. Furthermore, experimental NO3--N additions indicate that a small fraction of
NO3--N was transformed to N species that are retained on the floodplain
including: NH4+-N (3.4-11.3%), particulate N (6.3-6.6%), and dissolved
organic N (3.6-10.4%). Our results demonstrate that denitrification on the
Wisconsin River floodplain removes a considerable proportion of N derived
from the riverine flood waters.
FOSTER, BRYAN L. and TIMOTHY L. DICKSON. University of Kansas,
Lawrence, Kansas, USA. Multiple limitations on plant species richness
in a grassland community.
Local species richness is determined by the balance of local colonization
and extinction. Although extinction is largely governed by local ecological
processes, colonization may be determined by: (i) local processes that govern community susceptibility to invasion and (ii) broader-scale processes
that determine the pool of available colonists and rates of immigration. The
theory of fluctuating resources (Davis and Grime 1999) suggests that the
invasibility of a community will vary over time with resource availability,
such that windows of opportunity for colonization (invasion windows) arise
periodically when resource supply exceeds demand. We conducted a resource manipulation and propagule addition experiment in grassland to: (i)
examine impacts of resource availability on community invasibility; and
(ii) examine the extent to which colonization and species richness are limited by resources and the availability of establishment microsites versus
the availability of colonists from the regional species pool. Consistent with
the theory of fluctuating resources, community invasibility (measured as
the proportion of 32 sown species that successfully invaded a plot) was
significantly increased by irrigation and by the application of experimental
disturbances. Colonization rates and species richness were significantly enhanced by irrigation, disturbance and the addition of seeds. These results
suggest that species richness is jointly limited by local and regional processes in this grassland and that rates of colonization will be maximized
when periods of maximum invasibility coincide with periods of sufficient
propagule input.
FOSTER, SARA K.,* RICHARD N. MACK and R. ALAN BLACK.
Washington State University, Pullman, WA. The Barberry Eradication
Program in Whitman County, Washington: A reassessment.
The eradication of an invasive alien species, i.e., the total destruction of
all individuals at least below the level of detection, remains the most de-

sirable but a seldom attained goal in combating biological invasions. Eradication usually is possible only with small populations within well-defined
boundaries. However, the most ambitious alien plant removal campaign
ever undertaken, the Barberry Eradication Program, sought to extirpate
Berberis vulgaris, the alternate host for Puccinia graminis (stem rust),
across much of the northern U.S. In Whitman County, Washington, Barberry removal was both thorough and apparently exhaustive, as recorded
in the Programs exceptionally detailed field survey records dating from
1944 until 1978. Eradication teams searched the surroundings of all buildings of hundreds of farms and wood-lots in the county, seeking to destroy
all barberry, regardless of size or reproductive state. Barberries were initially destroyed with applications of salt, kerosene, and later with ammonium sulfamate, 2,4-D and 2,4,5-T. The Program ultimately destroyed
48,673 barberry plants on 1,373 properties in the county. Relying on the
original field survey records, in 2002-2003 we re-visited 100 sites that had
supported barberry as recently as 1978. We detected only eight shrubs at
four sites, representing a remarkably low level of occurrence 25 years after
cessation of the removal program for an invasive species that was once
widespread and abundant in the county. Virtually total eradication of a
widespread plant invader can be attained, provided a straightforward protocol is scrupulously followed: exhaustive and comprehensive field surveys, rapid destruction of all plants upon their detection, and long-term,
repeated searches for remaining or newly emergent plants.
FOSTER, TAMMY E.* and PAUL A. SCHMALZER. Dynamac Corporation, Kennedy Space Center, FL. The effect of season of fire on the
recovery of Florida scrub.
Florida scrub is a xeromorphic shrubland that is maintained by frequent
fires. Historically, these fires occurred during the summer due to lightning
ignition. Today, Florida scrub is often managed by the use of prescribed
burning. Prescribed burning of scrub has been implemented on Kennedy
Space Center/Merritt Island National Wildlife Refuge (KSC/MINWR) since
1981, with burns being carried out throughout the year. The impacts of the
season of burn on recovery are not known. Long-term monitoring of scrub
regeneration has been conducted since the early-19809s at KSC/MINWR
using permanent 15 m line-intercept transects. We obtained data from eight
transects that were subjected to a winter burn in 1986 and a summer burn
in 1997 and compared the recovery of the stand for the first five years
postburn. There was no difference in height growth between the winter and
summer burns; both were approximately 85 cm tall five years postburn.
Initially, the summer burn had a larger percentage of bare ground, but
within a year postburn the amount of bare ground was similar. Total percent
cover ( . 0.5m) was lower during the recovery from the 1997 summer
burn than from the winter burn, whereas total percent cover ( , 0.5m) was
greater. Some differences were found in recovery of the dominant species,
e.g., percent cover of Quercus geminata was lower after the summer burn
than the winter burn.
FOUFOPOULOS, JOHANNES,1,* MARMADUKE KILPATRICK2 and
ANTHONY R. IVES.2 1 University of Michigan, Ann Arbor, MI, USA;
2
University of Wisconsin, Madison, WI, USA. Extinctions of island reptiles under the combined effects of habitat fragmentation and a warming climate.
We examine the effects of a natural fragmentation process caused by rising
sea levels on the reptile communities of the Aegean Sea islands (Greece).
Following the formation of the islands since the end of the last ice age,
reptile populations have been disappearing in a predictable manner dependent on island size and age of isolation. We calculate extinction rates for
each species taking historical, geographical and phylogenetic non-independecies into account. Taxa with lower population densities, habitat specialists and species with more northern current distributions have higher extinction rates even after correcting for confounding factors. We show that
the elevated extinction rates of northern taxa are driven by their preferences
for northern habitats that have also been disappearing at disproportionately
high rates from the islands. As a result, habitat fragmentation under the
warming climatic conditions prevailing since the last glacio-pluvial maximum has resulted in disproportionate disappearances of northern species
from the region. Implications of these results for effective reserve design
under an anticipated warmer future climate will be discussed.

FOX, GORDON A.,1,* PAM TINGIRIS1 and BRUCE E. KENDALL.2

University of South Florida, Tampa, FL, USA; 2 University of Califonia,
Santa Barbara, Santa Barbara, CA, USA. Power analysis for matrix transition models.

Matrix transition models are a key tool for population biologists seeking
to make inferences about population growth rates and their components,
and about how changes in vital rates may affect growth rates and future
population sizes. As a result, these models play a large role in subdisciplines ranging from studies of natural selection to population viability analyses. We have conducted numerical simulation studies of how sample sizes
and the number of censuses in a study affect a number of quantities including the estimates of the matrices themselves, the growth rates l, eigenvectors, and senstitivities and elasticities. Necessary sample sizes and
numbers of censuses vary with the type of life history (structure of the
matrix). Transition probabilities near zero require large sample sizes but
can strongly affect estimates of many matrix quantities. These power analyses are likely to be useful in guiding study design; our code for conducting
them will be made publically available.
FOX, JENNIFER A.* and NELSON G. HAIRSTON. jaf38@cornell.edu,
Cornell University, Ithaca, NY. Temporal patterns of genetic change in
recently established Daphnia populations.
Much theoretical and empirical research has focused on the long-term genetic effects of the founding of new populations. Few studies, however,
have examined the short-term dynamics of genetic change in populations
immediately after establishment. Diapausing eggs of Daphnia remaining in
lake sediments provide a means of reconstructing population history and
fine-scale temporal changes. We examined the genetic trajectories of Daphnia mendotae populations that recolonized two lakes in upstate New York
in recent years. We compared patterns of genetic change in a population
founded by a large number of individuals hatching from a local egg bank
with those in a population founded by relatively few individuals dispersing
into the lake from other sources. We measured variation at microsatellite
loci in eggs stored in lake sediments to characterize population genetic
structure in each population at different time points following population
establishment. Preliminary results suggest that populations within a lake
are genetically differentiated over time and there is little evidence of a
genetic bottleneck in the population founded by dispersal. The results of
this study will help to determine the influence of a standing egg or seed
bank on genetic diversity and contribute to our understanding of how genetic variation is maintained in natural populations.
FOX, JEREMY W.* and CRISTINE BARRETO. Imperial College, Silwood Park, Ascot, Berkshire, UK. Effects of resource diversity on the
outcome of resource competition.
Many classical studies of resource partitioning demonstrate diet differences
among coexisting consumers, but demonstrating that observed diet differences are either necessary or sufficient to explain coexistence is difficult.
Here we take an alternative approach, by manipulating the diversity of
resources (bacterial taxa) available to a pair of ciliate bacterivores coexisting in laboratory microcosms. Classical theory and intuition suggest that
reducing bacterial diversity should reduce the scope for resource partitioning, and therefore coexistence. However, recent theory suggests that the
intensity of competition may be a non-linear function of resource diversity
when resources are self-reproducing. Self-reproducing resources can be
driven extinct by their consumers. The equilibrial intensity of competition
among consumers will depend on which resources persist at equilibrium,
which in turn depends in a complex fashion on initial resource diversity
and composition. We crossed four ciliate combinations (no ciliates, Tetrahymena, Colpidium, and both together) with five bacterial combinations
(monocultures of each of three taxa, all three together, and all three plus
10 additional taxa). We sampled ciliate and bacterial abundance and composition after four weeks (5dozens of ciliate generations). Bacterial and
ciliate composition have complex, reciprocal effects on one another. Resource partitioning only partially explains the observed coexistence of these
ciliates. We speculate on the nature of other mechanisms of coexistence,
and suggest that loss of biodiversity from one trophic level may have surprising consequences for biodiversity on adjacent trophic levels.

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FOX, JESSICA A.,* ADAM DIAMANT and ADAM KLEIN. EPRIsolutions, San Rafael, CA, USA. Species banking programs as a market
incentive for the recovery of T&E species: Indiana bat case study.
The ESA has been effective in identifying T&E species, but less successful
in ensuring species recovery. New incentives are necessary to encourage
landowners to create and/or enhance habitat for species recovery. One such
incentive is the development of bank and trade programs. A private party
requesting an incidental take permit could purchase species credits from
pre-established conservation banks, to provide mitigation for the take. We
evaluated the potential for establishing an Indiana bat conservation bank
in Ohio. We evaluated the potential to develop summer bat habitat, developed a methodology for determining bat credits, provided estimates of bat
credit value, and looked at the regulatory climate for the establishment of
species bank and trade programs. A breeding colony (composed of 40-300
individuals) and 10 acres with one primary roost tree was defined as a
credit unit, resulting in 1,200 potential Indiana bat credits. We estimate that
Indiana bat credits would be worth $20,000-$50,000, based on land value
and species credit prices in the U.S. Preliminary interviews with regulatory
agency personnel indicated a reluctance to adopt this approach. Pilot studies
on efficacy of species bank and trade programs will be necessary to evaluate the utility of such methods.
FRAGOSO, JOSE MANUEL V.1,2 and KIRSTEN M. SILVIUS.1,2 1 ESFSUNY, Syracuse, NY, USA; 2 Wildlife Conservation Society, New York,
NY, USA. Selective pressure on palm seed defenses: Interactions
among seed predators.

FRANCIS, TESSA B.* and DANIEL E. SCHINDLER. University of

Washington, Seattle, WA, USA. Consequences of lakeshore development
for littoral habitats in the Pacific Northwest.
Allochthonous inputs from riparian forests are critical to aquatic ecosystem
processes. One such key input is coarse woody debris (CWD). Much is
known about the function of CWD, and the consequences of its removal
by humans, in fluvial ecosystems. The function of CWD in lakes is less
well understood. Residential development on lakes in the Midwestern United States has been shown to reduce CWD densities in littoral habitats, but
similar patters have not been described for the Pacific Northwest. Here we
present results from a survey of 17 lowland lakes in western Washington
State and southern British Columbia. Across all lakes, residential development resulted in decreased densities of riparian forest and CWD in littoral habitats. Furthermore, organic matter content in littoral sediments declined with human density, and was positively correlated with CWD on a
whole-lake scale. The decoupling of this key terrestrial-aquatic linkage by
human development has important consequences for littoral nutrient cycles
and whole lake trophic interactions.

IT

A
W

Recent studies emphasize the selective pressure placed by seed mortality

agents on fruit structure, seed number and seed size in the tropics. In South
and Central America, the principal sources of mortality for seeds of Attalea
spp. palms are infestation by bruchid beetles, predation by rodents and
predation by white-lipped peccaries. At Maraca Island Ecological Reserve,
in northern Brazil, A. maripa trees produce fruits containing 1 (67 % of
endocarps, n 5 1455 endocarps from 20 trees and 12 dispersal sites), 2
(29 %) or 3 seeds (4 %). Individual trees may produce fruits with different
numbers of seeds in one fruiting season, but those that produce multiple
seeded fruits do so year after year. Single-seeded endocarps are significantly
thicker (. 4 mm) than multiple-seeded endocarps (, 4 mm). Seeds within
single-seeded endocarps are more likely to be killed by bruchid beetles
than seeds from multiple-seeded endocarps (100 % vs. 71 % mortality per
seed). In 8 % of multiple seeded endocarps gnawed by rodents, one or
more seeds also survived. In captive feeding trials, however, seeds in thick
(single-seeded) endocarps have a greater probability of surviving predation
by white-lipped peccaries than those in the thinner, multiple-seeded endocarps, indicating that predation by whites-lipped peccaries may select for
single-seededness, while infestation by bruchids may select for multipleseededness. At the above-species level, similar patterns of selection may
also have influenced trade-offs between seed size and seed number usually
ascribed to reproductive syndromes (e.g., small-seeded colonist vs. largeseeded shade tolerant species), selection by seed dispersers or conditions
for seedling establishment.

and in all possible combinations of three. After the stocked populations

reached carrying capacity, the mesocosms were invaded; each of the four
species invaded monocultures of the other three species as well as the
polyculture that did not originally contain it. I also used the number and
biomass of "volunteer" (naturally recruiting) invaders to determine the effect of consumer diversity on invasion success within lower trophic levels.
Increasing diversity of resident grazers reduced population growth rates
and biomass of invading grazers. This result parallels those found in plant
communities: at the local scale, invasion success is reduced by within trophic-level diversity. Resident grazer species composition, however, also
influenced invasion dynamics at all trophic levels examined. The resulting
differences in community composition and habitat structure subsequently
affected the success of mobile invaders. These results suggest that trophic
process and habitat engineering can strongly influence invasion success in
multi-trophic level systems.

FRANCE, KRISTIN E.* and J. EMMETT DUFFY. School of Marine Science & Virginia Institute of Marine Science, The College of William and
Mary, Gloucester Point, Virginia. Testing effects of consumer diversity
on invasion success in a multi-trophic level system.
Local scale experiments tend to support Eltons hypothesis that diversity
decreases invasion success. In contrast, regional studies indicate that diverse communities are more likely to be invaded. Most previous studies
have been conducted using plants or otherwise sessile organisms at basal
trophic levels. Effects of consumer diversity on ecosystem structure and
function, however, are likely to be qualitatively different from those of plant
diversity. Consumers can also affect habitat structure and resource availability for organisms at other trophic levels, which might mediate invasion
dynamics at those levels. Using estuarine seagrass beds as a model system,
I examined the relative importance of mobile consumer species richness
and composition in determining invasion success of other native consumers
in a flow-through mesocosm experiment. I varied species richness from 03 using four native grazer species (amphipods and isopods) placed singly

112

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FRANKLIN, JANET.* San Diego State University, San Diego, CA. Natural versus human disturbance and the structure of tropical rain forest
in the western Pacific islands.
On December 31, 2001, Tropical Cyclone Waka, a category 3 hurricane,
passed directly over the Kingdom of Tonga. During a previous study of
secondary succession, 44 forest plots (600-m2) on 13 islands had been
surveyed in 1995. Twenty-one of the plots were resurveyed in June 2002
in one of the only post-cyclone assessments of established lowland tropical
forest plots in the western Pacific. Tree mortality averaged 6%, varied from
0-7% for lowland late successional species, and tended to be higher for
early successional plots (8-16%) and species (up to 19%). Severe damage
(uprooting, snapped stems) affected 25% of the 2030 stems measured. The
greatest mortality and severe damage combined (.39% of stems) occurred
in plots that were early successional. The late-successional forest recovered
by direct regeneration showing little change in composition or structure.
While early-successional forest showed little species turnover, the cyclone
had a great impact on stand structure, setting back the successional clock.
There is, nonetheless, a much greater change to forest community structure
during secondary succession following agricultural abandonment than following this severe natural disturbance.
FRANKLIN, SCOTT B.,1,* WEI WANG1 and JOHN OUELLETTE.2 1 Department of Biology, University of Memphis, Memphis, TN, USA; 2 Research & Conservation Department, Memphis Zoo, Memphis, TN, USA.
Giant panda herbivory effects on bamboo dynamics.
Understanding small-scale patterns of disturbances will render a greater
understanding of local controls and mechanisms affecting bamboo regeneration dynamics. We studied herbivory and bamboo dynamics (growth and
regeneration) in the Foping National Nature Reserve (33845 N 107850 E)
on the southern face of the Qinling Mountains, Shaanxi Province, PR China. We set up nine paired sites in the summer forage habitat of giant panda,
where Fargesia qinlingensis is the dominant bamboo taxa. Each site contained a set of paired plots: one naturally foraged by giant panda, an ad-

jacent clipped (mimicking foraging) plot, and a control. Natural foraging

plot areas ranged from 0.8 m2 to 11.6 m2. Densities for both foraged and
adjacent control plots ranged from 12 to 97 stems m-2; culm heights ranged
from 1.9 m to 2.8 m. Total stem density was positively correlated with
dead culm density (r2 5 0.82) and new shoot density (r2 5 0.85). Average
culm diameter of foraged stems was 8.88 mm, and culms were clipped at
an average height of 0.68 m. We explored plot characteristics that might
affect foraging behavior. Foraging was biased toward culm sizes between
8 and 11 mm diameter, regardless of the size class distribution of the naturally foraged plot. However, there was no difference between the size class
distribution of the foraged plots and the adjacent control plots (x2 5 2.57).
Further, we found no significant differences in total culm density, dead
culm density, new shoot density, or broken culm density between foraged
and adjacent control plots. However, a pattern of greater foraging in plots
with a greater number of broken stems was observed and needs further
research.
FRANKS, STEVEN J.,* ANDREA M. KRAL and PAUL D. PRATT. Invasive Plant Research Lab, USDA/ARS, 3205 College Avenue, Ft. Lauderdale, FL, USA. The effects of insect herbivory on seedlings of the
invasive tree Melaleuca quinquenervia.
We investigated the influence of herbivory by two introduced biocontrol
insects on survival and performance of Melaleuca quinquenervia (Myrtaceae) seedlings, an invasive tree native to eastern Australia. In November
2002, Boreioglycaspis melaleucae (Psyllidae) nymphs and Oxyops vitiosa
(Curculionidae) larvae were transferred onto Melaleuca seedlings within
replicated 1/4 m2 caged plots in Palm Beach County, Florida. The treatments included three densities of 1st and 2nd Boreioglycaspis instars at 1,
15, and 50 nymphs per seedling, one treatment of a single Oxyops larva
per seedling, another treatment of both one Oxyops larva and one Boreioglycaspis per seedling, a caged plot without insects, and an uncaged
control. Yellow sticky cards were placed within each cage to trap out adult
Boreioglycaspis as they emerged, thereby maintaining uniform herbivore
densities among replicates. Leaf number and plant height were evaluated
monthly for four months. Herbivory by Oxyops did not affect seedling
height, leaf number, or survival. Boreioglycaspis herbivory significantly
decreased leaf number, and orthogonal contrasts showed significant differences among Boreioglycaspis density treatments. After completion of one
generation, Boreioglycaspis herbivory at the highest density caused mortality in 16 of 25 Melaleuca seedlings. The results indicate that Boreioglycaspis herbivory may be effective in reducing growth and survival of
Melaleuca seedlings, with the impacts on the plants related directly to
insect density. There was no evidence that Oxyops would be effective in
controlling Melaleuca seedlings, or that there were any interactions among
the biocontrol insects.
FRATERRIGO, JENNIFER M.,1,* MONICA G. TURNER,1 SCOTT M.
PEARSON2 and PHILIP DIXON.3 1 University of Wisconsin-Madison,
Madison, WI; 2 Mars Hill College, Mars Hill, NC; 3 Iowa State University,
Ames, IA. Impacts of past land use on spatial heterogeneity of soil
nutrients in Southern Appalachian forests.
Land-use practices can have a lasting effect on the distribution of soil
nutrients, which may in turn influence the recovery of native ecosystems
following land abandonment. The spatial structure of variation in soil resources may be particularly important for the establishment and persistence
of plants recolonizing disturbed areas, yet the longterm impacts of land
use on the spatial patterns of soil nutrients are poorly understood. We
studied forest soils along a gradient of disturbance history in the southern
Appalachian highlands to answer two questions: (1) How does prior land
use influence the amount and scale of variation in soil nutrients? and (2)
Do historic land-use practices alter the structure of fine-scale patterns in
soil resources? Mineral soil was sampled during 2001 and 2002 in 40 plots
from 13 forest stands that differed in land-use history (previously pastured,
logged, or undisturbed) in western North Carolina. Soil nutrients were most
variable in undisturbed stands and this variability was greatest at fine
scales. In contrast, formerly pastured and logged stands exhibited less variability overall, and soil nutrients were relatively more variable at coarse
scales. Geostatistical analysis of fine-scale patterns further revealed that
spatial structure in soil cations was more closely linked to former land use

than in other soil nutrients. These results suggest that land use has persistent
effects on the spatial distribution of soil resources and may also influence
scales and patterns of variability in the biota.
FREEMAN, MARY C.* U.S. Geological Survey, Patuxent Wildlife Research Center, Athens, GA, US. Effects of drought and policy on fish
assemblages in the southeastern United States.
The southeastern United States is a globally significant center of aquatic
biodiversity, the great majority of which evolved in flowing-water ecosystems. Over the past century, hydropower development has fragmented and
extensively inundated this stream habitat, and native faunal assemblages
have persisted primarily in the undammed tributaries of major river systems. These tributaries are coming under increasing pressure from water
supply development. Present policy focuses almost entirely on provision
of minimum flow levels at withdrawal points and downstream from reservoirs. Research conducted on the effects of municipal withdrawals in six
river systems across the Piedmont province of Georgia during the drought
of 2000 and 2001 has shown declining integrity of fish assemblages coincident with increasing relative withdrawal size. In contrast, reference
streams (having relatively unaltered flow regimes) have sustained high Index of Biotic Integrity scores despite drought flows. Additionally, fish assemblages downstream from water supply reservoirs exhibit declines in a
suite of native fishes relative to reference-stream assemblages. Fishes persisting and dominating assemblages downstream from reservoirs are species common in lentic and lotic habitats, including sunfishes (Centrarchidae) and mosquitofish (Poeciliidae). Assemblages downstream from water
withdrawals (i.e., without instream impoundments) are more similar to reference conditions, generally supporting larger diversity and abundances of
minnow (Cyprinidae), darter (Percidae), and sucker (Catostomidae) species, as well as in selected catfish (Ictaluridae) and lotic bass (Centrarchidae) species. The differences in fish assemblages downstream from water
supply reservoirs compared to other sites are not strongly related to differences in local habitat, catchment landuse, or minimum flow requirements. Understanding the biological consequences of alternative water supply strategies will require quantifying the cumulative effects of reservoir
proliferation and flow alteration at a landscape scale, issues that extend
beyond consideration of provision of minimum flows during drought or
low-flow periods.
FREIDENBURG, KEALOHA1,* and DAVID SKELLY.2 1 University of
Connecticut, Storrs, CT; 2 Yale University, New Haven, CT. Microgeographic variation in thermal preference by an amphibian.
Ectotherms use behavior to buffer effects of temperature on growth, development and survival. While behavioral thermoregulation is widely reported, localized adaptation of thermal preference is poorly documented.
Larval amphibians live in wetlands ranging from entirely open to heavily
shaded by vegetation. We hypothesized that populations undergo localized
selection leading to countergradient patterns of thermal preference behavior. Specifically, we predicted that wood frog (Rana sylvatica) larvae from
closed canopy ponds would be more strongly temperature selective and
would prefer higher temperatures than conspecifics from populations found
in open canopy ponds. In a study of 6 breeding ponds in northeastern
Connecticut, USA, these predictions were upheld. The countergradient, microgeographic variation in thermal preference documented here implies that
wood frog populations may have diverged rapidly in the face of contrasting
selection pressures. Rapid, behaviorally mediated responses to changing
thermal environment have important implications for understanding population responses to climate change.
FREY, SERITA D.* and MELISSA A. KNORR. University of New Hampshire, Durham, NH. Nitrogen Enrichment Effects on the Litter Decomposer Community.
Inputs to the global N cycle have increased significantly due to human
activities and there are many uncertainties as to how this N enrichment is
impacting terrestrial biotic communities and ecosystem function. Our objectives were to (1) synthesize the existing literature on the effects of N
additions on litter decomposition and (2) examine how N additions influence the structure and function of the decomposer community. To synthe-

Abstracts

113

size existing data, we conducted a meta-analysis of previously published

empirical studies that have examined the effects of N enrichment on litter
decomposition. When all studies were analyzed together, we found that N
additions did not have a significant stimulatory or inhibitory effect on litter
decomposition rates. However, interesting trends emerged when study sites
were separated by climate and N availability. Decomposition at sites exhibiting low N availability, as a result of climate and/or low N deposition
rates, was stimulated by moderate levels of N fertilization. In contrast,
moderate fertilization inhibited decomposition at sites where N availability
was already high. To address our second objective, we examined the effects
of N enrichment on the relative abundance, diversity, and activity of bacteria and fungi in forest soils exposed to chronic N fertilization. We collected and analyzed soil samples from the Chronic N Addition Plots at
Harvard Forest. The samples were analyzed for active bacterial and fungal
biomass, microbial functional diversity, microbial-derived organic matter,
and the activities of celluloytic and ligninolytic enzymes. Total microbial
biomass was 35% lower in the high N compared to control plots. The
proportion of the microbial community comprised of fungal biomass declined from 70-94% in the control plots to 45-53% in the high N treatments.
The metabolic diversity of the microbial community and lignin-degrading
enzyme activity were also reduced under long-term N additions. This talk
will synthesize the results from both studies.
FRIDLEY, JASON D.,* JOEL M. GRAMLING, R T. JOBE, JESSICA A.
KAPLAN, MEGHAN MCKNIGHT, AMANDA SENFT, DAVID B. VANDERMAST and ROBERT K. PEET. University of North Carolina at Chapel
Hill, Chapel Hill, NC. Fine-scale species-area relationships of the vascular flora of the Southeast.
Species-area relationships at fine scales (0.1 ha and below) have been neglected by ecologists, both for lack of appropriate data and the perception
that they are unrepresentative of larger-scale diversity patterns. We reassessed the meaning and importance of fine-scale species-area relationships
using the extensive database of the Carolina Vegetation Survey (NC and
SC, USA), including over 1300 plots with values of floristic richness at
each of 6 areas regularly spaced on a log-10 scale, from 0.01 to 1000 m2.
Contrary to the expectation that the standard Arrhenius function does not
fit fine-scale data, our data closely fit a straight line in log(species) vs.
log(area) space (as opposed to semi-log space), and log-log slope (z) values
fell within a narrow range (0.2-0.4) despite a 30-fold range in 0.1 ha richness. Variation in z between plots was largely a function of richness at the
smallest scale (0.01 m2), which was itself influenced by vegetation type,
edaphic gradients, and, most importantly, mean size of individuals. When
the slope of each area-richness transition was examined separately, the 0.1
to 1 m2 transition consistently had the highest rate of species accumulation,
with each subsequent transition accumulating slightly fewer species in loglog space. When estimates of state, continental, and global levels of floristic
richness were added to our results, a Preston-type triphasic curve emerged.
We suggest that 1) fine scale species-area relationships are remarkably consistent and are indicative of small-scale individual density patterns; 2) triphasic species-area curves should be expected from fine scales to the globe;
and 3) the first inflection point of the triphasic curve, where the density
constraint disappears, is commonly above the 0.1 ha scale in floristic data.
FRIEDENBERG, NICHOLAS A.* University of Florida, Gainesville, FL,
USA. The coincident evolution of dispersal and specialization: Consequences for diversity.
Both dispersal rates and local adaptation may evolve in response to the
spatial and spatiotemporal heterogeneity of a species potential habitats.
Increasing spatial heterogeneity should select for lower dispersal rates and
increased local adaptation. Increasing spatiotemporal fluctuations in the environment should select for a higher dispersal rate and favor a broad range
of tolerance over strong local adaptation. Both characters are proposed
mechanisms for the determination of species diversity. I allowed dispersal
propensity and the degree of local adaptation to evolve independently in
an individual based model. Individuals occupy a linear landscape of patches
that differ by a fixed degree, representing a transect along an environmental
gradient. Individuals belong to ancestral lineages and to derived lineages
that arise randomly with a fixed probability, much like a simplistic model
of speciation. I find that increased dispersal rates accelerate the loss of

114

Abstracts

lineages, ancestral and derived, whereas the relationship between local adaptation and diversity is contingent upon spatial heterogeneity. Above a
threshold level of spatial heterogeneity, an increase in the severity of patch
fluctuations leads to a negatively correlated change in dispersal and local
adaptation and a loss of ancestral and derived lineages. I will discuss these
results in terms of diversity both within and between species.
FRIGGENS, MICHAEL T.,1,2,* ROBERT R. PARMENTER,1,2 TERRY L.
YATES1,2 and JAMES R. GOSZ.1,2 1 Department of Biology, University of
New Mexico, Albuquerque, New Mexico; 2 Sevilleta Long Term Ecological
Research Project, University of New Mexico, Albuquerque, New Mexico.
Can simple correlation analyses detect "bottom-up" trophic cascades
among rodents on the Sevilleta NWR?
In habitats with relatively low primary productivity (i.e., deserts), there is
evidence that predators are largely donor controlled, leading predominantly
to a two-link, primary production-herbivore food chain wherein herbivore
abundance is mediated by primary production of vegetation. This "bottomup" trophic cascade hypothesis has recently been enlisted to explain Peromyscus rodent density increase and subsequent Hantavirus outbreak in the
Southwestern US. There is some debate as to the utility of simple linear
correlation in explaining trophic relationships or in detecting trophic cascade. In order to test for the presence of bottom-up, resource driven dynamics among 3 families and 20 rodent species occurring across varying
ecosystems with similar climatic regimes, we employed simple Pearson
correlation statistics, comparing rodent species density with seasonal precipitation and vegetation production (as measured from AVHRR NDVI) at
varying lags in five habitats from 1989-2000, as part of the Sevilleta Long
Term Ecological Research project. We did not detect overall patterns based
on taxonomic affinity or site. But although results were highly variable
among species and between habitats, 15 of 20 species exhibited positive
correlation to seasonal PPT and subsequent vegetation production in at least
one habitat. Our results suggest that while there are certainly more complex
explanations for density dynamics observed at our sites, simple correlation
statistics succeeded in highlighting positive relationships with available resources and the presence of bottom-up trophic cascades in many instances
on the Sevilleta National Wildlife Refuge.
FROST, CECIL.* North Carolina Plant Conservation Program, Raleigh,
North Carolina. Landscape fire ecology: Using fire compartment size to
predict presettlement fire frequency.
One of the emergent properties of the new field of Landscape Fire Ecology
is the apparent control of fire frequency in some regions by the size of
their fire compartments. Other things being equal in a given landscape, the
larger the fire compartment the higher the fire frequency. In the presettlement landscape, fire frequency may have been driven in some regions by
Native Americans and in others by lightning ignitions. Development of
mechanisms of fire dependence might be expected to take evolutionary
time. Immigration of man into the Western Hemisphere occurred only 1220,000 years ago. Since that length of time would provide too few generations for us to expect evolution of complex adaptations to fire, we might
expect that truly fire dependent species would be concentrated in those
areas where lightning dominated the fire regime. The national pattern of
lightning strikes illustrates hot spots for lightning activity in such places
as central Arizona and the Southeastern Coastal Plain. In the Southeast
there appear to be hundreds of fire dependent species. In 20 years of monitoring one of these, Venus flytrap has been shown to require a mean fire
frequency of 1-3 years for survival, making it the most frequent-fire dependent species yet documented. Further, the distribution of Venus flytrap
provides substantiation of the relationship of fire compartment size to presettlement fire frequency. In the West, the relationship of fire compartment
size to fire frequency may have been obscured by the very large size of
fire compartments, but a rare wetland plant, Huachuca water-umbel (Lilaeopsis schaffneriana, ssp. recurva) appears to be a frequent fire analog
of Venus flytrap. The pattern of such fire-frequency indicator species in
landscape fire compartments tells us enough about the original fire frequency that they can be used as one line of evidence for mapping presettlement fire regimes.

FROST, CHRISTOPHER J.* and MARK D. HUNTER. University of

Georgia, Athens, GA, USA. Canopy herbivory on red oaks increases
soil respiration and soil carbon independent of herbivore-derived inputs.
Increased nitrogen (N) mobilization and export from terrestrial forest ecosystems following canopy herbivory has been well documented, though the
mechanism behind the loss is not clear. Herbivore-mediated changes in N
distribution may affect above- and belowground trophic systems and carbon
(C) distribution. We initiated a replicated mini-ecosystem experiment to
test the hypothesis that insect droppings (frass) influence soil C and N
concentrations and turnover following insect defoliation. One hundred and
sixty nursery-grown Red Oaks (Quercus rubra) were transplanted to 7gallon pots with soil and litter from watershed 27 at the Coweeta Hydrologic Laboratory in North Carolina and overwintered in experimental pot
stands. Approximately 6 weeks following budburst, trees were subjected to
insect damage, mechanical damage, or left as undamaged controls. Frass
from the herbivore-damaged trees was collected and redistributed to half
of each of the damage groups, creating a 3x2 factorial experimental design.
Herbivory increased soil respiration independent of frass additions. We
hypothesize that the increased soil respiration results from enhanced root
exudate C, which is used by rhizosphere microbes as an energy source. To
our knowledge, a link between damage and soil microbes has not been
demonstrated in a hardwood system, and experimentally separating herbivory from other factors when measuring soil respiration has not been attempted. A root exudation hypothesis is developed, future experiments are
outlined, and links between canopy herbivory and ecosystem function are
discussed.
FROST, JODY L.* and MARY C. CURRAN. Savannah State University,
Savannah, Georgia. Diel behavior of blackcheek tonguefish Symphurus
plagiusa.
Blackcheek tonguefish Symphurus plagiusa, are abundant in shallow coastal waters, but have rarely been the subject of scientific study. In order to
assess the behavioral patterns of one of the least understood flatfishes,
observations of their burial and activity patterns were performed over a
40-h period in a controlled environment. Individuals were caught, measured, and randomly assigned to seperate experimental chambers (83 x 43
x 34 cm) within 6 hours of capture. Chambers contained 1 cm of clean,
sorted sand and 10 cm of water (23-298C, 24-25 ppt). To determine if any
diel or tidal rhythmicity existed, fish were observed in approximately 3-h
intervals in correspondence with high, ebb, low, and flood tides. Diel activity was quantified, in part, by determining the percent of the tonguefish
body that was covered by sediment. Movement was assessed based on
whether fish had changed position or orientation between readings. Results
indicated that blackcheek tonguefish had significantly more sediment coverage during the day than at night. The fish changed position significantly
less during the day than at night. Tidal stage had little effect on tonguefish
movement. Future studies will focus on the interactions of light level, orientation, and lunar phase on activity patterns.
FUHRMAN, NICHOLAS E.,* CAROLYN A. COPENHEAVER, PAUL
A. GELLERSTEDT, NICHOLAS A. POVAK and LAURA S. STEPHENS.
Virginia Polytechnic Institute and State University, Blacksburg, Virginia,
USA. Forest encroachment into glades on Buffalo Mountain, Virginia.
Buffalo Mountain Natural Area Preserve is a 400 ha preserve managed by
the Virginia Natural Heritage Program. Buffalo Mountain is the highest
mountain (1,190 m asl) in Floyd County, Virginia and home to 14 rare
plant species and six different vegetation communities. The Natural Heritage stewards have expressed concerns that the grass-dominated glade communities on Buffalo Mountain are succeeding to forest. The objective of
this study was to evaluate the risk and rate of forest encroachment within
these glade openings. Three meter wide belt transects were established from
the center of 6 glades passing through the glade-forest interface and into
the interior of the forest. Within these belt transects, increment cores and
disks were taken from all trees. These cores and disks were then aged to
evaluate whether a relationship between distance from edge of glade and
age of tree existed. In addition to the belt transects, a master chronology
from twenty of the oldest white oak (Quercus alba L.) trees was created

from cores sampled both within and outside of the belt transects. This
allowed us to evaluate whether recent growth patterns in the dominant tree
species on Buffalo Mountain have changed within the last 130 years.
FUJIWARA, KAZUE.* Yokohama National University, Yokohama, Kanagawa, Japan. Warm-temperate deciduous forests in East Asia and the
southeastern US: Response to global warming?
Warm-temperate deciduous forests (sensu Kira) in East Asia occur between
the warm-temperate evergreen and temperate deciduous broad-leaved forest
regions. Species of evergreen broad-leaved forests are climatically excluded
from these transitional forests. Over the past 50 years evergreen broadleaved species have invaded the warm-temperate deciduous forests of Japan, including drier forests, some of which have had their canopy layers
occupied by evergreen Quercus myrsinaefolia and understoreys by Aucuba
japonica, Hedera rhombea, Trachycarpus fortunei, etc. which escaped
from gardens or hedges. Warm-temperate deciduous forests (in this sense)
also occur in China and Korea, but warm-temperate evergreen species apparently have not invaded them yet. In Japan these warm-temperate deciduous forests occur as secondary (successional?) forests in the evergreen
forest region, and as such they may correspond to coastal-plain deciduous
forests in the southeastern USA. Deciduous forests on the southeastern
coastal plain include dry Quercus forests, semi-evergreen floodplain and
wetland forests, and Fagus forests as far south as Florida. Some warmtemperate evergreen species, such as Quercus virginiana, Q. hemisphaerica, Sabal palmetto, Magnolia grandiflora, Ilex vomitoria, and Tillandsia
usneoides, as well as some exotics, appear to occur in wet forests more
than in dry forests. In China and Korea there appears to be a similar tendency for evergreen broad-leaved species to be precluded from drier forest
climates.
FULTON, MARK R.1,* and PAUL A. HARCOMBE.2 1 Bemidji State University, Bemidji, Minnesota, USA; 2 Rice University, Houston, Texas, USA.
Concisely characterizing forest dynamics with change vectors.
Changes in a forest stand or a set of monitoring plots can be envisioned
as vectors in a multidimensional space defined by the abundance of tree
species. These vectors can be used to concisely characterize aspects of
forest chance that are more difficult to recognize in the trends of individual
species. A vector of change is characterized by a direction and a magnitude.
The direction of a change vector is determined by relative amount of
change in each species; the magnitude is the total amount of change. One
can compare directions of change among several monitoring plots by comparing angles between change vectors, either pairwise between individual
plots or between individual plots and the overall trend. If one plot is monitored over an extended period of time, one can compare directions or
magnitudes of change from one survey interval to the next to reveal the
effects of changing conditions. Randomization tests can be used to detect
consistent temporal trends in response to past disturbance or changing climate, thereby distinguishing trends from fluctuations of an essentially stable forest. Plotting changes in ordination diagrams has been used in the
past to approach these kinds of questions, but comparing change vectors
directly with each other avoids the distortions that are inevitable when a
high-dimensional space is projected onto a low-dimensional space. We
demonstrate this approach using long-term data from two stands in eastern
Texas: a mesic forest disturbed by a moderate hurricane, and a bottomland
hardwood forest with a variable flooding regime.
FUNK, JENNIFER L.,1,* CLIVE G. JONES,2 MANUEL T. LERDAU,1
DENNIS W. GRAY,1 HEATHER L. THROOP3 and LAURA A. HYATT.4
1
State University of New York, Stony Brook, NY; 2 Institute of Ecosystem
Studies, Millbrook, NY; 3 Beloit College, Beloit, WI; 4 Rider University,
Lawrenceville, NJ. Variation in isorpene emission from Quercus rubra:
sources, causes and consequences for estimating fluxes.
Because studies examining the patterns and regulation of isoprene emission
from plants usually monitor only a few individuals and/or causal factors,
it is not known at what scale plants show the greatest variability in basal
emission rate (BER) of isoprene. We measured BER from a total of 40 red
oak (Quercus rubra) individuals from 2 field populations from 1997 to
1999. We found inter-plant variation to explain 60% of the total variance

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115

with only 40% being explained by leaf and branch combined. Absolute
flux rates from the 2 populations, which differed in plant age, were similar.
To examine temporal variation in BER, 10 individuals were measured repeatedly during the 1998 and 1999 growing seasons. During 1998, interplant variation explained most of the variance. Drought conditions during
July of 1999 caused a substantial decline in BER for all plants. Thus,
seasonal effects explained most of the variance in the 1999 dataset. Absolute flux rates were similar between years, however individual plants did
not maintain consistent average flux rates between years. The causal factors
for the large inter-plant variation observed in this study are uncertain but
may include differences in plant history, microclimate or genotype. Seasonal patterns of BER were best explained by temperature and PAR integrated over the previous 6 hours (r2 5 0.40). Overall, we found BER to
be highly variable between plants and within the growing season. Consequences of this leaf-level variability for modeling fluxes across larger spatial and temporal scales will be discussed.
FUREDI, MARY ANN* and JAMES B. MCGRAW. West Virginia University, Morgantown, WV. Deer browsing effects on American ginseng:
Better late than never?
American ginseng (Panax quinquefolius L.) is a long-lived perennial herb
found in the deciduous forests of the northeastern United States and southern Canada. Although American ginseng is widely distributed, it is considered rare within much of its range. This rarity may be due to harvesting
for the medicinal market and browsing by white-tailed deer. A simple conceptual model was developed that predicted less pronounced deer browsing
effects on population growth as the season progressed. The objective of
this study was to test this model in a large natural population of American
ginseng. Size, reproduction, survival, and browsing status were measured
during the summers of 2001 and 2002. The data were used to parameterize
a population matrix model. The effect of deer browsing was quantified by
calculating the sample influence function for each plant. In 2001, 24.1%
of all plants were browsed. The effect of timing of browse on plant height,
reproduction, and influence on population growth was slight. However,
growth and reproduction of browsed plants was significantly reduced relative to unbrowsed plants. The net effect of browsing on population growth
was negative when compared with unbrowsed plants of comparable size.
This conclusion, however, may depend on the level of harvest pressure in
the population since browsed plants are invisible to harvesters. Efforts to
enhance wild ginseng populations through management may be hampered
by monospecific management for white-tailed deer that does not account
for collateral effects of that species.
FUSSMANN, GREGOR F.* Universitat Potsdam, Potsdam, Brandenburg,
Germany. Sensitivity to model structure and the paradox of enrichment.
Rosenzweig (1971) showed mathematically that enrichment of a predatorprey system can lead to dynamical destabilization. The implications for
natural communities have been debated up to the present day. I show that
the choice of function, which describes the uptake of prey by the predator
(functional response), critically determines whether and to what degree the
paradox of enrichment occurs. I use graphical isocline analyses to demonstrate that two popular functions (Holling II, Ivlev), which may provide
virtually identical mathematical descriptions of the functional response, can
cause drastically different dynamical behavior. Isocline analysis also reveals
that subtle differences between the two functions become exceedingly important when predator and prey populations interact. Sensitivity to model
structure may be the reason why destabilization does not occur in many
natural and experimental communities although simple models predict it.
GABBARD, BETHANY L.* and NORMA L. FOWLER. University of
Texas-Austin, Austin, TX, USA. The joint effects of competition and
simulated grazing on an exotic grass and a co-occurring native species.
Bothriochloa ischaemum (King Ranch bluestem grass) is a Eurasian species
that has become common throughout central Texas. To investigate its effects upon the related native dominant, Schizachyrium scoparium (little
bluestem grass), the separate and joint effects of competition and simulated
grazing upon both species were measured in a fully factorial garden ex-

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Abstracts

periment. Plants were grown in pots for 5 months. Half of the 360 pots
had B. ischaemum as the target species; the other half had S. scoparium as
the target species. Each pot received one of three competition treatments
(target plant grown alone; target plant grown with a conspecific plant; target
plant grown with a plant of the other species). Each of the pots also received one of three clipping treatments (unclipped; clipped to 5 cm once;
clipped to 5 cm twice). The final above-ground dry biomass of each species
was greatly reduced by the presence of a B. ischaemum plant in the pot:
B. ischaemum has a very strong competitive effect. In contrast, there was
no significant effect of the presence in the pot of a S. scoparium plant on
either species: S. scoparium appears to have a relatively weak competitive
effect. The competitive response of S. scoparium was greater than the competitive response of B. ischaemum. Thus B. ischaemum outcompeted S.
scoparium. Unfortunately, central Texas grasslands have few other species
that are more likely than S. scoparium to outcompete B. ischaemum. Clipping had a significantly negative effect on both grass species; as expected,
the negative effect of clipping was greater on S. scoparium, which is taller.
We expected that, in the absence of clipping, the taller S. scoparium would
outcompete the shorter B. ischaemum, but there was no interaction between
the effects of clipping and competition: cessation of grazing is not likely
to halt B. ischaemum.
GAGNON, PAUL R.* and WILLIAM J. PLATT. Louisiana State University, Department of Biological Sciences, Baton Rouge, LA, USA. Clonal
demography and multiple disturbances in Louisiana cane stands.
Species-rich and monodominant plant communities sometimes occur side
by side with no obvious environmental differences at their boundary. These
community differences may be the result of variation in the effects of
natural disturbances. We are exploring how natural windthrow affects population and genet dynamics of cane (Fam. Poaceae, Arundinaria gigantea,
W. Muhlenberg). Cane occurred historically in extensive monospecific
stands called canebrakes alongside diverse bottomland forests throughout
the southeastern U.S. Our fieldsite in northeastern Louisiana is on the Buckhorn Wildlife Management Area within the Lower Mississippi Alluvial
Valley. Here a massive tornado blowdown provides an opportunity to examine cane response to large-scale disturbances. We are conducting life
table response experiments (LTREs) in the blowdown and adjacent forest
to quantify 1) effects of natural windthrow gaps on cane ramet demography,
2) ramet demography of flowering cane stands, and 3) ramet demography
in small, discrete stands compared to dense, continuous (canebrake-like)
stands. Initial results indicate that cane culms in blowdown plots are shorter
in stature and have smaller diameter than those in forest-grown plots. Cane
culms in blowdown plots are younger, more vigorous and less-often damaged than those growing under forest canopy. Cane genets in blowdown
plots therefore have higher average l (lambda) than those in forest-grown
plots. These results may indicate mechanisms underlying potential environment-mediated changes between species-rich and species-poor communities, and will be useful for canebrake and bottomland hardwood forest
restoration efforts.
GAINES, KAREN H.* University of New Mexico-Albuquerque, Albuquerque, NM, U.S.A. Does size really matter? Factors affecting larval
odonate diversity patterns in a desert sinkhole complex.
The Bitter Lake National Wildlife Refuge in southeastern New Mexico
contains an unusual ecological mosaic consisting of desert scrub and grassland surrounding dozens of saline water-filled sinkholes of varying sizes,
geomorphologies, and water chemistries. Nearly one hundred species of
adult dragonflies and damselflies (Order Odonata) have been collected on
the refuge, representing the highest diversity of odonates in the state of
New Mexico. In order to characterize the distribution and diversity patterns
exhibited by the breeding odonate population, thousands of exuviae (final
instar larval exoskeletons) were periodically collected at over thirty sinkholes in 2000, 2001, and 2002. Multivariate analyses of sinkhole dimensions and water quality data revealed that while larger sinkholes did tend
to support more diverse larval odonate communities, average winter salinity
levels explained nearly 70% of the variation observed in species diversity,
as species with broad larval salinity tolerances successfully bred in most
sinkholes visited by adults regardless of sinkhole size. In contrast to predictions of island biogeography theory, this study suggests that in certain

cases, individual species characteristics (e.g. larval stenotopy) may have a

far greater influence on community diversity patterns than does habitat
patch size.
GAISER, EVELYN E.,* PABLO RUIZ, ANNA WACHNICKA, ANGELIKIE A. ZAFIRIS and MICHAEL S. ROSS. Southeast Environmental
Research Center, Florida International University, Miami, FL. Paleoecological determination of effects of salt-water encroachment on community migration in coastal South Florida wetlands.
Coastal communities in South Florida are migrating landward due to the
combined effects of freshwater diversion and sea-level rise. We examined
the current distribution of plants, periphyton and mollusks among 112 sites
in a 4 km2 area of coastline in the southeast saline Everglades, and compared it to past distributions determined from soil cores taken along transects in the same wetland band. Soil profies show an upper mangrovederived peat layer that thickens toward the coast, overlying a marl layer
on top of the limestone bedrock. Because marl-producing periphyton communities are associated with freshwater gramminoid and mixed gramminoid-mangrove communities, this transition likely represents an expansion
of the peat-producing fringing mangrove forest into areas previously occupied by Everglades marl prairie. The mollusk record confirmed this hypothesis, as sites located between the drainage canal and the coast showed
significant upcore increases in the ratio of marine to freshwater taxa, while
sites to the interior of the canal show the opposite trend. Terrestrial taxa
have also increased in the interior sites, indicating encroachment of remaining shallow gramminoid marsh by a shrub-forest community. Maps
showing the distribution of ecotones (including, in order from coast to
interior, fringing mangrove, dwarf mangrove, brackish gramminoid, freshwater gramminoid, and interior shrub-forest) in surface (modern) and subsurface layers have been used to calculate spatially explicit rates of ecotonal
migration in this hydrologically complex band of wetland basins.
GALLERY, RACHEL E.* and JAMES W. DALLING. University of Illinois Urbana-Champaign, Urbana, IL. Pathogens and seed mortality in
neotropical soils.
Successful recruitment for many light demanding tree species is dependent
upon seed survival in the soil seed-bank. Seed-bank experiments conducted
to date implicate soil-borne fungi as predominant mortality agents of smallseeded tropical tree species. To investigate the sources of variation in seed
susceptibility to pathogenic fungi, we conducted reciprocal transplant experiments between populations of common light demanding tree species in
lowland Neotropical forests. Seeds were collected from individuals of five
species from La Selva Biological Station, Costa Rica, and at Barro Colorado Island (BCI), Panama and incubated in the soil for five months. We
have found that: (1) fungi significantly limit survival in the soil-seed bank
for these species; fungicide experiments at both sites increased seed survival by 28-50% (2)overall seed mortality is significantly higher at La Selva
than at BCI (F549.0, df51, p,.0001), and (3) higher seed mortality for
one species, Cecropia insignis (Cecropiaceae), buried beneath conspecific
versus heterospecific crowns (F528.53, df5 1, P ,.0001). We also found
evidence for differences in seed survival among individuals, meaning seed
source and burial location contributed significant variation to overall seed
mortality. These results indicate the potential for host-specificity of soilborne fungi in this system, which could influence local species composition
through host-specific recruitment limitation. These preliminary results suggest that sources of variation in fungal-mediated seed mortality are evident
across multiple scales.
GALLUZZI, LAURIE A.* and ROBIN W. KIMMERER.* SUNY College
of Environmental Science and Forestry, Syracuse, NY. The basketry
plant: Hierochloe odorata; Sustainable managment through indigenous
harvesting practices.
The reintroduction of Native American management practices may be necessary to maintain existing wild and restored sweetgrass (Hierochloe odorata) populations in the Northeast. Viable inland sweetgrass populations are
essential to preserve the art of traditional basketry, as practiced by the
Haudenosaunee people of New York State. Yet, the supply of this important
cultural resource is threatened by various forces, including successional

change. In this study, hypotheses regarding the effect of different indigenous harvesting regimes on sweetgrass populatiob growth were tested using
a controlled field experiment comparing three different harvesting methods.
The results of this experiment suggest that sweetgrass, a disturbance adapted species, may show and increase in population density as a result of
indigenous harvesting practices. These harvesting practices may help to
maintain ecological conditions favorable for reproduction, thus preserving
community integrity. Traditional harvesting regimes, as well as other indigenous management practices, are a part of a larger body of Traditional
Ecological Knowledge that can offer valuable insight to those involved in
the field of native plant restoration.
GAMON, JOHN A.,1,* FAIZ RAHMAN,2 DANIEL SIMS,1 KARL F. HUEMMRICH3 and WALTER C. OECHEL.4 1 Center for Environmental Analysis (CEA-CREST), & Department of Biological Sciences, California State
University, Los Angeles, Los Angeles, CA; 2 Department of Geography,
Ball State University, Muncie, IN; 3 Code 923.4, NASA/GSFC, Greenbelt,
MD; 4 Global Change Research Group, San Diego State University, San
Diego, CA. SpecNet: Integrating optical remote sensing with ecosystem
flux measurements at FLUXNET sites.
Ecosystem carbon flux is often measured by eddy covariance, providing
temporally rich (but spatially poor) measurements of carbon and water
vapor flux at selected terrestrial ecosystems. On the other hand, remote
sensing provides a synoptic view of the world, yet has difficulty providing
temporally continuous data at a scale that can be related to flux measurements. SpecNet (Spectral Network) is designed to better integrate optical
remote sensing with ecosystem flux data at flux tower (FLUXNET) sites
as a basis for cross-ecosystem analysis of carbon and water vapor flux. A
unique feature of SpecNet is the provision of repeated optical sampling at
a scale that closely matches that of the flux tower footprint. Here we report
initial results comparing optical and flux data from several SpecNet sites,
including Alaskan tundra, Californian chaparral, and deciduous forest. This
combined analysis of optical and flux data reveals contrasting and dynamic
ecosystem behavior that reflects underlying limitations to ecosystem productivity. For example, water limitations are apparent in both Arctic and
chaparral ecosystems. Statistical analyses suggest optimal sampling and
aggregation methods for optical and flux data that best reveal temporally
changing flux patterns. These results demonstrate the complementary nature of optical and flux sampling, and provide a basis for improving remotely sensed estimates of carbon and water vapor flux from terrestrial
ecosystems.
GANGER, MIKE T.* Department of Biology, Massachusetts College of
Liberal Arts, North Adams, MA, USA. Inferring genet development:
Interplay of meristem commitment and genet integrity.
Canada mayflower (Maianthemum canadense Desf.) is a rhizomatous, perennial herb common to the understory of mixed coniferous-deciduous forests in New England. Mayflower genets consist of multiple ramets. In a
given year, ramets may exist as either flowering (2-3 leaves and a terminal
inflorescence consisting of 4-35 perfect flowers) or vegetative (1 leaf)
shoots. Production of a flowering shoot requires commitment of a terminal
meristem. If this meristem is not committed then the ramet will exist as a
vegetative shoot in the following year. The recruitment of flowering shoots
into Mayflower populations has relevance to their ecology and evolution.
A two-year field study was undertaken to identify factors related to recruitment of flowering shoots. Ages of ramets varied significantly, with
those existing as flowering shoots older on average than those existing as
vegetative shoots. In both years, greater than 50% of ramets existing as
flowering shoots were more likely to exist as isolated ramets (not connected
to other ramets). Isolated ramets existing as vegetative shoots were more
likely than expected to flowering in the following year. Ramets existing as
vegetative shoots connected to a ramet both basipetal and acropetal were
less likely than expected to flower in the following year.
GARCIA-NOVO, FRANCISCO and JUAN BAUTISTA GALLEGO-FERNANDEZ. University of Seville, Sevilla, Sevilla, SPAIN. Plant functional
types in response to global climate change.
Plant functional types (PFTs) reveal the association patterns of plant organs
exhibited by those species sharing a common environment, and PFT de-

Abstracts

117

scription is performed against a limited set of environmental variables.

Vegetation response to climate change may be explored trough the forecast
of changes in environmental variables and the PFTs response to them.
Coastal dune fields are suitable areas to monitor global climate change
because sea level rise, littoral current transport and sand movement, amplify
climate fluctuations inducing broad changes to the plant environment. The
Gulf of Cadiz (SW Spain) offers a coastal sector over 100 Km long dominated by sand beaches associated to dune fields. The vegetation of the area
has been thoroughly studied and the PFTs have been described for high
beach and embryo dunes, active and stabilized dune fields. To monitor
climate change effects on vegetation, a test area has been chosen in the
Gulf of Cadiz at the Rio Piedras estuary. It presents a spit 12 Km long and
300 to 700 m wide, with active dune fields and diverse coastal vegetation.
The identification of PFTs and their associated environmental variables let
the interpretation of successional processes in terms of species sequences,
PFTs substitution and dominant physical processes (sand transport, soil salt
leaching, groundwater availability, soil surface instability and others).The
study of the dynamics of Rio Piedras spit, associated the environmental
physical processes to the littoral evolution of the area. The relationships of
spit building and sea level rise; river discharge and littoral transport have
been established, thus linking PFTs to climate change effects. The contemporary patterns of PFTs distribution over the spit will be discussed against
some climate change hypothesis.
GARDINER, NED,1,* ANDREW SUTHERLAND,2 JUDY MEYER,2 CATHY PRINGLE,2 GENE HELFMAN,2 FRED BENFIELD,3 PAUL BOLSTAD4 and DAVID WEAR.5 1 American Museum of Natural History, New
York, NY; 2 Institute of Ecology, University of Georgia, Athens, GA; 3 Virginia Polytechnic Institute, Blacksburg, VA; 4 University of Minnesota, St.
Paul, MN; 5 US Forest Service, Raleigh, NC. Linking stream ecosystems
and landscape trajectories in the southern Appalachians.
We established an innovative procedure to examine relationships between
catchment land use and stream ecosystem structure which guided predictions of future stream ecosystem states in western North Carolina. We chose
eight stream sampling sites with similar elevation and catchment sizes representing three distinct categories of land-use history: "forested" (n52),
"rural" (n53), and "suburban" (n53). Algal biomass (Chlorophyll-a mg m2
and ash-free dry mass mg m-2), median substrate particle size, sediment
core particle size distributions, and total suspended solids (mg l-1) measures
did not differ among these groups. By incorporating previously-collected
data from the study area, clear patterns were evident for fish assemblages
and other water quality parameters. Ordination of fish collections data suggested four categories of streams with distinct catchment land uses in both
1970 and 1993: "forested", "rural", "suburbanizing farmland", and "suburban/urban". The percentage of omnivorous and of widespread fishes increased at sites with "forested", "rural", "suburbanizing farmland", and
"suburban/urban" catchments, respectively. Invertivores, lithophilic nest
spawners, and fishes with restricted ranges decreased in percentage across
that set of categories. As expected, NO3, K, Ca, Mg, and mean summer/
fall temperature each increased in the stated order. Ecological integrity was
maximal in rural watersheds and minimal in urban ones. We used these
observations to extrapolate potential stream ecosystem states in 149 catchments of similar size throughout four large basins in western North Carolina. Based on our observations, we infer two trajectories of possible future
stream ecosystem responses to likely future land use changes. If building
and road densities increase, streams in forested watersheds are likely to
become warmer, higher in nutrients, and support more omnivorous and
widespread fishes; proportions of invertivores, lithophilic nest spawners,
and endemics would decrease. In suburbanizing farmland catchments that
become more like suburban/urban ones, the same predictions hold.
GARMESTANI, AHJOND S.,1,2,* CRAIG R. ALLEN1,2 and K. MICHAEL
BESSEY.2 1 South Carolina Cooperative Fish and Wildlife Research Unit,
Clemson University, Clemson, SC, USA; 2 Clemson University, Clemson,
SC, USA. Discontinuities in urban systems over time.
Complex systems, such as ecosystems and urban systems, emerge unpredictably without the influence of central control but as a result of adaptive
behavior by their component, interacting agents. Urban systems exhibit
spatial patchiness in their social and economic infrastructure. Thus, urban

118

Abstracts

systems, much like ecosystems, are subject to a hierarchy of structure and

processes that govern the function and growth of cities at a variety of
scales. Urban size distributions have been described by Zipf9s law or the
rank-size rule, which imply invariant growth processes across scales. Zipf9s
law predicts that city-size distributions will have a continuous distribution,
and conform to the restraints of a power law. We analyzed city size distributions, by decade, from the southwestern region of the United States
for the years 1890-1990. We determined if the distributions were discontinuous and changes in the pattern of discontinuities over time. Discontinuities were determined with computer simulations utilizing a null model,
cluster analysis and SMW analysis. The data were discontinuous as determined by all 3 methods, with agreement among all comparisons. In 1890,
the southwestern United States region consisted of 48 cities, but by 1990
this had grown to 161 cities. Likewise, the maximum city size increased
from 38,067, to 3,198,259 during the same time period. Despite these enormous changes, our analyses identified 5 or 6 aggregations and discontinuities in each of the decades analyzed. Our analyses suggest that the
spatial and temporal processes shaping urban size distributions at different
scales are conserved over time despite great change in city function and
size.
GARNEAU, DANIELLE E.1,* and ERIC S. POST.2 1 The Pennsylvania
State University, University Park, PA, USA; 2 The Pennsylvania State University, University Park, PA, USA. Spatio-temporal niche partitioning
among three sympatric predator species in a single-prey system.
Terrestrial investigations of multiple predators sharing one prey are rare in
natural systems. This study offers an explanation as to how sympatric black
bears, grizzly bears and gray wolves coexist on a seasonally limiting prey
item, specifically moose calves. Spatial and temporal separation within a
shared niche dimension has classically been offered as an explanation for
sympatric species coexistence. Niche partitioning, resulting from differential hunting behaviors among members of the predator guild, is often used
as a means of capitalizing on a common resource. Our findings suggest
that black bears and grizzly bears temporally separate hunting during
moose parturition, due to their spatial overlap in habitat. Similarly during
periods of increased inter-predatory competition, black bears and gray
wolves exhibit temporal overlap in hunting bouts; therefore, they must
spatially segregate within different hunting habitats. Consequently, findings
from this study indicate the need to consider each member of the predator
guild individually, complete with its own suite of behavioral, physiological
and foraging strategies.
GARRETT, KAREN A.* Kansas State University, Manhattan, KS, kgarrett@ksu.edu. Accounting for disease in communities: the fourth horseman jockeys for position in the LTER network.
Patterns of fire, precipitation, and grazing have been key factors in the
maintenance of tallgrass prairie in the Great Plains of the USA. Konza
Prairie Biological Station (KPBS), a Long Term Ecological Research Site
in Kansas, has environmental manipulations in place for studying the effects of precipitation changes at the plot level and burning and grazing at
the watershed level. I will present results from experiments that a number
of collaborators and I have performed to evaluate how these environmental
drivers affect plant disease in tallgrass prairie at KPBS. Barley yellow
dwarf virus is common in several of the major grass species of KPBS and
more abundant in areas without experimentally increased precipitation. Increased precipitation favors abundance of the rust fungus Uropyxis petalostemonis, however, which has a dramatic effect on the reproduction of
its host, the legume Dalea candida. Increased precipitation also favors
abundance of the rust fungus Puccinia dioicae infecting Solidago canadensis, but proximity of a primary host species was a more important factor
in some years. Annual burning greatly reduced infection of the composite
Erigeron strigosus by Puccinia dioicae, but had little if any effect on rates
of infection of the grass Andropogon gerardii by Puccinia andropogonis.
GARTNER, TRACY B.* and ZOE G. CARDON. University of Connecticut, Storrs, CT. Neighbors on the forest floor: consequences for mixes
of leaf litter on differing soil types.
Leaf litters do not segregate neatly in forests; composition of litter mixes
and the stratification of leaf types on the forest floor depend on community

structure and timing of leaf drop. Also, soil properties influence decay of
litter mixtures by providing nutrients, influencing moisture and temperature
regimes, and housing differing decomposer communities. We studied decomposition of mixed sugar maple (Acer saccharum) and red oak (Quercus
rubra) litters in four northwestern Connecticut field sites, on calcareous and
acidic soils, using compartmentalized litterbags. Each litterbag contained
sugar maple in equal masses in both layers, red oak in both layers, or sugar
maple in the bottom compartment and red oak in the top (mimicking natural
leaf stratification in the forest). Litterbags were deployed at two calcareous
and two acidic sites; subsets were harvested and analyzed for mass and
nitrogen changes every three months for one year. Overall patterns in decomposition across sites correlated well with soil temperature and moisture.
Sugar maple decayed more quickly than red oak, but the rate of decay for
both litter types changed depending on the identity of neighboring leaves.
Maple decay was significantly increased (2-3%) in the presence of oak,
while oak decay was either unaffected or slowed (1-2%) in the presence
of maple. Changes in decay within the mixtures varied with site; decay
rates in the mixture were most altered on acidic soil. Spatial heterogeneity
in distributions of leaf litters could have significant impacts on nutrient
cycling in forests, and the extent of the impact can vary with the soil
substrate.
GAVIN, DANIEL G.* and FENG SHENG HU. Department of Plant Biology, 265 Morrill Hall, University of Illinois, Urbana, IL. Climatic vs.
non-climatic control of western hemlock distribution in its coastal and
interior ranges.
The role of climate in determining species distributions at large spatial
scales is rarely tested rigorously. We used three bioclimatic models (climatic envelopes, sequential maximum a posteriori classification (SMAP),
and response surfaces) to assess climatic vs. non-climatic controls of the
range of western hemlock (Tsuga heterophylla), a late-successional species
that occurs in disjunct coastal and interior ranges in northwestern North
America. The three bioclimatic models differ in the level of generalization
of the species-climate relationship. Each model was calibrated with both
the observed coastal and interior ranges of hemlock and then applied to
the entire region to predict the species distribution. All models predicted
the coastal range more accurately than the interior range. The response
surface model was most accurate in its calibration range but failed to predict the interior range using the coastal calibration. In contrast, the climatic
envelope model was the least accurate in its calibration range but predicted
a southern portion of interior range using the coastal calibration. The
SMAP model was moderately accurate within its calibration range and
could most accurately predict the interior range using the coastal calibration, suggesting that it has an appropriate level of generalization. The most
suitable models predicted the potential interior range of hemlock to be ca.
40% larger than its observed range. The poorer fit of all models in the
interior range than in the coastal range, and the agreement in substantial
areas of overprediction in the interior, suggest that non-climatic factors
affect range limits in the interior to a greater degree than near the coast.
Dispersal limitation and/or competition following fire probably exert important constraints on the distributional limits of hemlock in the interior
range.
GEDDES, NEOMA A.* and SUSAN MOPPER. University of Louisiana
at Lafayette, Lafayette, LA, USA. Deer florivory influences arthropod
biodiversity in a southern brackish wetland.
Interactions between plants and herbivores are multifaceted. Herbivores can
drastically alter the appearance of landscapes, and their overabundance not
only affects primary production but can also have far-ranging effects on
other trophic levels. Although herbivory has been a traditional research
focus, florivory may alter community dynamics uniquely. Here, we report
on an interaction between white-tailed deer (Odocoileus virginianus), blue
flag iris (Iris hexagona), and arthropods associated with I. hexagona. Reproducing both sexually and clonally, I. hexagona is a native perennial
wild flower that occurs throughout the southeastern Gulf Coast. Our study
focused on the recently isolated I. hexagona populations that are found on
Marsh Island, Louisiana. After establishing remote infrared camera systems
at two Marsh Island sites, the organism responsible was determined conclusively to be white-tailed deer. First, we report on a field survey of the

occurrence of complete florivory (entire flowers and immature seed capsules are browsed and seed production eliminated) at a total of ten I. hexagona populations located on Marsh Island, LA. Second, we present arthropod data collected from within and without experimental deer exclosures that were established to protect flowers from browsing. We discuss
the implications of deer florivory on the more than forty-five arthropod
species collected, among them native ants (Crematogaster sp.) that are
unique colonists of iris seed capsules. This research was supported by National Science Foundation grant DEB-0124901.
GEIGER, ERIKA L.* and GUY R. MCPHERSON. University of Arizona,
Tucson, Arizona, USA. Changes in structure in semi-desert grasslands
following restoration of fire.
In an effort to recreate pristine semi-desert grasslands in the southwestern
United States, land managers have touted restoration of fire as the most
appropriate tool. Yet, fires often are implemented contrary to historic fire
regimes and into vastly altered systems. In southwestern grasslands, Eragrostis lehmanniana, a nonnative perennial grass from southern Africa, is
implicated as the cause for loss of native organisms. Currently, land managers use fire in controlling this nonnative species despite considerable
evidence demonstrating the benefits of fire to E. lehmanniana. We designed
a large-scale experiment in the semi-desert grasslands of southeastern Arizona to quantify changes in plant community structure following fire treatments (spring, summer, no fire) across a gradient of dominance by a nonnative grass. There was a slight change in the proportion of E. lehmanniana
following fires (increase following spring fire, decrease following summer
fire), but these differences faded within 1 year. Richness in these grasslands
was consistently negatively correlated with proportion of E. lehmanniana
biomass to total biomass regardless of year, season, community, or fire
treatment. Additionally, richness appeared to be influenced more by yearto-year variability than by fire treatment. A decrease in total biomass was
maintained for two fall seasons following fire, a response that varied by
treatment season and year. Because of variability of responses across years,
continued experimental research is needed before fires may be restored on
a large scale.
GEIGER, JOHN H.* Florida International University, Miami, Fl., USA.
Investigation of Allee effects in the endangered vine Ipomoea microdactyla (Convolvulaceae).
Empirical evidence showing habitat fragmentation results in population genetic consequences for plants is rapidly accumulating. The effects have
been variable, most often negative but also positive. Habitat fragmentation
followed by reductions in population size generally leads to decreases in
genetic variation, both allelic richness and heterozygosity of individuals.
The danger to the fragmented populations may be reductions in the fitness
of individuals and the immediate viability of these populations. The purpose of my study is to gauge the relationship between genetics and demography for a threatened species. Ipomoea microdactyla (Convolvulaceae) is a hermaphroditic perennial vine. In the United States, it occurs
only in the pine rockland habitat of Miami-Dade county; it also occurs in
Cuba and the Bahamas. This species is Florida state listed as endangered
and has populations at ten conservation areas in the county, including Everglades National Park. Presently, there is less than 2 % of the original
pine rockland habitat left, outside of the large habitat block protected in
Everglades National Park. The goal of my project is to conduct a population
viability analysis (PVA) to determine which factors are important for the
continued persistence of this species in Florida. The first part of my research will be a breeding system experiment to gauge self-compatibility/
self-incompatibility. This entails a protocol of hand pollinations at several
levels: the individual, within population, and among populations. The second part of my project will be a multi-year demographic study at all ten
conservation areas. The final component will be a genetic study to determine the spatial pattern of genetic variation and then incorporate this into
the PVA. Preliminary results from the breeding system experiment suggest
this species is self-incompatible and unable to set fruit without a pollen
vector. This implies potentially severe Allee effects for these low-density
populations.

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119

GENKAI-KATO, MOTOMI* and STEPHEN R. CARPENTER. University

of Wisconsin-Madison, Madison, Wisconsin. Effect of macrophytes on
lake eutrophication and restoration in relation to lake morphometry.
In eutrophic lakes, phosphorus (a primary cause of algal blooms) is supplied internally from the lakes sediment, as well as externally from the
lakes watershed. Phosphorus recycling from the sediment can cause two
different states of water clarity: clear-water and turbid-water states. Since
shifts between the states are often abrupt events, phosphorus recycling
makes restoration management more difficult. In shallow lakes, an inverse
relationship between water clarity and macrophyte dominance is often reported. Macrophytes in the littoral zone can prevent phosphorus from resuspension by stabilizing the sediment. However, the effect of macrophytes
should depend on lake morphometry determining the proportion of the
littoral zone. In addition, water temperature in the hypolimnion is known
to be an important determinant of phosphorus recycling. Here we look at
the effect of macrophytes on water clarity along a gradient of eutrophication, in relation to lake morphometry and water temperature. The macrophyte effect was critically dependent on mean depth and temperature, while
lake area hardly altered the effect. Lakes with intermediate mean depth
(;10 m) are most susceptible to eutrophication. In these lakes, restoration
by reducing phosphorus input is most likely to end in failure. Shallow lakes
(,2 m in mean depth) are resistant to eutrophication due to the presence
of macrophytes. A deep lake (.20 m) is also resistant because of its great
water mass in the hypolimnion. Eutrophication is more likely and therefore
restoration is more difficult in a warmer lake.
GEORGE, KATE,1 NATHAN PHILLIPS,2 THOMAS PYPKER,1 ANDREW SCHAUER3 and BARBARA J. BOND.1 1 Oregon State University,
Corvallis, OR; 2 Boston University, Boston, MA; 3 University of Utah, Salt
Lake City, UT. Environmental regulation of water flux in young and
old growth Douglas-fir trees over five years.
In order to predict vegetation responses to climate change in the future we
need to understand how dominant species at different developmental stages
will respond to variations in environmental conditions. We analyzed the
extent to which year-to-year variations in water flux could be explained by
differences in vapor pressure deficit (VPD) and soil moisture. We measured
daily water flux per unit sapwood area in young (approximately 25 years
old) and old growth (approximately 450 years old) Douglas-fir (Pseudotsuga menziesii) trees over five years (1998-2002) using constant heat sap
flow sensors. Water flux of old growth Douglas-fir became asymptotic at
higher values of VPD, whereas young Douglas-fir did not reach an asymptote. At higher values of VPD water flux in young Douglas-fir was
approximately 50% greater than old growth trees. The water flux of young
and old growth trees was significantly different between years at high values of VPD. The difference in water flux between years in a high VPD
environment can be explained in part by soil moisture. When soil moisture
was 30% or greater for part or all of the growing season maximum water
flux was higher compared to drier years for both young and old growth
trees. When water flux is scaled to the water use of all Douglas-fir present
within a stand, young Douglas-fir use approximately four times more water
than old-growth Douglas-fir on a ground area basis. On a stand level basis
the water use of young Douglas-fir stands are constrained by soil moisture,
whereas water use of old growth Douglas-fir stands are not altered by
seasonal and annual changes in soil moisture, even though there are no
differences between soil moisture at both sites. The increasing dominance
of young Douglas-fir stands and reduction of old growth stands in the
Pacific Northwest may significantly alter the hydrology of these ecosystems.
GEORGE, SOPHIE B.* Georgia Southern University, Statesboro, Georgia,
USA. The ectosymbiont crab Dissodactylus mellitae-sand dollar Mellita
isometra relationship.
This study investigated dispersion and reproduction of a small commensal
crab and their effects on the sand dollar Mellita isometra. The number of
crabs found on the sand dollar, stage of maturity, sex, numbers of eggs,
and size of eggs produced by crabs were noted over a two year period.
The test diameter of 2648 sand dollars, the number of eggs produced, and
the diameter of 4259 eggs was also noted. Infestation rates of crabs on

120

Abstracts

sand dollars varied over time. The three types of population dispersion;
random, uniform, and clumped were observed for crabs on sand dollars
throughout the sampling period. Clumped distribution was observed when
juvenile crabs were abundant, uniform and random distribution when adult
crabs were abundant. Female crabs were significantly larger than male
crabs. Crabs produced between 80 and 300 eggs ranging from 0.188 mm
to 0.291 mm in diameter. Bigger crabs produced significantly more eggs
than smaller crabs. The presence of crabs on sand dollars affected total egg
production of sand dollars in May, the peak of the spawning season, with
sand dollars carrying one or two crabs having a lower egg production than
those without crabs. The present study though not conclusive clearly demonstrates that a small percentage of the variation in egg size observed for
M. isometra might be due to seasonal variation in the distribution and
abundance of the commensal crab Dissodactylus mellitae.
GERACI, CHRISTY J.1,2,* 1 University of North Carolina at Chapel Hill,
Chapel Hill, NC, USA; 2 Clemson University, Clemson, SC, USA. Land
cover effects on benthic macroinvertebrate community structure in
Orange County, NC.
This study examined the relationship between land cover and benthic macroinvertebrate community structure in the Little Creek and Morgan Creek
Watersheds of Orange County, North Carolina. Rootwad and riffle habitats
were sampled at 18 sites along five creeks using D-frame sweep nets and
kick nets. Land cover percentages in the upstream drainage basin and 60m
riparian corridor of each site were calculated from classified Enhanced
Thematic Mapper (ETM) imagery of the Chapel Hill Area from July 1999
(Row 35, Path 16). Linear regression models showed stronger relationships
between land cover and benthic macroinvertebrate community structure in
rootwads than in riffles. At the upstream drainage basin scale, percent forest
cover and percent urban land cover were both strong predictors of taxa
richness, Shannon diversity, and abundance in rootwads, but were only
weakly related to taxa richness and diversity in riffles, and were not related
to abundance in riffles. Similar results were found at the 60m riparian
corridor scale. The results of this study suggest that 1) the benthic macroinvertebrate community in rootwads is more reflective of altered hydrology and non-point source pollution than the riffle communities in these
watersheds, and 2) land use planning techniques that preserve forest patches throughout the watershed are equally important to preserving aquatic
biodiversity as those that restrict development only within a specified distance from stream channels.
GERHARDT, FRITZ* and SHARON K. COLLINGE. University of Colorado, Boulder, Colorado. Causes of rarity in the vernal pool endemic
Lasthenia conjugens (Asteraceae).
Rare species are usually defined by limited distributions and narrow habitat
specificities, yet the mechanisms causing rarity remain unclear. Narrow
habitat specificity can arise because species are adapted to a particular set
of environmental conditions or because these habitats serve as refuges for
species that are poor competitors elsewhere. In this paper, we report on
field surveys and greenhouse experiments examining rarity in the vernal
pool endemic Lasthenia conjugens (Asteraceae, Contra Costa goldfields).
We examined the effects of abiotic conditions, especially water depth and
duration of inundation, and interactions with other native and exotic plants
on the performance of L. conjugens. Field surveys showed that L. conjugens was most abundant in natural pools and in pools that burned in a
wildfire in June 2000. Furthermore, L. conjugens abundance increased with
increasing water depth. In a greenhouse experiment examining the effects
of water depth and duration of inundation, L. conjugens had low overall
survival (47%), but height, number of leaves, total biomass, and number
of inflorescences all decreased with increasing water depth. Duration of
inundation had no significant effects on the performance of L. conjugens.
In a greenhouse experiment examining the effects of biotic interactions,
the biomass of L. conjugens decreased sharply as the biomass of neighboring plants increased. Collectively, these results show that, although L.
conjugens occurs at greater water depths in the field, this species actually
grows larger and reproduces more when not inundated. However, competition with native and exotic plants appears to limit L. conjugens to more
stressful, inundated sites. Thus, we conclude that L. conjugens and possibly
many other rare species are not narrow habitat specialists but rather are

refugees restricted to more stressful habitats by competition in other, more

favorable environments.
GERLACH, JOHN P. and MICHAEL B. WALTERS.* Michigan State University, Dept. of Forestry, East Lansing, Michigan, USA. Productivity and
leaf/canopy traits for deciduous and evergreen conifers over a soil resource gradient.
Studies have shown that productivity can be similar for evergreen and
deciduous conifers on a given site, but how productivity and related leaf
and canopy traits change over soil resource gradients is less well known.
For the deciduous species European larch (Larix decidua) and the evergreen species red pine (Pinus resinosa), based on reported differences in
leaf and canopy traits, we hypothesized a site quality by species interaction
with larch having greater productivity at high soil resource availability and
red pine having greater productivity at low soil resource availability. We
tested this hypothesis with soil, vegetation, and productivity data collected
from 27 geographically paired larch and pine plantations distributed over
the Great Lakes region. Our gradient was characterized by autocorrelation
of potential N mineralization, and nitrification with available soil water
holding capacity (AWC). Over the entire range of AWC, larch had greater
productivity (site index and net primary productivity), and there was a weak
species by AWC interaction with larch site index increasing more with
AWC than pine. Over the AWC gradient larch had greater % leaf N, and
annual N loss, and lower area based leaf N, and canopy N mass than red
pine. Foliar carbon isotope discrimination (D) indicated that larch was less
water use efficient that pine over the AWC gradient, but D did not vary
with AWC. For both species, leaf N (both area and mass bases) was more
strongly related to site index than any other leaf/canopy characteristic and
leaf N increased with AWC. Our results do not support a productivityresource availability tradeoff between larch and pine.
GERMINO, MATTHEW J.,1,* WILLIAM J. SMITH,2 BRODERSEN
CRAIG2 and DAN JOHNSON.2 1 Idaho State University, Pocatello, ID;
2
Wake Forest University, Winston-Salem, NC. Photosynthetic variation
in young and old conifers across an alpine-treeline ecotone.
Whether reduced carbon assimilation contributes to exclusion of trees from
subalpine and alpine meadows is currently debated, yet there have been
few or no reports of photosynthesis in treeline conifers across the gradient
from forest to alpine. Moreover, most studies of factors affecting treeline
have focused on responses of adult trees to high-elevation climates, even
though young seedlings may represent a critical life stage for determining
conifer establishment above forest elevation limits. Microclimate, photosynthetic carbon uptake, chlorophyll fluoresence, and plant water potential
were measured in newly-germinated seedlings (germinants), 3-10 yr old
seedlings, and adult Picea engelmannii from subalpine forest across the
treeline ecotone up to the alpine of the Snowy Range of Wyoming, USA.
Mean photosynthesis was greater for adult trees, marginally lower for seedlings, and ;50% lower for germinants near the alpine compared to lower
elevation sites. Similarly, predawn Fv/Fm (an indicator of photosynthetic
light-use efficiency) did not vary in adults along the gradient, but was over
18% lower in germinants in the alpine compared to lower sites. Mean Fv/
Fm remained above 0.75 for adults, but became as low as 0.55 for germinants, indicating greater photoinhibition in the colder, brighter alpine
compared to more-forested sites, as well as in the younger compared to
older individuals. These results indicate that young, establishing seedlings
may be more likely than mature trees to experience carbon limitations
above the forest elevation limit.
GHASSEMZADEH, FERESHTEH,* M.H. ARBAB ZAVAR and M.H.
KARIMPOUR. Faculty of Science, Ferdowsi University of Mashhad,
Mashhad, Khorasan, Iran. Environmental studies of arsenic in the Chelpo area, northeast Iran.
Arsenic is ubiquitous in the environment, and exposure to it can occur
from natural and anthropogenic sources. Human exposure occurs from air,
food and drinking water. Drinking water is the most significant source
worldwide. The aim of this study was to determine arsenic concentration
in drinking water sources in the Chelpo area of northeast Iran. This region
is naturally rich in arsenic. We also conducted this study to analyze mor-

tality and causes of death, especially from cancers and noncancer diseases.
Arsenic concentration was determined in fourteen water samples and sixteen soil samples from the study area. Four villages located in the study
area were selected for the study on the basis of a survey of arsenic pollution
and mortality from cancer and noncancer diseases. The total arsenic concentration in the water samples ranged from 8 to 27 ppm and in the soil
samples ranged from 210 to 26x104ppm. Arsenic concentrations in the
drinking water mostly exceeded WHO guidelines. Arsenic toxicity consists
mostly of skin lesions and carcinogenesis. Digestive cancer is the most
clearly associated mortality related to arsenic pollution. However, skin lesions and other diseases have also been found in the study area. About
20% of the mortality is due to cancer, which is probably related not only
to arsenic pollution but also to nutritional deficiencies. Greater mortality
was found in males with digestive cancers, which is probably related to
the drinking more of water and tea by men.
GIBSON, DAVID J.,1,* BETH A. MIDDLETON,1,2 KARI FOSTER,1 YOHANES A.K. HONU,1 ERIK HOYER1 and MARILYN J. MATHIS.1,3
1
Southern Illinois University, Carbondale, IL; 2 National Wetlands Research Center, Lafayette, LA; 3 Howard Payne University, Brownwood, TX.
Core and satellite species dynamics in response to disturbance and
fertilization during early old-field succession.
The Core-Satellite Species hypothesis (CSS) provides a metapopulation
perspective for understanding community patterns. We used the CSS to
evaluate the pattern of species richness dynamics at two scales during early
old-field succession in southern Illinois. Species richness was recorded
from the initial seedbank and during the first 7 years following plowing of
both a post-agricultural upland and lowland field. Fertilizer and disturbance
treatments (mowing and rototilling) were established in 25 m 2 plots arranged according to a randomized complete block design in 225 m2 blocks.
There was a significant interaction between disturbance and time
(p,0.0001) on species richness in both fields, between disturbance and
fertilizer in the lowland old-field (p50.004), and fertilizer alone in the
upland old-field (p,0.0001). At the block scale, a bimodal pattern of species richness consistent with the CSS was observed both in the initial seedbank and through the course of early succession in both fields. The identity
of native and exotic core species (present in $ 90% of blocks) changed
through time, which was consistent with predictions from established models of succession. Some core species from the seedbank became core species in the vegetation, albeit after several years. At the smaller plot scale,
the CSS model rarely applied. In the lowland field, species richness patterns
were predominantly unimodal reflecting large numbers of satellite species
(present in #10% of plots). In the upland field, species patterns were also
predominantly unimodal, but exhibited bimodalality consistent with the
CSS during the first two years following plowing, becoming more strongly
unimodal through time. These findings reflect the need for a spatial and
temporal scaling perspective in utilizing the CSS model. The results suggest
the operation of regional processes (e.g., propagule availability) at the largest scale in secondary succession. At smaller scales, local processes (disturbance and soil nutrients) are important and place a constraint on the
colonization of both native and exotic species.
GILADI, ITAMAR* and H. R. PULLIAM. Institute of Ecology, The university of Georgia, Athens, GA, USA. Habitat specific dispersal and its
implication for population dynamics.
As many other ecological processes, dispersal exhibits spatial and temporal
heterogeneities. The significance of dispersal events for population dynamics can be evaluated only from studies that include the spatial and temporal
variation in dispersal and population demography. Over the last three years,
I have studied habitat-specific demography and habitat-specific dispersal of
a small herbaceous ant-dispersed forest plant, Hexastylis arifolia. The study
was conducted at Whitehall forest, near Athens, GA. I have studied demography and seed dispersal in mature oak-hickory forest, the typical habitat type of H. arifolia. In addition, I have studied dispersal and population
demography in three marginal habitats where H. arifolia is usually absent.
Population demographic parameters in the main habitat were estimated
from annual census of individual plants in permanent plots. Plant introduction experiments provided the data for estimating demographic parameters in the marginal habitats. In all habitat types, seed dispersal was mea-

Abstracts

121

sured directly by following ants dispersing seeds from randomly located

points. The results from the dispersal experiments show that the proportion
of dispersed seeds and the distribution of dispersal distances differ among
habitats. These differences were mainly attributed to differences in ants
activity in different habitats. Population growth rate in the permanent plots
and one marginal habitat was close to one, while populations in the other
marginal habitats have growth rates lower than one. Combined, these results suggest that H. arifolia is present in mostly suitable habitat, is absent
from low quality habitat, but is also absent from high quality habitat that
lack the appropriate dispersers.
GILBERT, BENJAMIN D.* and MARTIN J. LECHOWICZ. McGill University, Montreal, Quebec, Canada. The relative importance of spatial
and environmental processes in structuring diversity on a forested
landscape.
Niche specialisation and neutral interactions among dispersal limited species are two competing hypotheses that attempt to explain plant species
diversity and community membership. We tested these hypotheses at Mont
St. Hilaire: a temperate, old growth forest near Montreal, Quebec that contains approximately half of the regional flora in an area of 10 km2. We
used a sampling technique that decoupled spatial and environmental patterns on the landscape and incorporated diverse plant functional groups
(trees, shrubs and herbs) and phylogenetic groups (Asteraceae, Cyperaceae,
Poaceae, and seedless vascular plants). Constrained ordinations and distance decay tests were used in conjunction with path analyses to determine
the relative importance of each contributing factor. Tree community composition was structured mainly along environmental gradients, but also
showed evidence of dispersal limitation. The tree functional group was the
only group that showed support for the neutral theory. The other groups
were predominantly structured by environmental gradients, although a distance by environment interaction was sometimes present. Niche partitioning
differed among groups, with trees and shrubs mainly structured by light
availability, and understory plants mainly structured by water and pH or
nitrate. Our results indicate that dispersal limitation in this old growth forest
is either nested within or secondary to environmental gradients. Models
that attempt to predict species richness and species turnover must incorporate a better understanding of fine scale environmental differences, and
incorporate space by environment interactions.
GILBERT, JANICE,1,* SERITA D. FREY2 and VIRGINIE BOUCHARD.1
Ohio State University, Columbus, OH; 2 University of New Hampshire,
Durham, NH. Linking macrophyte diversity and denitrification in wetland ecosystems.

Our objective was to determine the impact of macrophyte functional group

diversity on N2O and N2 fluxes in two mesocosm experiments in which we
manipulated the number (0-5) of macrophyte functional groups (tussock,
reed, facultative annuals, obligate annuals, and clonal dominants). Our hypothesis was that a shift in functional group diversity would significantly
impact N2O and N2 fluxes due to changes in the quantity and quality of
plant biomass inputs into the soil and porewater organic matter pools. Experiment 1 included the 5 functional groups listed above while Experiment
2 included the same groups minus the clonal dominants. We measured in
situ N2O and N2 production under nitrate limiting and non-limiting conditions, denitrifying enzyme activity (DEA), pore water organic C, sediment labile C, and sediment bacterial biomass. We found distinct differences in results between the two experiments. In Experiment 1 DEA was
significantly higher in the highest diversity treatment (P,0.05). There was
a strong correlation (R25 0.63) between bacterial biomass and root biomass
(the combination with 5 functional groups supported the highest bacterial
biomass) and labile C and root biomass (R25 0.70). For Experiment 2, we
found no relationship between functional group diversity and DEA fluxes.
The highest DEA rates occurred with the reeds (179 mg N m22day21), and
lowest with the facultative annuals and controls (68 and 85 mg N
m22day21). We found negative correlations between DEA and bacterial biomass (R25 0.36), bacterial biomass and root biomass (R25 0.28) and DEA
and TOC (R25 0.69). We also found no significant differences between
treatments for labile C and total plant biomass (p.0.05). We conclude that
macrophyte community composition rather than functional group diversity
impacts denitrification and has potentially important implications for wetland mitigation and global climate change.

122

Abstracts

GILL, RICHARD A.,1,* H. W. POLLEY,2 HYRUM B. JOHNSON2 and

ROBERT B. JACKSON.3 1 Washington State University, Pullman, WA;
2
USDA-ARS, Grassland, Soil & Water Research Lab, Temple, TX; 3 Duke
University, Durham, NC. Short- and medium-term carbon and nitrogen
dynamics in a grassland exposed to past and future atmospheric CO2.
Many important ecosystem processes have been influenced by historical
increases in atmospheric CO2 which may lead to carbon sequestration in
terrestrial ecosystems. Most plants exhibit increased C assimilation in response to experimentally doubled carbon dioxide, but it is unclear whether
increased production can be maintained long-term because of decreases in
mineral nutrient availability. It is also unclear whether increases in photosynthesis will result in ecosystem sequestration of C in soil pools with
turnover times of decades to centuries. Our field experiment in an intact
C3/C4 grassland in central Texas is unique in providing a continuous gradient of atmospheric CO2 from 200 to 550 ppm, allowing us to examine
critical threshold and nonlinear responses to past, present, and future atmospheric CO2. Along this continuous gradient, increased CO2 promoted
higher rates of decomposition in lab incubations for green, aboveground
tissue from the dominant C3 forb but had no significant effect on decomposition rates of the dominant C4 grass. We found a positive, linear increase
in the storage of C in fractions of soil organic matter (SOM) with decadal
scale residence time in soil, with the SOM pools with the shortest residence
time being most sensitive to changes in atmospheric CO2. However, the
increases in soil C with decadal scale residence time were offset by losses
in older soil C under elevated CO2, leading to no significant increases in
total soil organic matter. We also suspect that the previously reported decrease in soil N availability with rising CO2 may be explained by higher
rates of N immobilization during the decomposition of the dominant C3
forb.
GILLIAM, FRANK S.1,* and WILLIAM J. PLATT.2 1 Marshall University,
Huntington, WV; 2 Louisiana State University, Baton Rouge, LA. Distinguishing between desirable and undesirable old-growth overstory systems as goals of conservation and management.
Knowledge of the structure and composition of old-growth longleaf pine
stands is imperative in the conservation and management of longleaf ecosystems. Among the challenges to our understanding of the ecology of oldgrowth longleaf sites, however, is the great degree of variability among
sites, something that arises out of spatial variation in several factors, the
most important of which is disturbance regime. Although once viewed as
early-successional forests of low diversity, longleaf pine ecosystems are
now seen as high-diversity, persistent savannas maintained by disturbance
and pyrogenic vegetation, containing numerous endemic species. We distinguish between desirable versus undesirable characteristics of old-growth
longleaf pine systems by comparing two sites that differ in disturbance
regime: the Boyd Tract (NC) and the Wade Tract (GA). The Boyd Tract
has undergone chronic fire suppression and occurs in the Sandhills, a region
that experiences hurricanes infrequently. The Wade Tract has undergone
annual prescribed burning and has historically experienced relatively frequent hurricanes. Differences between sites were seen more in stand structure than in overstory composition. Basal area of the Boyd Tract was over
three times that of the Wade Tract. Relative dominance of longleaf pine
also varied between sites, approximately 50 and 95% at the Boyd and Wade
Tracts, respectively. These sharp contrasts in stand structure and longleaf
pine dominance between the Boyd and Wade Tracts demonstrate the importance of large-scale disturbance, especially hurricanes and fire, in shaping the structure and function of longleaf pine ecosystems of the southeastern United States. In particular, long-term fire exclusion the Boyd Tract
has altered stand structure substantially by decreasing longleaf pine regeneration. Reintroduction of fire accompanied by physical removal of dense
turkey oak stems at the Boyd Tract resulted in only transient increases in
longleaf seedling density. Thus, chronic fire exclusion can bring about longterm changes in longleaf pine stand structure.
GILLOOLY, JAMES F. Biology Dept., Univ. of New Mexico, Albuquerque, NM, USa. The allometry of stoichiometry.
Recent work in stoichiometry has highlighted the relationship between
growth rate, RNA content and whole body P concentration in aquatic or-

ganisms (i.e.,"growth rate hypothesis). Here we show how growth rate and
the related stoichiometry is related to metabolic rate, and thus to body size.
We present a model that predicts the scaling of RNA content and whole
body phosphorous concentration to body size. We test the model using data
from various aquatic organisms that vary in trophic level. Results suggest
that energy and materials are not alternative or complimentary currencies,
but that they are inextricably linked.
GINZEL, MATTHEW D.,1,* JOCELYN G. MILLAR2 and LAWRENCE
M. HANKS.1 1 University of Illinois at Urbana-Champaign, Urbana, IL;
2
University of California - Riverside, Riverside, CA. Chemically-mediating mate recogntion in the longhorned beetles (Coleoptera: Cerambycidae).
The wax layer on the cuticle of insects is comprised of a complex mixture
of long-chain fatty acids, alcohols, esters, aldehydes, ketones, and hydrocarbons that protect insects from desiccation. Components of the wax layer
also may have a secondary role as contact pheromones. There is a growing
body of evidence that mate recognition in cerambycid beetles is mediated
by such contact sex pheromones. We add to our previous work on cuticular
hydrocarbons of cerambycids and indentify the contact pheromone of the
locust borer Megacyllene robiniae (Forster). Male M. robiniae respond to
females only after touching them with their antennae, indicating that mate
recognition is mediated by a contact sex pheromone. GC-MS analyses of
whole-body extracts revealed compounds present in the wax layer of females that are absent or present in smaller quantities in males. We reconstructed blends of the most abundant female specific straight-chain (nC23,
nC24, nC25, nC26), methyl-branched (3me-C23, 3me-C25), and unsaturated
(Z9:C23, Z9:C25, Z9:C27) compounds and assessed their bioactivity in arena
bioassays. In these bioassays, males showed the greatest response to Z9:
C25 alone. This compound elicited the complete sequence of mating behaviors, up to and including coupling the genitalia. Z9:C25 comprised 16.4
6 1.3% of the total hydrocarbons while in solid phase microextraction
wipe samples of the cuticle, Z9:C25 comprising 34.6-38.3% of the sampled
hydrocarbons. Our data indicate that Z9:C25 is the contact sex pheromone
of M. robiniae, being the most abundant hydrocarbon on the cuticular surface of females where it is readily accessible to the antennae of males.
GITTINGER, LORI S.,1 DAVID M. SOBALLE2 and JOHN H. CHICK.1
1
Illinois Natural History Survey, Great Rivers Field Station, 8450 Montclair Avenue, Brighton, Illinois, USA; 2 United States Geological Survey,
Environmental Management Technical Center, 575 Lester Avenue, Onalaska, WI, USA. Twelve years of water quality trends in Pool 26 of the
upper Mississippi River.
The upper Mississippi River is a major resource for multiple uses, including
navigation, water supply, habitat for fish and wildlife, and recreation. In
order to effectively manage this resource, the Long Term Resource Monitoring Program (LTRMP) was established to enhance understanding of this
ecosystem and provide information needed for sound management. The
LTRMP has accumulated over 10 years of limnological data, providing a
comprehensive and unequalled view of the water quality of the upper Mississippi River. The Great Rivers Field Station, as part of the LTRMP, has
performed basic field and laboratory water quality measurements on Pool
26 of the Missisippi River, along with its tributaries and backwater areas.
One of the benefits of such a long-term database is the ability to detect
trends or change over time. We used time-series analysis on a number of
important water quality measurements (e.g. dissolved oxygen, nitrogen,
phosphorous, etc.) to determine whether there were any detectable trends,
either poolwide or habitat-specific, over the past 12 years. We found a
number of distinct trends: total nitrogen, soluble reactive phosphorous and
chlorophyll-a showed distinct downward trends poolwide over the past 12
years. Habitat-wise, dissolved oxygen showed a distinct upward trend in
backwaters over the past 12 years. Several water quality parameters also
showed distinct seasonality, some of which varied among habitats. We believe the ability to detect trends attests to the value of the LTRMP water
quality database. Whether these trends will continue can only be determined through continued monitoring of this important resource.

GIVLER, KIM A.,1,* SCOTT C. NEUBAUER,1 MASOUMEH SIKAROODI,2 DAVID EMERSON,3 PATRICK M. GILLEVET2 and PATRICK MEGONIGAL.1 1 Smithsonian Environmental Research Center, Edgewater,
MD; 2 George Mason University, Manassas, VA; 3 American Type Culture
Collections, Manassas, VA. Microbial carbon metabolism along a salinity gradient: coupling biogeochemistry to microbial community composition.
The dominant pathways of anaerobic carbon metabolism follow wellknown trends across a salinity gradient. Methanogenesis dominates where
salinity is low, but is suppressed by sulfate reduction as salinity increases.
In mineral soils, Fe(III) reduction is expected to dominate both methanogenesis and sulfate reduction. Our goal was to determine whether changes
in microbial community composition reflect these well-understood shifts in
biogeochemical activity. Microbial communities were studied by Most
Probable Number (MPN) culture techniques and Amplicon Length Heterogeneity (ALH) fingerprinting, and standard biogeochemical techniques
were used to quantify methanogenesis, Fe(III) reduction, and sulfate reduction. Duplicate cores were taken at a series of eight tidal wetland areas
ranging from fresh to mesohaline along the Patuxent River estuary, Maryland. Sulfate reduction rates increased three orders of magnitude from the
least saline to the most saline site (0.00031 mgSO4 cm23d21 at 0.4 to
0.12 mgSO4 cm23d21 at 15), with an associated two-fold decrease in
methane production (0.0032 umolCH4 g21 hour21 at 0.4 to 0.44 umolCH4
g21 hour21 at 15.2). However, these trends were not reflected in the MPN
cell counts, which showed no obvious variations in sulfate-reducing bacterial communities. MPN counts for Fe-reducing bacteria were below detection in most of the freshwater marsh samples, although iron reduction
rates (13 umol Fe/g soil/d) were among the highest observed at any of the
sites. Large variations in biogeochemical activity were not reflected in the
community composition of culturable microorganisms. We anticipate that
the more sensitive ALH fingerprinting technique in progress will be able
to detect changes in microbial biological diversity.
GLADWELL, RANDI R.1,* and YENNY IHUE-UMIRE.2 1 University of
New Mexico, Albuquerque, NM, USA; 2 Universidad de San Augustin,
Arequipa, Arequipa, Peru. From coast to highlands: An elevation gradient survey of rodent communities in southwestern Peru.
A small-scale survey of rodent communities was conducted during July
2002 in three locations (Camana, 60 masl; Aplao, 650 masl; and Chuquibamba, 2700 masl) in the department of Arequipa on the western flanks of
the Andes Mountains in Southern Peru. The goal was to explore ecological
and historical impacts on plant and animal species by humans and human
associated activities. We documented the species diversity within rodent
and plant communities present in the narrow river valleys of the Majes
Drainage System; an area highly modified by humans practicing agriculture
ino events, there was
for more than 1500 years. With the exception of El N
no significant precipitation on the west coast of Peru due to geographic
and climatic factors that involved the combination of the Humboldt Current
and rapid ascent of the Andes Mountains. As a result, these western slope
river valleys are ecologically constrained due to this limited access to water
and the biota depends on water originating from the highlands. Vegetation
is limited only to the bottom of narrow river valleys and rarely grows on
valley slopes. With supporting archaeological evidence, our findings suggest that commensal rodents (e.g., Mus musculus and Rattus rattus) that
arrived with Europeans have aggressively replaced endemic rodent communities from these low elevation valley bottoms. Higher elevation rodent
communities have greater species diversity and no commensal rodents were
encountered, possibly due to geographic barriers. The hypothesis that climatic change is an agent and factor of ecological changes suggested here
has not been falsified.
GLEESON, SCOTT K.1 and DAVID TILMAN.2 1 University of Kentucky,
Lexington, KY; 2 University of Minnesota, St. Paul, MN. Nitrogen saturation and resource isoclines during grassland succession.
Although nitrogen is often retained in terrestrial ecosystems at low rates
of deposition, at some point, the point of nitrogen saturation, most added
N is leached from the ecosystem. Just such a pattern was observed in the
grasslands of Cedar Creek Natural History Area in response to experimen-

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123

tal N addition. Here we propose that the N-release threshold corresponds

to the switch point on a community-wide zero net growth isocline (ZNGI)
where limitation changes from nitrogen to light. Community-wide isoclines
for three of four grassland communities are sharply rounded, indicating a
nitrogen saturation threshold and suggesting that vegetation dynamics, rather than microbial processes, may be driving nitrogen retention and leaching
in these ecosystems. The shapes of the observed community-wide isoclines
are consistent with the levels of limiting resources being determined by N
versus light limitation and resulting shifts in plant mass in roots versus
shoots. N leaching roughly corresponds the expected transition to light
limitation at higher rates of N addition.
GLOWACKI, GARY A.* and YOUNG D. CHOI.* Purdue University Calumet, Hammond, IN, USA. Restoration model for a degraded riparian
wetland.
Wetlands in North America have suffered great losses. While restoration
of wetlands is a management priority, a large number fail due to lack of
adequate models to follow. Many restoration efforts focus solely on vegetation composition and do not attempt to emulate the overall structure and
dynamics of the ecosystem. To develop a restoration model for a degraded
58 ha riparian forest along the Little Calumet River, we examined two
remnant riparian forests within the boundaries of the Indiana Dunes National Lakeshore. The water table, soil characteristics and vegetation were
measured at 80 plots at both the remnant and degraded sites. Large canopy
trees (.2m) were dominated by Acer rubrum and Ulmus americana at the
remnant sites, while Populus deltoides and Fraxinus americana dominated
the degraded sites. Beneath the canopy, herbaceous species diversity was
greater at the remnant site (H95 2.63) compared to the degraded site
(H951.86). Non-metric multidimensional scaling (NMS) of herbaceous
vegetation showed that axis 1 was mainly factored by Typha spp.
(r520.621) and Phragmities australis (r520.571), while axis 2 was mainly factored by Urtica procera (r5 0.673) and Poa pratensis (r5 20.637)
at p,0.05. Water tables of the remnant sites (mean 6.25cm + 3.7 below
ground) were significantly higher and more fluctuating than the ones of the
degraded sites (mean 16.16cm +4.2 below ground) particularly in early
spring. Spring floods likely discouraged invasions of shrubs in the remnant
sites, as density of shrubs was greater at the degraded sites (1.58 stems/
m2) as opposed to the remnant site (0.53 stems/m2). It appeared to be
essential to restore natural hydrology prior to removal of exotic species
and restoration and native vegetation in our degraded sites.
GODDARD, KELLY L.,* DAVID D. BRESHEARS,* SCOTT N. MARTENS,* KATHERINE E. DAYEM and CLIFTON W. MEYER. Los Alamos National Laboratory, Earth and Environmental Sciences division, Los
Alamos, NM. Drought recovery via foliar water absorption: Bypassing
the soil uptake pathway.
Woody plants benefit form large pulses of water that penetrate deeper into
the soil profile. During drought, such pulses are non-existent and the only
precipitation that may occur is likely to be in the form of small precipitation
events. Small precipitation events are likely to penetrate the soil only to
near-surface depths and to be subject to rapid soil evaporation. Here we
document a mechanism for addressing drought recovery via foliar absorption in a woody juniper species, Juniperus monosperma. We excluded precipitation by installing a shelter located beneath the tree canopies and above
the soil. Predawn plant water potentials of treated trees tracked control trees
during the first monthly interval after initiation of rain, suggesting that
100% of the recovery could be due to foliar absorption. They exhibited
;33% of the recovery of control trees the following month. Hence nearly
50% of the overall recovery was attributable to foliar absorption. To confirm that foliar absorption was occurring and was important during drought,
we compared control branches with treated branches, which we poured
water over to establish foliar uptake. We found that treated branches responded significantly relative to control branches. This work highlights an
important mechanism of water uptake in a semiarid species that allows
bypassing of soil water uptake.
GOERIZ, RACHEL E.,* MARK FOX and ROBERT F. DENNO. University of Maryland, College Park, MD. Omnivory enhances growth and
diminishes mortality in a marsh-inhabiting grasshopper.
The mismatch in C:N stochiometry between insect herbivores and their
host plants strongly suggests that herbivores are nitrogen limited. Omni-

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vory has evolved in some herbivore lineages, and the occasional consumption of nitrogen-rich prey may help offset the inherent penalty many herbivores pay for consuming only plant food. Thus, dietary supplementation
through the consumption of prey items may increase fitness. We tested the
consequences of diet on the growth and mortality of an omnivorous grasshopper, the tettigoniid Conocephalus spartinae, that feeds on Spartina cordgrass as well on the herbivores (Prokelisia planthoppers) associated with
this marsh dominant. Grasshoppers were fed either Spartina or a combination of Spartina and planthoppers. Fed a cordgrass diet only, grasshoppers grew slowly and survived poorly compared to when they were offered
a mixed diet of cordgrass and planthoppers. The disappearance of planthoppers from treatments confirms that grasshoppers attack and consume
planthopper prey. In the field, grasshoppers occur most abundantly in lowmarsh habitats where cordgrass is most nutritious and planthoppers abound.
Moreover, the frequency of invertebrate parts in the gut contents of Conocephalus is positively related to the density of planthoppers across habitats. These data suggest that by tracking planthopper populations and selectively colonizing nutritious Spartina, grasshoppers can more easily meet
their nitrogen demands. Thus, dietary supplementation with prey and the
occurrence of omnivory may have been promoted in certain groups of
"herbivores" as means of increasing nitrogen intake in a carbon-dominated
world.
GOHEEN, JACOB R.,1,* ROBERT K. SWIHART2 and JAMES H. ROBINS.3 1 University of New Mexico, Albuquerque, NM; 2 Purdue University,
West Lafayette, IN; 3 Northern Illinois University, DeKalb, IL. Ecological
implications of red squirrel range expansion into the central hardwoods region of the United States.
Species with expanding ranges provide unique opportunities to examine
environmentally induced adaptations in ecological characteristics and behaviors. Since the late 1800s, the North American red squirrel (Tamiasciurus hudsonicus) has expanded its range into the central hardwoods of the
United States in conjunction with increasing agricultural fragmentation. We
conducted two experiments with regard to this range expansion. In the first
experiment, we investigated the dispersal behaviors of red, gray (Sciurus
carolinensis), and fox (S. niger) squirrels in west-central Indiana using a
series of experimental releases. We hypothesized that all species would
display increased willingness to disperse in the pre-harvest season and with
decreasing distance to forest patches. We hypothesized further that gray
squirrels would be inferior dispersers relative to red squirrels, given the
recent range expansion of red squirrels. All three hypotheses were supported. In the second experiment, we examined whether red squirrels from
the central hardwoods displayed differences in foraging behaviors and morphology relative to red squirrels from conifer-dominated environments, a
biome in which red squirrels evolved, in the upper peninsula of Michigan.
Red squirrels from Indiana and Michigan differed significantly in the efficiency with which they used food items, with individuals from each region more efficiently using those items to which they previously had been
exposed. The enhanced efficiency of southern red squirrels feeding on black
walnuts (Juglans nigra) presumably was due to differences in cranial morphology; skulls of southern squirrels were larger, with longer jaws and
higher metrics associated with greater mandibular force than northern squirrels. Contrary to our expectations, red squirrels from Indiana and Michigan
did not differ qualitatively in preferences for food items, suggesting that
diet choice is constrained by perishability of food items.
GOLUBIEWSKI, NANCY E.1,2,* and CAROL A. WESSMAN.1,2 1 University of Colorado-Boulder, Boulder, CO; 2 Cooperative Institute for Research in Environmental Sciences, Boulder, CO. Are urban forests in the
western United States a panacea for carbon sequestration?
As the discussions over the potential consequences of climate change intensify, so do the debates over how to mitigate emissions of carbon dioxide
and other greenhouse gases. Afforestation has been proposed as one of
several mitigation strategies, and terrestrial carbon sequestration offers an
apparent panacea due to its provision of a potentially large carbon sink. At
the same time, urban sprawl continues apace, and the subsequent land transformation captures the attention of both the public and researchers. In the
semi-arid western U.S., this land transformation results in the afforestation
of metropolitan landscapes. This study examined the temporal and spatial

shifts in carbon storage in the Front Range of Colorado due to urban

sprawl. Previous field studies indicate that biomass and carbon differ significantly in grassland and urban environments. Whereas soil carbon decreases in human-dominated systems, vegetative carbon storage increases
substantially due to increased woody plant biomass. In this research, spatial
analyses of modern shifts in carbon storage were examined via spectral
mixture analysis of a series of satellite images,1970s-present. Field data
were used to validate results from the most recent imagery. A historical
analysis of land-cover change from aerial photos extended the study to an
aggregated evaluation of changes in carbon storage from 1930s-1990s. During the twentieth century, urbanization transformed a semi-arid grassland
into a deciduous forest. However, carbon sequestration resulting from this
afforestation may not be so much a panacea for greenhouse gas emissions
as a Pandoras box.
GOLUBSKI, ANTONIO J.* University of Illinois at Chicago, Chicago, IL.
Mutualistic traits can confer a selective advantage in a simple consumer-resource feedback model.
A central question regarding mutualisms remains what prevents "cheaters",
which reap mutualistic benefits but provide nothing in return, from enjoying a selective advantage. Consumer-resource feedback models are developed here that simulate two competing consumers; the more mutualistic
consumer gains less from a given resource patch but has a greater positive
impact on future resource availability, and the less mutualistic consumer
benefits the resource less but gains more from any given patch. Consumers
disperse inoculum (and win patches next time step) in the proportions with
which they obtain resources, and die at the end of each time step. In spatially explicit simulations, the more mutualistic consumer can exclude its
competitor; spatially implicit versions of the model show that local dispersal is required for this outcome. These results show that in principle a
mutualist can gain a selective advantage by sacrificing its ability to acquire
resources in order to stimulate future resource production. This mechanism
parallels those suggested to promote prudent predation or intermediate parasite virulence, but, as shown here, can operate without consumers having
net negative effects on the resource or displaying any particular behaviors
such as defending resource patches or abandoning patches with low resource abundance. Further, neither partner fidelity nor partner choice was
explicity included. The effects of consumers were also integrated over several patches to explore the significance of co-infection of a host by multiple
consumer types and the significance of non-additive effects of multiple
consumers; the results suggest that added benefit of multiple partners can
allow consumer coexistence. These models broaden the mechanisms by
which and conditions under which mutualistic traits might be expected to
be advantageous.
GOMEZ-SAPIENS, MARTHA* and ANGELINA MARTINEZ-YRIZAR.
Instituto de Ecologia, UNAM-Mexico, Hermosillo, Sonora. Woody species leaf traits: Changes from the desert to the tropical deciduous forest
in Sonora, Mexico.
Leaf structure, morphology and nutrient content have been used to establish
relations that can provide information about different plant strategies of
resource use in the ecosystem. In this study sun leaves of five perennial
woody species present along a precipitation gradient in Sonora, Mexico
were analyzed to test the relationship between leaf trait variability and
precipitation and nutrient availability patterns from a desertscrub to a tropical deciduous forest ecosystem. We expected i) a positive correlation between leaf nutrient concentration (total N y P) and available N and P soil
forms, and ii) the lowest specific leaf area (SLA) values in arid or low
nutrient availability sites. Fully expanded randomly selected leaves from
15 adult trees of each species were measured for thickness, total length,
leaf area, SLA, dry matter content (DMC) and total N and P content. Also,
ion exchange resin bags were used to determine the relative availability of
soil NH4, NO3 and PO4. We found that intraspecific differences in leaf
length and leaf area were significant across the sites, with the longest and
largest leaves in the tropical deciduous forest. Also, leaves with the highest
SLA and lowest DMC ratios were found in the deciduous forest. The trend
of change in total P and N concentrations along the gradient varied depending on the species. Relative availability of soil nutrients did not correlate with foliar nutrient concentrations. The variation in SLA seems to

have a strong relation with precipitation than with the relative nutrient
availability patterns.
GONELLA, MICHAEL P.1,2,* 1 Miami University, Botany Department, Oxford, Ohio, USA; 2 Myaamia Project, Miami Tribe of Oklahoma & Miami
University, Miami University, Oxford, Ohio. Ethnobotany of the Miami.
Documentation of traditional ecological knowledge (TEK) holds particular
value for indigenous communities pursuing academic venues of cultural
education and for determining the applicability of TEK to conservation on
tribal lands. The Miami communities of Oklahoma and Indiana have formalized their desire to document plant-related TEK and other cultural information in the formation of the Myaamia Project. Ethnographic research
on ethnobotanical TEK of the Miami and interviews with tribal members
has been initiated to document cultural uses of plants. Gathered data includes over 100 culturally important plant species, and related TEK, organized into a ethnobotanical database. Specific applications of ethnobotanical Miami TEK are being investigated through experiments reintroducing the traditional harvesting regimes on two common meadow plants,
common milkweed (Asclepias syriaca) and indian hemp (Apocynum cannabinum), and one common wetland plant, arrowhead (Sagittaria latifolia).
Experimental data are predicted to help management of these culturally
important plants and ensure that their historic abundances remain available
for use in contemporary Miami cultural practices.
GONZALEZ, ANDREW.* Universite Pierre et Marie Curie, Centre National de la Recherche Scientifique, Ecole Normale Superieure, Paris,
75230, France. Diversity, abundance and biomass dynamics in fragmented landscapes: insights from a bryophyte-based natural microcosm.
I provide a synthesis of the key results obtained from a series of experiments, using a bryophyte-based microecosystem, conducted to examine the
manifold effects of habitat fragmentation. This microecosystem is conceived as an experimental model system, and I will argue that the general
nature of the observed patterns and dynamics highlight its value as a natural
analogue for the effects of fragmentation at larger spatial scales. In particular I will focus upon the role dispersal plays in mediating the interaction
between local and regional community pattern and dynamics. Specifically
data will be presented showing how habitat fragmentation as a perturbation
induces a disassembly dynamic that is reflected in changes at the population
(abundances), community (diversity and biomass production) and metacommunity (range-abundance relation) level, and how habitat inter-connection can ameliorate these effects.
GONZALEZ-MELER, MIQUEL A.* Department of Biological Sciences,
University of Illinois at Chicago, Chicago, IL, USA. Introduction: Respiration across scales.
Terrestrial ecosystems exchange about 120Gt of carbon (C) with the atmosphere, through the processes of photosynthesis (leading to gross primary productivity, GPP) and ecosystem respiration (Re). Increasing evidence indicates that raising atmospheric CO2 enhances carbon uptake in
most ecosystems, however, responses of Re to elevated CO2 or temperature
are still unresolved due, largely, to our inability to reliably determine Re
and its autotrophic and heterotrophic components. Sources of Re are measured over a variety of spatial and time scales making integration of respiratory processes difficult. Rate-based measurements of components of
Re and meteorological-based measurements of nighttime Re have encountered technical caveats that has limited our potential for reliably quantifying
Re. Mechanistic understanding of respiratory processes at any temporal and
spatial scale is crucial for building robust process based models used in
prediction of the impact of global change on the terrestrial biosphere.
GOODELL, KAREN. Department of Ecology, Evolution, and Natural Resources, Rutgers University, New Brunswick, NJ, USA. Structure of bee
communities in calcareous fens invaded by purple loosestrife compared
to uninvaded fens.
Invasive plants with copious insect-pollinated flowers potentially alter the
nectar and pollen available to bees within invaded communities. The intro-

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125

duced invasive plant purple loosestrife (Lythrum salicaria) forms dense

stands in wetland fens in New Jersey, USA. Its flowers create an attractive,
long-lasting resource for bees and other pollinators. Purple loosestrife may
replace species that offer little resource for bees, resulting in more resources
for bees during its bloom. Invaded areas may offer fewer resources early
in the season before the loosestrife bloom if purple loosestrife crowds out
native flowering plants. I tested the hypothesis that bees are less abundant
and less diverse in invaded than uninvaded fen communities early in the
season, but more abundant in invaded sites coinciding with the loosestrife
bloom. I also hypothesized that large, social bees are more abundant in the
invaded areas during the loosestrife bloom because they have high food
requirements and can fly far. I sampled bee communities in six invaded
and six uninvaded calcareous fens in northwestern New Jersey. Pan traps
were set out in each site at three times over the summer: before the loosestrife bloom began (late May/early June), and twice during the loosestrife
bloom (July and August). Pan traps yielded more bees in May/June than
in July or August. Uninvaded sites tended to yield more bees especially in
the first sample, although these differences were not significantly. Richness
of bee genera was twice as high in uninvaded sites than invaded sites
(marginally significant) in May/June, but not in July or August. Extensive
pollinator observations of flowering plants revealed a higher proportion of
honey and bumble bee visitors to flowers in invaded sites than uninvaded
sites.
GOSLEE, SARAH C.,1,* BENJAMIN F. TRACY2 and MATT A. SANDERSON.1 1 USDA-ARS Pasture Systems and Watershed Management Research Unit, University Park, PA, USA; 2 University of Illinois at UrbanaChampaign, Urbana, IL, USA. Plant community structure of managed
grasslands at a range of spatial scales.
The composition and structure of managed grassland communities can have
a large impact on their productivity and sustainability. As part of a project
studying the role of diversity in grasslands, plant community composition
was surveyed at 88 sites between 1998 and 2002. A modified Whittaker
plot sampling design was used, allowing the analysis of community structure at different spatial scales. All potential pairwise relationships were
quantified using an asymmetric coefficient. A graph-theoretic framework
was employed to visualize the structure of these relationships at spatial
scales from 1m2 to 1000m2. At the finest scale, only 2% of the possible
pairwise relationships were significant, and 75% of those were negative.
At the coarsest scale, less than 1% of the possible relationships were significant, and 63% of those were negative. The coarse scale showed a lower
proportion of symmetric relationships. Analysis of indegree (association to)
and outdegree (association from) in the graphs of these relationships
showed a few key species of high degree. Associations with these species
were primarily negative at the fine scale, and positive at the coarse scale.
Relationships at one scale were poor predictors of relationships at other
scales. The coarse-scale patterns respond to environmental conditions and
management effects, while the fine-scale patterns are mainly determined
by local plant interactions. Development of management practices to affect
grassland community composition must take into account processes at both
scales. The graph-theoretic approach presented here provides an analytic
framework that allows the examination of complex community structure.
GOTELLI, NICHOLAS J.1,* and AARON M. ELLISON.2 1 University of
Vermont, Burlington, Vermont; 2 Harvard Forest, Petersham, Massachusetts. Tanking up: Drought, disturbance, and structure of Sarracenia
food webs.
Food webs in rainwater-filled pitchers of the carnivorous plant Sarracenia
purpurea are a model system for investigating how disturbance regulates
food web structure. This food web is supported by a base of insect prey,
and includes prey-shredding midge larvae; detritivorous protozoa and bacteria; omnivorous rotifers, mites, and mosquito larvae; and top-predator
sarcophagid larvae. In an ecological press experiment, we disturbed this
system by altering water quantity in Sarracenia leaves in a Vermont bog
and observing changes in food web structure. During a single field season,
one of five manipulations was applied semi-weekly to all leaves of each
experimental plant: 1) unmanipulated controls; 2) pitcher water and food
web removal; 3) food web removal; 4) pitcher water addition; 5) pitcher
water addition and food web removal. Conventional ANOVA revealed that

126

Abstracts

individual taxa responded idiosyncratically to water quantity, although

some taxa responded more to variation in leaf age and prey availability
than to experimental treatments. Path analysis was used to test simple colonization models based on water volume or prey resources, versus models
that incorporated known trophic relationships of the food web. Models
without trophic structure fit the data poorly. The best-fitting model was a
mixed trophic model that included both top-down and bottom-up effects.
Although habitat volume and basal prey resources have important effects,
trophic interactions account for more of the variation in food web structure
and abundance of individual taxa.
GOTTDENKER, NICOLE L.1,2,* 1 Southeastern Cooperative Wildlife Disease Study, The University of Georgia, nicoleg@vet.uga.edu, Athens, GA;
2
WCS Bolivia, Casilla 3-35181, La Paz, Bolivia. Resource availability
relative to White-lipped peccary (Tayassu pecari) populations in a Bolivian tropical forest landscape.
White-lipped peccaries (Tayassu pecari) are ecologically and socially important components of Neotropical forests. The objective of this study is
to understand patterns of resource availability in relation to white-lipped
peccary populations in a Bolivian Amazonian landscape. During peak wet
and dry seasons, fruit and seed patches were counted along fixed width
transects in five different forest types (sartenejal, Beni alluvial plain, two
geographically different tierra firme forests, and piedmont forest) within
Madidi National Park and the Tacana Tierra Comunitaria de Origen, La
Paz Province, Bolivia. Indices of local abundance and habitat use by whitelipped peccaries were estimated by observation of sign (i.e. peccary tracks)
and encounter rates with peccary herds. The overall number of available
fruit patches was greatest in tierra firme forests, intermediate in Beni alluvial plain and sartenejal forests, and relatively low in piedmont forest.
Preliminary results suggest a positive correlation between the number of
available ground fruit/seed patches and local white-lipped peccary abundance. Palms constituted a large proportion (0.53 to 0.98 palm patches/
total number of patches) of available fruit/seed patches in all forest types
during wet and dry seasons. Palm fruits and seeds also provided a majority
of available ground fruit/seed calories in all habitat types. Members of
Palmae and Moraceae were also primary resources in all forest types during the peak dry season. These results suggest that a better understanding
of the spatial distribution of lowland tropical forest palms at different scales
will allow for predictions of white-lipped peccary foraging behavior, densities, movements, and dynamics. This information is needed to develop
effective peccary conservation and management strategies.
GOVINDARAJULU, PURNIMA* and BRADLEY ANHOLT. University
of Victoria, Victoria, British Columbia, Canada. Factors influencing daily
survival rates of Pacific treefrog tadpoles (Hyla regilla) in natural
ponds.
Documented declines in amphibian populations have made it particularly
urgent to develop methods for accurately estimating their survival rates in
the wild, as well as identifying the factors influencing this rate. Numerous
theories and experimental studies have identified factors influencing survival of tadpoles but few have been assessed under field conditions. In this
study we obtained the first estimates of daily survival rates of tadpoles
under field conditions using capture-mark-recapture methods and quantified
the effect of abiotic and biotic factors on this survival rate. Specifically,
we assessed the effect of temperature, intra-specific density and the presence of introduced competitors (Rana catesbeiana), on the survival rate of
Pacific treefrog (Hyla regilla) tadpoles in Victoria, BC, Canada. We found
that daily survival rates of tadpoles were relatively constant within a season
and among years, but varied between ponds even if they were in close
proximity to each other. We found that temperature and intra-specific density interacted strongly to affect survival rates. At low densities survival
rates increased with temperature, but at high densities survival decreased
with increasing temperature. We were unable to detect an effect of the
introduced competitor over and above the variation accounted for by differences in temperature and intra-specific density in the ponds.
GOWE, AMY K.* and STEVE BREWER. University of Mississippi, University, Mississippi. The evolution of fire-stimulated flowering in the
genus Pityopsis (Asteraceae).
One hypothesized benefit of ecological restoration of fire-dependent ecosystems is increased reproductive success of fire-adapted species. Unequiv-

ocal evidence for the existence of fire-adapted species is lacking in many

systems however. The genus Pityopsis (Asteraceae) has been hypothesized
to contain at least one fire-adapted species, P. graminifolia. This species
dramatically increases flowering in response to lightning-season fires. In
this study, we documented the occurrence of fire-stimulated flowering in
all Pityopsis species and examined relationships between fire-stimulated
flowering, historic fire frequency, and the light environment for each species. In addition, we constructed a phylogeny of the genus based on published morphological data and then mapped fire-stimulated flowering on
the resulting topology. Floral induction and tree and groundcover canopy
coverage within each species9 preferred habitat were quantified for all but
one species (P. ruthii, an endangered species) at field sites in Massachusetts, Georgia, Florida, and Mississippi. Photographs and published data
were used to infer flowering patterns and the light environment for P.
ruthii. Results of our investigations support the following the hypotheses:
1) Fire-stimulated flowering occurred in one subgeneric taxonomic group
(sect. Graminifoliae) but not the other (sect. Pityopsis). 2) Fire-stimulated
flowering tended to occur in habitats or microsites that contained a dense
groundcover canopy during years without fire. 3) Fire-stimulated flowering
was more likely to occur in areas of relatively high pre-Colonial fire frequency in the United States. The phylogenetic analysis tentatively suggested that the two sections of Pityopsis were not distinct clades. Furthermore, the character used to distinguish sections in previous taxonomic treatments may, in part, be an artifact of describing a disproportionately large
number of non-reproductive plants in years without a lightning-season fire.
If our phylogenetic hypothesis is correct, then some species may have
gained or lost fire-stimulated flowering during diversification of the group.
GRABOWSKI, JONATHAN H.,* ERIKA J. CLESCERI, ADAM BAUKUS and PHIL O. YUND. University of Maine, Darling Marine Center,
Walpole, Maine, Walpole, Maine. Are we using herring to farm lobsters?
The effects of herring bait on Lobster growth and diet composition.
Lobster (Homarus americanus) landings in the past two decades have been
higher than traditionally thought to be sustainable in the Gulf of Maine.
Given the economic value of the lobster fishery in Maine and the socioeconomic consequences of a potential fishery collapse, determining the
importance of herring bait to lobster population dynamics is of critical
importance. To evaluate whether the annual subsidy of herring bait in the
nearshore waters of Maine are contributing to lobster production, we compared stomach contents and stable-isotope ratios in the tissues of lobsters
from closed (Monhegan Island) and open (Georges Islands) fishing areas
off the coast of Maine in the summer and fall of 2002. We also quantified
growth rates in both regions using mark-recapture experiments to assess if
the presence of herring bait in fished areas increases short-term growth of
lobsters. Diet composition of lobsters in fished areas consisted of substantially more herring bait than lobsters in areas closed to fishing. Larger sublegal lobsters (65-83 mm carapace width) in closed areas consumed more
of their natural prey (i.e., crabs, urchins and mollusks) than similar sized
lobsters in fished areas. Recaptured lobsters from fished areas grew substantially more than those in areas closed to fishing, indicating that in the
absence of herring bait, food may limit lobster growth in the Gulf of Maine.
In addition, these results suggest that herring bait influences the diet and
growth of lobsters as well as may impact benthic community structure in
the Gulf of Maine.
GRACE, JAMES B. USGS, Lafayette, LA. The utility of multivariate
hypothesis testing as illustrated through the study of plant diversity.
The thesis of this talk is that taking a multivariate approach to developing
and testing hypotheses can provide unique insights into the nature and
behavior of ecological systems. To illustrate this point, I present the results
of a sustained application of multivariate analyses to understanding the
regulation of plant diversity. These studies have used a variety of techniques, including structural equation modeling, iterative partial least
squares modeling, nonlinear signal strength modeling, and system simulation, often in combination. Examples of the discoveries made in these
studies are organized into (1) those that involved partitioning effects, (2)
those that involved detection of residual effects, (3) those that involved the
study of composite effects, (4) those that involved cross-site comparisons,
(5) those that involved comparing experimental results to multivariate ex-

pectations, and (6) those that involved partitioning reciprocal influences.

Specific discoveries of special note include (1) the value of light penetration
as a predictor of competitive effects, (2) the role that disturbance can play
in altering the competitive effect of a unit of biomass, (3) how a nonsignificant correlation between grazing and richness can hide offsetting positive and negative effects, and (4) how residual analysis can reveal the
continued impact of past disturbances. General discoveries of note include
(1) most current models of diversity regulation fail to incorporate the effects of nonresource stresses on richness, (2) nonresource stress effects
constitute the largest and most consistent cause of variations in species
richness in nature, and (3) field data reveals little evidence of positive
effects of richness on biomass production. I conclude that multivariate hypothesis testing can allow ecologists to develop more mature theories that
possess greater reality, superior operational meaning, a higher empirical
content, and a greater predictive capability than obtained with traditional
approaches.
GRAHAM, JOHN H.,1,* HOYT H. HUGHIE,1 SUSAN ROTH,1 KERRI
WRINN,1 ANTHONY J. KRZYSIK,2 JEFF J. DUDA,3 CARL FREEMAN4
and JOHN M. EMLEN.3 1 Berry College, Mount Berry, Georgia, USA;
2
Prescott College, Prescott, Arizona, USA; 3 Western Fisheries Research
Center, Seattle, Washington, USA; 4 Wayne State University, Detroit, Michigan, USA. Effects of habitat disturbance on diversity and abundance
of ants in the Southeastern Fall-Line Sandhills.
We examined the effects of habitat disturbance on diversity and abundance
of ants in the Fall Line Sandhills, at Fort Benning, Georgia. We collected
ants from highly, moderately, and lightly disturbed sites, using pitfall traps,
sweep nets, and by active searching. Disturbed areas were used for military
training; the canopy and soil was damaged by tracked and wheeled military
vehicles. Highly disturbed sites had fewer trees, more bare soil, a shallower
A-horizon, and soils that were highly compacted. In three years of sampling, we collected 109,690 ants (30 species in 5 subfamiles). Highly disturbed areas had fewer species than did moderately or lightly disturbed
areas. The disturbed ant communities were also less equitable, and were
dominated by active Dolichoderinae, especially Dorymyrmex pyramicus.
Prescribed burning at several of our sites had no detectable influence on
either species diversity or abundance.
GRAMLING, JOEL M.,1,* ROBERT K. PEET,1 JASON D. FRIDLEY1 and
THOMAS R. WENTWORTH.2 1 Department of Biology, University of
North Carolina, Chapel Hill, NC, USA; 2 Botany Department, North Carolina State University, Raleigh, NC, USA. Species richness and species
pool size variation across pH gradients in southern Blue Ridge forests.
Partel and Ewald have hypothesized that the relationship between local
species density and soil pH is determined by regional species pool size,
which in turn reflects the relative abundance of soil types during the evolutionary history of a flora. Ewald observed that calcareous sites in Central
Europe have higher species density and larger species pools than acidic
sites, and argues that this is the consequence of a Pleistocene bottleneck
for acidiphiles. The flora of the Southern Blue Ridge Mountains, USA has
always been associated with primarily acidic soils. Given the Partel-Ewald
hypothesis, one would expect the Blue Ridge region to contain a limited
pool of calciphiles. We used vegetation and soil data from 3328 100 m2
Blue Ridge forest plots to examine the generality of this hypothesis. The
Blue Ridge flora with less than 20% of species confined to sites above pH
4.7 contrasts dramatically with that of Europe. However, regional species
pool size increases with pH and related soil factors. Mean species density
is also strongly positively correlated with soil pH. Thus, both regional
species pool size and plot species density of Southern Blue Ridge forests
fail to match the Partel-Ewald predictions. The increase in species pool
with increase in pH and correlated soil factors exhibited by Southern Blue
Ridge forests appears to reflect broad species distributions and tolerance
for high pH conditions among species that grow predominantly under acid
conditions. We conclude that high species pool size and species density on
higher pH sites are consequences of generally more favorable conditions.
It may be correct that the abundance of high pH specialists in Central
Europe is an historical artifact, but our data contradict the assertion that
evolution of a flora in a primarily low pH environment will translate into
a negative correlation between pH and species density.

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127

GRANDY, A. STUART and G. PHILIP ROBERTSON. W.K. Kellogg Biological Station, Michigan State University, Hickory Corners, MI. Soil
carbon, nitrous oxide and aggregation changes immediately following
cultivation of an undisturbed soil profile.
Cultivation of previously untilled soils commonly results in the loss of 4060% of initial C and N from surface horizons. In the tropics losses typically
occur within months of cultivation, but a lack of data for temperate ecosystems prevents similar generalizations on a time scale less than about
five years. If the consequences of land-use intensification occur more rapidly than usually assumed, our understanding of agricultural expansion and
its impact on soil communities and processes may be altered. We investigated changes in soil C, aggregation, respiration, N2O production, and denitrification in the first sixty days following cultivation of a previously
untilled mid-successional community in southwest Michigan. Cultivation
had an immediate impact on all measured properties; within two weeks we
found differences in dissolved organic carbon, inorganic N, trace gas fluxes,
and field-moist aggregates due to plowing. Cultivation increased the average carbon dioxide flux during the sixty day study by over 50% and was
associated with a decline in water-extractable C. Large, water-stable aggregates rapidly declined following cultivation, likely providing increased carbon availability for respiration via aggregate destruction. Cultivation increased nitrous oxide fluxes from about 0.62 to 3.3 g N2O-N/ha/d. Differences in inorganic N pools appeared to drive these results: between 30-60
days after tillage, when differences in N2O fluxes were greatest, soil nitrate
concentrations averaged 0.18 micrograms NO3-N/g in the uncultivated sites
and 6.0 micrograms NO3-N/g in the cultivated sites. Cultivation also reduced denitrification enzymes likely due to rapid aggregate turnover, which
reduced the frequency and persistence of anaerobic microsites. The rapid
changes in nutrient mobilization observed here have significant implications for understanding the historical impact of agricultural expansion and
for mitigating that impact via soil C sequestration and other means.
GRANT, BRUCE W.,1,* CHARLENE DAVANZO,2 JASON TAYLOR3
and DAN UDOVIC.4 1 Department of Biology, Widener University, Chester, PA; 2 School of Natural Science, Hampshire College, Amherst, MA;
3
Ecological Society of America, Washington, DC; 4 Department of Biology, University of Oregon, Eugene, OR. Teaching Issues and Experiments
in Ecology (TIEE): Its foundations in cognitive and learning theory.
Teaching Issues and Experiments in Ecology (TIEE) is an NSF-funded
collaborative project to create a new peer-reviewed on-line and CD ROMbased publication of innovative ecological educational materials published
by the Ecological Society of America (www.ecoed.net/tiee). The principal
goal of TIEE, beyond that of materials dissemination, is to help ecology
faculty to improve their teaching and include more inquiry, active learning,
scientific thinking, collaborative group work, authentic assessment, formative evaluation, alignment, and other pedagogical principles and innovations - all of which are based upon a solid foundation of research and
theories about cognition and learning. This talk will explore this foundation
and demonstrate how the site design embodies (1) key features of constructivism, cognitive processing, and metacognitive activities such as reflection
and self-regulation of cognition, (2) the importance of building professional
learning communities, and (3) the elevation of the science of teaching as
scholarship. For example, a key idea in TIEE is that student activities (e.g.
describe and then analyze data with a peer) should reflect the cognitive
skills students should learn. Many "traditional" curricula present ideas like
this linearly. In contrast, when actively navigating the TIEE resources, faculty are guided to construct their own understanding of ideas such as the
link between student interaction and cognitive development. TIEE seeks to
catalyze a transformation among undergraduate ecology teachers from materials adopters to scholarly curriculum developers. We believe that the
ESA, through its digital library resources such as TIEE, is in an important
position to affect "top-down" catalysis of systemic reform among ecological educators. Further, we enjoin that research and theories in learning and
cognitive science must be central to the design of digital libraries and other
web resources for science faculty.
GRANT, NIKKI M.* and JAMES K. DETLING. Colorado State University, Ft. Collins, Colorado, USA. Effects of Gunnison9s prairie dogs on
vegetation structure and composition in a drought year.
Prairie dogs have been shown to alter their environment through burrowing,
grazing, and other activities, but most existing studies of prairie dogs con-

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Abstracts

centrate on the black-tailed species (Cynomys ludovicianus). We studied

the effects of Gunnison9s prairie dogs (C. gunnisoni) on plant species diversity, percent cover of plant species, and biomass of plant functional
groups (grasses, forbs, sub-shrubs). This study was done in two areas in
southwestern Colorado: Gunnison and Monte Vista. In Gunnison, litter,
Elymus elymoidies, Picradenia richardsonii, and Boechera drummondii
were more abundant off town than on (p,0.1). No differences were found
in the percent cover on and off town of the dominant species Muhlenbergia
filiculmis and Festuca samimontana at these sites. In Monte Vista, lichen
(Xanthoparmelia chlorochroa), and Bouteloua gracilis were more abundant
off town (p,0.1) and bare ground was more abundant on town (p,0.1).
When data from these two areas were combined more bare ground was
found on town and Chrysothamnus nauseosus was more abundant off town
(p,0.1). Contrary to many studies of other prairie dog species, few significant differences (p,0.1) were found in standing biomass on and off
town. Our results may have been influenced by severe drought during 2002
that limited the growth of many plants.
GRAY, JANET B. and THOMAS R. WENTWORTH.* XVIII Airborne
Corps and Fort Bragg, Fort Bragg, NC, USA. Rare vascular flora of the
longleaf pine-wiregrass ecosystem: Spatial structure of populations.
We examined the effects of patch isolation on extinction, colonization, and
persistence rates of rare vascular flora contained in the longleaf pine-wiregrass ecosystem. Two comprehensive surveys for rare flora were conducted
on Fort Bragg Military reservation in 1991-93 and 1998-99. Presence/absence data from the two surveys, comprising 1,192 observations, were used
to test the hypothesis that with increasing distance from its nearest neigbor,
the probability of persistence declines and the probability of extinction
increases. Analyses were conducted across species, among functional
groups, and at the species level. Across species, persistence of populations
declined and extinction increased with increasing distance from the nearest
neighbor. At the functional group level, monocots and clonal perennials
demonstrated isolation effects with respect to extinction and persistence.
For four of eight species examined individually, persistence and extinction
varied as functions of distance to nearest neighbor, indicating a metapopulation structure. This study shows the importance of spatial structure in
the persistence of rare flora in this landscape.
GREATHOUSE, EFFIE A.,1,* CATHERINE M. PRINGLE,1 WILLIAM H.
MCDOWELL2 and JEFF G. HOLMQUIST.3 1 Institute of Ecology, University of Georgia, Athens, GA; 2 Department of Natural Resources, University of New Hampshire, Durham, NH; 3 Sierra Nevada Aquatic Research
Lab, University of California, Santa Barbara, Mammoth Lakes, CA. Smallscale exclusion experiments indicate ecological effects of tropical migratory fauna extirpation.
While extinction and local extirpation of freshwater species is a global
phenomenon, ecological effects of species losses are poorly known. Previous research in Puerto Rico has documented that: (1) all native fishes
and freshwater shrimps have been extirpated from rivers upstream of large
dams without spillway discharge because these taxa are diadromous (i.e.
migrate between fresh and salt water); (2) shrimps reduce stream algal
biomass and deposition of fine particulate organic matter (FPOM). In this
study, we experimentally examined ecological effects of native fish and
shrimp extirpation above large dams. We used electricity to exclude macrofauna (fishes, shrimps and tadpoles .1.5 cm in length) from individual
boulders and small areas of bedrock in 2 streams of comparable size, watershed land use, habitat and slope. The first stream was above a large dam
that has caused the upstream extirpation of native fishes and freshwater
shrimps and has a high abundance of exotic non-migratory tadpoles and
poecilid fishes. The second stream had no large dam and was dominated
by native migratory fishes and shrimps. In the dammed stream, exotic tadpoles and fishes had no effect on chlorophyll a or FPOM. In the stream
with no large dam, native macrofauna reduced chlorophyll a by a factor
of 5 and FPOM by a factor of 11. These results indicate that large dams
have altered upstream biotic interactions between macrofauna, algae and
FPOM.

GREAVER, TARA,* LEONEL STERNBERG and ALEX DIAZ. University of Miami, Department of Biology, Coral Gables, Fl. Water uptake
and use efficiency of fore and back dune vegetation: integrating plant
physiology and hydrology.
Coastal beaches are transitional landscapes at the interface of land and sea.
As such, beaches are characterized by gradients of environmental factors
and zonal plant distributions that change with increasing distance from the
ocean. Zonal patterns of vegetation in coastal communities are often attributed to the quality and availability of underground water resources, however the relationship between the hydrologic conditions and vegetation of
beach dunes have not been closely examined, particularly in tropical and
subtropical beaches. This study investigated the interaction between vegetation and hydrology of subtropical beaches in South Florida and the Bahamas. Although environmental gradients are continuous, here we compare
the extremes of the gradient to use natural conditions as in situ experimental treatments. Thus, zones closest and furthest from the ocean were
demarcated as the fore dune and the back dune, respectively. Vadose soil
water conditions, plant water uptake, and plant water use efficiency (WUE)
were compared between dune positions. Results indicate that fore dune soil
waters were consistently more saline that those of the back dune at every
depth. Natural abundance of D and 18O, used as water tracers, identified
ocean water deposition in shallow soil waters of the fore dunes at both
sites and ocean water intrusion into the fresh water lens of the fore dune
at the Bahamian site. Mixing models calculated the proportion of environmental water sources in stem water, indicating that fore dune species are
likely to use some proportion of ocean water whereas those of back dune
do not. Thus their water harvesting strategies differ. Accordingly, d13C values of leaf tissues suggest fore dune species have higher WUE than those
from the back dune. Our findings suggest fore and back dune plant associations differ in their ability to utilize ocean water, therefore predicted sea
level rise will affect these species differently.
GREEN, DOUGLAS M.1,* and DOUGLAS E. JOHNSON.2 1 Arizona State
University East, Mesa, AZ, USA; 2 Oregon State University, Corvallis, OR,
USA. Long-term changes in species richness, diversity and eveness in
a northeastern Oregon riparian area.
Species richness and diversity of undisturbed sites have been suggested as
one of many metrics to evaluate the health or condition of managed ecosystems. We compared species richness, species diversity (H), and species
evenness (J) over a twenty year period from eight late season grazed and
ungrazed plant communities in a northeastern Oregon riparian zone. Communities measured were dry meadows, moist meadows, cheatgrass (Bromus
tectorum) Douglas hawthorn (Crataegus douglasii), ponderosa pine (Pinus
ponderosa), black cottonwood (Populus tricocarpa), thinleaf alder (Alnus
incana), gravel bars. Six stands of each community were measured with
thirty 25X25cm quadrats in late June or July of 1979, 1980, 1987, 1989,
and 1999. Mean species richness was greatest on gravel bars (33.0), moist
meadows (28.4), Douglas hawthorn (24.9), thinleaf alder (24.9), dry meadow (21.0), ponderosa pine (21.0, black cottonwood (18.9) and cheatgrass
(17.0). Gravel bars and moist meadows had the highest mean H (3.03 and
2.75 respectively) and mean species evenness (J) 0.8732 and 0.8640 respectively). Lowest mean diversity and evenness was found in black cottonwood (H 5 2.36, J 0.8229) and cheatgrass (H 2.22, J 0.8196). With
the exception of dry meadow communities no community had a consistent
grazing effect over the study period. This may be due to application of
sustainable stocking rates. Large year-to-year variation in species richness,
diversity, and evenness independent of the grazing treatment may be due
to study design, proper stocking rates, climatic variations or random error.
The large year-to-year variation observed raises questions about the usefulness of the selected indicators to evaluate ecosystem health in this riparian system.
GREEN, WALTON A.* and LEO J. HICKEY. Yale University, New Haven, CT. Using leaf architecture to compare fossil and modern forests:
Preliminary results from hierarchical cluster analysis and graphical
representation.
We have developed a methodology for numerically describing dicot forests
that allows direct comparison of fossil and modern stands. We use readily

observed characteristics of the form and venation of angiosperm leaves as

proxies for environment because comparable leaf-architectural data are easily obtainable from imperfectly preserved or insufficiently described fossil
and modern floras, and because a strong prima facie case has been made
for the presence of an ecological signal in leaf morphology. Descriptions
of forest stands based on both the two main published systems of leaf
architectural analysisClimate Leaf Analysis Multivariate Program
(CLAMP), and Compendium Index Categories (CICs)confirm expected
relationships among modern forests and allow semi-quantitative analogies
to be drawn between, for instance, the Puryear flora from the Lower Eocene
of Tennessee and a dry tropical forest currently growing in lowland Cuba.
We are working to expand this strategy into a standard methodology for
reporting paleoecological data on leaf litter assemblages that (1) provides
a readily visualized way of comparing forest ecosystems on a meso- or
macroscopic spatial scale, (2) allows us to track vegetational changes
through geological time, and (3) enables data from the leaf fossil record to
inform explanations of modern vegetation patterns like the similarity of
mixed mesophytic forests in Asia and North America.
GREENBERG, CATHRYN H.1,* and DOUGLAS J. LEVEY.2 1 USDA Forest Service, Southern Research Station, Bent Creek Experimental Forest,
Asheville, NC; 2 University of Florida, Gainesville, FL. Fruit production
in mature and harvested cove and upland hardwood forest of the
southern Appalachians.
Fleshy fruit is an important dietary component of many vertebrates, yet
little is known about fruit availability or management strategies. Since
1999, weve monitored fleshy fruit abundance monthly in 31 plots (0.1
ha)within cove hardwood forest (CHF)and upland hardwood forests
(UHF)and recently harvested areas of each(CH2 and UF2, respectively).
Thirty-nine species produced fruit during 1999-2002. Production ranged
570 (C56) to 1,629 (CHF) g (dry mass of pulp)/ha in 1999, and 796 (C53)
to13,644 (UF2) g/ha in 2002. Total fruit production increased during the
second year post-harvest, and remained highest in the harvested treatments.
Phytolacca fruit production peaked in CH2 and UH2 2 and 3 years postharvest. Rubus also increased only in harvested treatments 3 and 4 years
postharvest. Gaylussacia production was highest in both UHF and UH2
but varied among years. A few species dominated fruit production, but
dominant species varied among treatments. In 2002 Gaylussacia composed
67% of biomass in UHF, 31% in UH2, 20% in CHF, and 7% in CH2;
Rubus was absent from CHF and UHF but composed 45% (UHF) and 54%
(CH2) in harvested treatments. Fruit was available from June through December, but peak availability differed among treatments due to differences
in species composition and fruiting phenology. We conclude that fruit abundance and composition are influenced by forest type and age, and fruit
production varies in space and time.
GREENBERG, DAVID B.,1,* JOSEPH H. CONNELL1 and LLOYD
GOLDWASSER.1,2 1 Department of Ecology, Evolution, and Marine Biology, University of California, Santa Barbara, CA, 93106; 2 Department of
Demography, University of California, Berkeley, CA, 94720-2120. Direct
competition for space between coral colonies at Heron Island, Australia.
Competition for space on the sea floor, both within and between species,
is an important factor governing the abundance and species diversity of
reef corals. Corals may compete for space via overgrowth and overtopping,
or via physical and chemical attacks between neighboring colonies. We
have studied the extent to which competition via direct attack occurs between coral colonies at Heron Island, Great Barrier Reef, Australia, using
census data collected from fixed square meter plots at intervals since 1963.
Attacks between coral colonies occur along neighboring portions of their
perimeters, where each colonys polyps can physically reach each other.
Competition for space should then constrain growth along these closelyneighboring perimeters more than along other regions further from neighbors. We tested this hypothesis using maps drawn from color photographs
which show the perimeters of all colonies within each plot for each census
year. The maps allow us to track changes in size and position of individual
colonies as a function of their proximity to other colonies. We regard parts
of colony perimeters relatively far from any neighbors as controls for the
neighbored portions, but also analyze the extent to which this "neighbor

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129

effect" varies continuously, rather than discretely, along a colonys border

as a function of distance from its nearest neighbored region. For each
colony in each census year, the analysis involves: a) viewing its perimeter
as a series of evenly-spaced points, b) computing each points distance from
the nearest point on the perimeter of its nearest neighbor, and then c) measuring its growth at that point to the next census. Under competition,
growth at points on a colony border should vary as a continuously increasing function of each points distance (measured in the first census) from a
neighbor.
GREENLEE, ERIN S.,* WILLIAM L. GANNON and TERI J. ORR. University of New Mexico, Albuquerque, New Mexico, USA. Accumulation,
analysis, and web dissemination of bat call sequence files.
In the last 5 years, uses of the Anabat echolocation system to detect, archive, and analyze calls emitted by bats has increased dramatically. Recent
advances have allowed for the rapid and constant accumulation of bat call
files. Here we describe a system of filters and storage media for the rapid
and accurate analysis of call files. A call sequence for a selected bat can
be examined for species-specific characteristics such as slope, duration, and
frequency. These then can result in measurements that can be used to form
a species-specific filter. Filters can be constructed based on captured, then
hand-released bats so that species-specific values are used to construct a
filter. Those filtered files can then represent a particular species or sonic
group depending on the goals of the investigation. In order to handle large
quantities of call data, filters increase the speed and accuracy of call sorting
and organization. They are also extremely plastic and can allow filtering
for a variety of purposes. Once organized, call files are available on the
web stored in an acoustic library. The bat call library is archived and
available to any qualified researcher following the model of natural history
museum collection data.
GRIES, CORINNA,1,* DIANE HOPE,1 BROOKE L. STABLER,2 CHRIS
A. MARTIN2 and JOHN M. BRIGGS.2 1 Center for Environmental Studies,
Arizona State University, Tempe, AZ, USA; 2 Dept. of Plant Biology, Arizona State University, Tempe, AZ, USA. The manmade plant communities in a desert metropolitan area.
In a stratified tessellation random sampling design perennial plant abundance and richness was measured as percent cover of non-paved area within
30x30 m plots in the Phoenix, Arizona metropolitan area including surrounding natural desert. Of the 204 total plots that were investigated, 166
plots contained plant cover of a total of 188 unique genera. Average a
diversity was 8.4 and b diversity 23 with 95% empty cells and an average
total cover per plot of 35%. After eliminating rare taxa and appropriate
data standardization, a cluster analysis revealed a clear division between
mostly urban and mostly desert plots with only 25% information remaining.
Eight urban sites were classified in the desert cluster all of which had
original desert vegetation and 10 desert sites were classified as similar to
urban sites. Natural desert communities are characterized by dominance of
Larrea, Ambrosia or Encelia. Three of the urban sub-clusters were dominated by the genera Cynodon/Pinus/Morus, Prosopis, and Syagrus, respectively, while a fourth urban sub-cluster was highly diverse with no clear
indicator genus. The last two urban sub-clusters were examples of desert
landscape design while the first sub-cluster represents an oasis landscape
design type. The Prosopis dominated and the desert landscape sub-clusters
also contained highly disturbed desert sites that lacked the typical desert
genera.
GRIEVE, KATIE A.,1,* GREG A. BARRON-GAFFORD,1 EDWARD G.
BOBICH,1 RANDY L. PATTERSON,1 JOHN D. MARSHALL2 and RAMESH MURTHY.1 1 Biosphere 2 Center, Columbia University, Oracle, AZ,
USA; 2 University of Idaho, Dept. of Forest Resources, Moscow, ID, USA.
Canopy transpiration and conductance during increasing soil water
stress and cycles of VPD.
In order to separate the individual effects of atmospheric vapor pressure
deficit (VPD) and drought on leaf and stand level sap flux, conductance,
and transpiration of eastern cottonwoods (Populus deltoides Bartr.) growing
under ambient and elevated levels of CO2, cottonwoods were examined
under increasing soil water stress and cycles of high and low atmospheric

130

Abstracts

VPD. Rates of sap flux were measured on six trees within each of three
stands of fast-growing cottonwoods established in 1998 in the controlled,
sealed greenhouse system at the Biosphere 2 Center (Oracle, Arizona).
Atmospheric pressure, VPD, relative humidity, and temperature were measured continuously and averages were stored every 15 minutes in dataloggers. Stand level (canopy) transpiration was calculated using measures
of sapwood area index and sap flux. Stand-level conductance was calculated using a mass balance model for this particular closed system. Stand
level transpiration rates for plants growing under 400, 800, and 1200 ppm
CO2 averaged 0.562, 0.953 and 0.933 meters per day, respectively, under
low VPD (1.0 kPa) and high soil water content (0.3 v/v). Stand level transpiration rates for plants growing under 400, 800, and 1200 ppm CO2
averaged 0.518, 0.661, and 0.848 meters per day, respectively, under high
VPD (3.1 kPa) and high soil water content (0.3 v/v). After experiencing
four weeks of drought, volumetric soil water content had declined to 0.1
v/v, resulting in a 57, 69, and 43% decrease in stand level transpiration
under low VPD for the 400, 800, and 1200 ppm CO2 treatments, respectively. Under high VPD, stand level transpiration decreased by an average
of 63, 57, and 46%, respectively. Stand level conductance and leaf level
transpiration and conductance also decreased dramatically as soil water
content declined for trees under all CO2 treatments. Results at leaf and
stand levels will be compared and discussed.
GRIFFIN, CELINE A.* and JENNIFER S. THALER. Department of Botany, University of Toronto, Toronto, Ontario. Carnivore induced densityand trait- mediated indirect effects cause changes in plant resistance
to herbivores.
Indirect effects are interactions between two species that occur due to the
intervention of one or more species. These effects are determined both by
changes in density and traits of the intervening species, such as behaviour,
morphology and physiology. Such indirect effects are known to be important in many ecological systems. Their influence on plant resistance to
herbivores, however, has never been tested. I present results demonstrating
that the presence of the carnivore Podisus maculiventris caused changes
both in the density and behaviour of the herbivore Manduca sexta, and
these density- and trait-mediated indirect effects result in changes in resistance in Solanum ptychanthum. In the field, I tested the indirect effects of
carnivores on plants using five treatments: Control, Herbivore only, Lethal
carnivore and herbivore, Herbivore density control (herbivores were artificially removed at a rate to match removal on the Lethal treatment), and
Non-lethal carnivore and herbivore (carnivores had their beaks snipped).
In the field, M. sexta larvae feeding on S. ptychanthum in the presence of
lethal P. maculiventris caused 18% less damage relative to control plants
(Herbivore density control), whereas the presence of non-lethal carnivores
caused a 39% decrease in damage relative to controls (Herbivore only).
Consequently, both lethal and non-lethal carnivores indirectly caused significant decreases in the level of herbivore damage. This affected resistance
to subsequent herbivores feeding on the plant, as seen by M. sexta lab
performance bioassays. Furthermore, assays of several defensive proteins
revealed that carnivore presence indirectly suppressed plant resistance. In
the field, both lethal and non-lethal carnivores caused density changes in
herbivores, but lethal P. maculiventris caused much higher mortality than
did non-lethal carnivores. Behavioural observations in the lab showed that
the presence of a carnivore (either lethal or non-lethal) caused M. sexta to
feed less and spend more time in defensive postures. Overall, the results
from this experiment indicate that both density- and trait-mediated indirect
effects are important for plant resistance.
GRIFFIN, KEVIN L.,1,* NATALIE T. BOELMAN,1 MATTHEW H.
TURNBULL,2 DAVID T. TISSUE,3 SARA J. RICHARDSON,4 DUANE
A. PELTZER4 and DAVID WHITEHEAD.4 1 Lamont-Doherty Earth Observatory, Columbia University, Palisades, NY, USA; 2 Department of Plant
and Microbial Sciences, University of Canterbury, Christchurch, Canterbury, New Zealand; 3 Department of Biology, Texas Tech University, Lubbock, TX, USA; 4 Landcare Research, Lincoln, Canterbury, New Zealand.
Relationships between leaf physiological activity and spectral reflectance in vegetation along a soil chronosequence with a gradient in nutrient availability.
We compared spectral reflectance and physiological function in upper canopy leaves collected along a chronosequence with a gradient in nutrient

availability resulting from glacial activity in the Franz Josef system, New
Zealand. Specifically, leaf-level photosynthetic (Amax) and respiratory (Rd)
capacity, chlorophyll fluorescence and spectral reflectance of several species occurring at nine sites that vary from early colonizers to mature podocarp forest were measured. Spectral indices examined include the adjusted normalized difference vegetation index (aNDVI), the photochemical
reflectance index (PRI) and the water band index (WBI). We found that
across all sites, aNDVI, WBI, PRI and a combination index incorporating
both WBI and PRI were all correlated with Amax (r2 5 0.2 to 0.6). and that
two indices, aNDVI and PRI were correlated with Rd(r2 5 0.6 to 0.7).
Interestingly, there was no relationship between PRI and maximum electron
transport rate (ETRmax) across sites, but there were strong relationships
among species within all but two sites. In general, these results provide a
mechanistic basis for linking remotely sensed vegetation indices to photosynthetic and respiratory activity at increasing spatial scales. When
placed in the context of the chronosequence these data may provide a better
understanding of the processes regulating succession along gradients of
nutrient availability.

GRIFFIS, PAIGE A.* and JOEL C. TREXLER. Florida International University, Miami, FL. Experimental test of the effects of omnivory on
stable isotopic labels.

An organism9s trophic level can be identified by observing ratios of isotopic forms of carbon (13C/12C) and nitrogen (15N/14N) relative to other
components of a food web. Ratios of these isotopes in tissues of plants
and animals reflect those in their C and N sources, with a shift toward
dominance of the increasingly lighter form of N, but not C, with each
trophic level. Omnivores (animals feeding at multiple trophic levels either
serially during life or simultaneously) provide a special challenge for interpretation of isotope data. To explore the effect of omnivory on interpretation of isotopes, we raised neonate sailfin mollies (Poecilia latipinna)
over an 80-day period on diets with three isotopic values (a vegetable diet
d15N 5 2.8, a carnivore diet d15N 5 8.2, and a mixture of these two d15N
5 5.4). We also switched the diets after the first 40-days to test for the
persistence of diet shifts on tissue isotope signatures. We found that diet
affected growth rate, with herbivores growing the slowest and carnivores
the fastest. The isotopic signatures of fishes sacrificed after 40 days were
consistent with those of their diets (vegetable d15N 5 7.8, mixed d15N 5
10.1, animal d15N 5 12.0). After 40 days, we switched the diets and recorded the isotopic values at 80 days. Isotopic values of these fish were
strongly affected by the second diet, though a residual effect of the initial
diet was noted. These results indicate that simultaneous consumption of
foods from multiple trophic levels yields tissue isotopic signatures close to
a simple mean of the diet components weighted by their relative contribution. However, interpretation of serial diet shifts is more complicated,
though residual effects were greatly diminished after 40 days.

GRIFFIS-KYLE, KERRY L.* Syracuse University, Syracuse, NY, USA.

Hatching and early larval survival of amphibians in response to nitrate
enrichment.

There has been considerable concern related to how anthropogenic activities, such as pollution, affect organisms. Excess nitrogen entering the environment is causing changes in ecosystems worldwide, has been generating concern on a global scale, and is a possible culprit in the global
decline of amphibian populations. Research suggests that increases in the
concentration of nitrate in surface waters may alter the behavior and physiology of organisms and alter species interactions in aquatic communities.
We conducted field and laboratory toxicology studies on the effects of
agriculture and nitrogen enrichment on the hatching and early larval survival of several species of early breeding amphibians. We found that concentrations of nitrate in the environment during the early spring depress
the number of eggs that hatch and decrease larval survival. These results
have important implications for the long-term stability of amphibian populations in agricultural lands.

GRIFFITH, MICHAEL B.,1,* F. B. DANIEL,2 JAMES M. LAZORCHAK2

and MICHAEL TROYER.1 1 USEPA, ORD, National Center for Environmental Assessment, Cincinnati, OH, USA; 2 USEPA, ORD, National Exposure Research Laboratory, Cincinnati, OH, USA. Nutrients, canopy
cover, and grazers: Their effects on summer periphyton in small Midwestern streams.
Numerous studies in artificial streams suggest the relationship between nutrients and periphyton biomass (AFDM) and chlorophyll a in streams is
affected by ambient light, which is influenced by canopy cover, and by
grazer densities. To assess the relationships between nutrients and eutrophication-type effects in small streams, we created a model to describe
spatial and among year variations in periphyton collected during the summer of three years from 35 2nd-3rd order tributaries (Strahler order from
RF3 digital maps) of the Little Miami River, Ohio. These streams are characterized by high concentrations of dissolved P (tdP, 0.026-0.30 mg L21)
and nitrate-N (0.016-14 mg L21), agricultural land cover (row crop and
grassland: 50-95%) and variable riparian canopies (6.3-93%). The basin
also includes two ecoregions that differ in their relationships between land
cover and nutrient inputs to streams. Grazer densities during the three years
of study have ranged from 0.44-900 individuals m22. Regression analysis
showed that periphyton AFDM was positively related to tdP and inversely
related to canopy density (decreasing light). Periphyton chlorophyll a was
positively related to grazer density and nitrate-N, but relationship with nitrate-N differed between the ecoregions. Grazer density was positively correlated with nitrate-N (r50.35, p,0.001), suggesting that grazers suppress
the effects of nutrients and reduce the increase in periphyton accumulation
as nutrient concentrations increase.
GRIFFITH, TIMOTHY M.* and SONIA E. SULTAN. Wesleyan University, Middletown, CT. The effect of differing plasticity levels on population divergence in two Polygonum species.
Increasing attention is being given to the role of phenotypic plasticity in
evolutionary diversification. One possible consequence of adaptive plasticity is to reduce the extent of evolutionary divergence among populations
due to local selection. Alternatively, greater plasticity may broaden population divergence by allowing genotypes to spread to a wider range of
habitats in which subsequent selection can occur. To evaluate these possible
effects of plasticity on population diversification, we compared population
divergence within two closely related annual plant species that differ in
their degree of plasticity in response to light level, for traits of known
functional value. Polygonum persicaria and its less plastic congener, P.
hydropiper, both grow in a variety of disturbed habitats that range from
full sun to partial shade. We ask; does a species with greater plasticity for
traits associated with shade tolerance (P. persicaria) show different patterns
of population divergence in contrasting light habitats than a species with
less individual plasticity for these traits (P. hydropiper)? We grew inbred
lines from five P. persicaria and four P. hydropiper populations in high
light and simulated shade environments in the greenhouse, and measured
the degree of plastic response and the extent of population divergence within each species for functional and fitness traits. We discuss the results of
this experiment and their implications for the role of plasticity in evolutionary diversification and species ecological amplitude.
GRISWOLD, MARCUS W.1,* and L P. LOUNIBOS.2 1 University of Florida, Gainesville, FL; 2 University of Florida, Florida Medical Entomology
Laboratory, Vero Beach, FL. Functional responses and species coexistence in members of a treehole community.
Larvae of Toxorhynchites rutilus and Corethrella appendiculata are aquatic
invertebrate predators commonly found in treeholes and artificial containers
in the southeastern U.S. Both predators may co-occur. The functional responses of each predator to two prey species, Aedes albopictus and Ochlerotatus triseriatus, were estimated in laboratory experiments. Relative
prey vulnerability was also determined by offering the two prey species to
the predators in seven fixed ratios with a fixed total density of 100 prey
larvae. Both T. rutilus and C. appendiculata exhibited an inversely densitydependent Type II functional response to each prey species, taken alone.
This protocol provides a null model for examining the functional responses
on both prey, when encountered together. Analyses using Manlys a suggest

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131

that the full functional response may be more complex than predicted from
the null model. These results may help to explain the coexistence of the
native treehole species O. triseriatus with the invasive A. albopictus. Although A. albopictus escapes some predation by developing at a faster rate
than O. triseriatus, the greater vulnerability of A. albopictus to both predators in early instars counteracts the developmental advantage that A. albopictus has over O. triseriatus.
GRISE, DAVID J.* and AMANDA M. KENNEY. Department of Biology,
Southwest Texas State University, San Marcos, TX. Light-treatment-dependent phenotypes of Chenopodium album.
In an experiment to examine the role of nighttime water loss, we observed
two distinct forms of Chenopodium album, exemplified by a bimodal height
distribution. Although all plants were exposed to red/far red light (about
750-800 nm) during the overnight period, about half the plants displayed
a shorter bushy phenotype, similar to plants in their natural environment,
while the other half were taller and less branched, a typical shade response.
In a separate experiment designed to quantify differences between these
phenotypes, in which we did not expose plants to red/far red light, we
failed to observe these differences and plant height was normally distributed. We thus conclude that the expression of these two phenotypes is
dependent upon the presence of red/far red light, as might be expected if
it is related to shade response. Chenopodium album normally grows in high
light, non- shaded environments. However, the presence of two phenotypes
that occur in response to red/far red light may indicate genetic differences
that would impact fitness of plants growing in sub-optimal light conditions
including situations where plants are shaded by competitors. We propose
potential mechanisms and discuss the possible benefits of light-induced
phenotypes and ways to test these ideas.
GROFFMAN, PETER M.* and ANN M. DORSEY. Institute of Ecosystem
Studies, Box AB, Millbrook, NY, USA. Hyporheic denitrification in urban streams.
Hyporheic zones are areas where the water in a stream diverges from the
main channel into sediments below the stream, gravel bars next to the
stream or organic debris dams in the middle of the stream. Denitrification
is an anaerobic microbial process that converts nitrate, a cause of eutrophication in salt waters, into nitrogen gas. Hyporheic zones can be important sites for denitrification if they contain enough organic matter and support enough microbial activity (that consumes oxygen) to allow for anaerobic conditions to develop. We measured denitrification in hyporheic features in four streams in and around Baltimore, MD USA. Sediments from
pools, riffles, gravel bars (vegetated and non-vegetated) and debris dams
were sampled in streams in forested reference and suburban watersheds.
One stream (Mine Bank Run) is a highly degraded (extensive stream channel incision) stream that is undergoing restoration. Samples were taken
from both restored and degraded sections of this stream. Denitrification,
and a series of related variables (organic matter content, potential net mineralization and nitrification, respiration) were measured in a 10-day laboratory incubation of unamended sediments and denitrification potential
were measured in soil slurries amended with nitrate and carbon and incubated under anaerobic conditions. Denitrification potential was highest in
organic debris dams and organic-rich gravel bars - features with high organic matter content. Organic debris dams in suburban streams had higher
denitrification than debris dams in the forested reference stream, likely due
to the higher nitrate content of the suburban streams. This result suggests
that hyporheic denitrification increases in response to high nitrate levels
and that this process may be an important sink for nitrate in urban or
suburban streams. It is important to note however that high denitrifying
features (e.g. organic debris dams) are rare in urban streams with high
storm flows (e.g. Mine Bank Run).
GROGAN, WHITNEY N.,1,* ELIZABETH A. WALLS2 and JIM BERKSON.1 1 Department of Fisheries and Wildlife Science, Virginia Polytechnic
Institute and State Universtiy, Blacksburg, VA; 2 Center for Environmental
Studies, Virginia Commonwealth University, Richmond, VA. Movement
patterns and population dynamics of the American horseshoe crab,
Limulus polyphemus.
The American horseshoe crab is a multiple-use resource with both ecological and economic value. The horseshoe crab acts as an important food

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resource for shorebirds, bait source for eel and conch fisheries, and the
only source of Limulus Amoebocyte Lysate (LAL) for the biomedical industry. Horseshoe crabs require 10 years to reach sexual maturity, therefore
overexploitation of the population could have severe consequences. The
development of a management plan has been difficult due to lack of information regarding the status of the horseshoe crab population. We are
conducting a tagging study to investigate the movement range of the horseshoe crab. Preliminary results indicate that horseshoe crabs can move as
far as 195 miles over the course of 2 years. In another study, we are
examining the demography of the catch of horseshoe crabs by location.
Results show dramatic differences in age structure between locations. This
information, together with the existing literature, provides an increased understanding of the ecology of the horseshoe crab species. In addition, information gathered from these studies will be useful in the development or
modification of strategies to effectively manage the horseshoe crab population.
GROOTJANS, AB P. and ERWIN B. ADEMA. University of Groningen,
A.P.Grootjans@biol.rug.nl, Groningen, Groningen, The Netherlands. Sea
level rise: new opportunities for natural dune slacks in NW-Europe.
During the last decades, sea level rise has become a potential problem for
the preservation and maintenance of the Dutch coast. Sea level rise in North
Sea is estimated to be 20 cm/century, partly due to geological process and
partly due to an increment in global temperature. Other scenarios predict
a more extreme sea level rise, ranging from 60 cm/century to 85 cm/
century. Such changes will lead to increased erosion and coastal retreat
along large stretches of the NW-European coast. This calls for enforcement
of the coastal defence, but at the same time it may provide new opportunities for natural coastal wetlands in areas without human settlements. Until
recently most dune systems along the North Sea coast were fixed, thus
preventing the formation new dunes and dune slacks. The result was a
large loss in biodiversity during the last decades. We will report on experiments in which natural processes in the fore dune area have been promoted, by stopping the regular maintenance of sea defence structures (sand
dikes) or by creating small holes in the fore dunes to permit reflooding of
degraded dune slacks and beach plains. However, restoring a natural flooding regime does not automatically lead to the development of wetlands
with a high biodiversity. Knowledge on the effects of local hydrological
systems and willingness to promote renewed sand blowing in dune areas
are decisive factors for the creation of young dune slacks with a high
biodiversity.
GROSHOLZ, EDWIN D.,1,* LISA LEVIN,2 CARLOS NEIRA2 and A. C.
TYLER.1 1 University of California, Davis, Davis, CA, USA; 2 University
of California, San Diego, Scripps Institution of Oceanography, La Jolla,
CA, USA. Multi-trophic level and ecosystem changes following the
Spartina invasion of San Francisco Bay.
The 1975 introduction of Spartina alterniflora introduction into San Francisco Bay, CA (USA) and subsequent hybridization with the native S. foliosa produced a hybrid that has dramatically altered the historically unvegetated mudflats below the tidal distribution of native plants. The invasive hybrid reduces light (100X), reduces water flow (2X) and reduces
variation in sediment accretion (2-5X). In areas colonized by Spartina,
faunal densities have declined by .70%, diversity and richness are lower
and species composition has shifted significantly. For selected taxa, experimentally determined recruitment and growth rates are lower in Spartina
compared to unvegetated mudflat. However the abundances of several exotic molluscs are higher (2-10X) in the expanding margin of the hybrid
indicating it is facilitating some introduced species. Experimental sediment
transplants from mudflat into the hybrid and vegetation removal experiments provide support for the conclusion that the hybrid invasion is responsible for these changes. Changes in faunal abundances, increases in
total organic matter, silt/clay fraction, soil temperature, sulfide, ammonium
and chl a all occur rapidly post manipulation. At higher trophic levels,
grazing Canada geese are being affected by declining distributions of native
S. foliosa. Field data and exclosure experiments in the field show hybrid
Canada geese graze heavily and selectively on native S. foliosa during
winter nesting periods and ignore the hybrid Spartina. Feeding experiments
with captive geese support these observations. These results suggest that

hybrid Spartina is dramatically altering community structure, trophic transfer and ecosystem processes in this system.
GROSS, KATHERINE L.* and RICHARD G. SMITH. Michigan State
University, Kellogg Biological Station and Dept Plant Biology, Hickory
Corners, MI. Determinants and benefits of weed diversity in row-cropping systems.
Row crop agricultural systems in the USA typically are managed to minimize the biotic diversity within the field so that yield losses to competitors
(weeds), predators, pathogens and disease are reduced. A consequence of
this management is a reduction in field-level diversity that can potentially
provide ecosystem services to the crop. Concerns about the environmental
and economic costs of pesticides and increased interest in the potential
ecosystem services that can be provided by diverse communities has increased interest in the potential benefits that managing weed diversity within row-crops may provide. We have been monitoring the impacts of different agricultural management on weed communities over the past 15
years as part of the LTER project in agricultural ecology project at the
Kellogg Biological Station (KBS). We have shown that low chemical input
systems have more diverse and predictable weed communities than high
input systems. Cropping system diversity (through crop rotation and cover
crops) also influences the composition and abundance of the above and
below ground weed community (emergent and seed bank communities,
respectively). However, these differences in weed community composition
(and biomass) have little or no effect on crop yields. We recently initiated
an experiment in which crop rotation and cover crops are manipulated in
a way that will allow us to measure the direct effects of cropping system
diversity (continuous monocultures to 5 species mixtures) on the associated
weed community, crop yield and soil nutrients and microbial communities.
This experiment will provide us with a powerful tool for testing the potential role of biotic diversity in a cropping system on ecosystem function
and services in row-crop agricultural systems.
GROW, DAVID E. University of Arizona, Laboratory of Tree-Ring Research, Tucson, Arizona. Fractal modification of tree-ring chronologies
for streamflow reconstructions.
Long-term streamflow estimates are prerequisite to successful water management and stream restoration projects. Dendrochronological streamflow
reconstruction is a viable tool to estimate pre-gauged discharge, and fractal
analysis as applied to dendrochronology is a new approach to refine streamflow reconstructions. Six pinon (Pinus edulus) tree-ring chronologies were
developed within the Paria River watershed in southern Utah for the period
1702 to 1997. The fractal parameters (fractal dimension and Hurst exponent), calculated using the roughness-length method, describe the long-term
persistence of each tree-ring series and of the hydrologic record (19241998). The fractal dimensions range from 1.739 to 1.939 for the tree-ring
series for the calibration period (1924-1998), and from 1.884 to 1.946 for
the entire chronology lengths (1702-1997). The fractal dimension for the
annual hydrologic record is 1.802, and 1.819 for October 1 through May
31 discharge. Exponential smoothing of each tree-ring series based on the
ratios of the Hurst exponent of each series and the hydrologic record forced
the fractal dimensions of the tree-ring series to be closer to that of the
hydrologic series. Streamflow reconstructions using the fractally transformed chronologies resulted in less variance being explained than when
using the original series (Ra2 5 0.57 vs. 0.59 for October 1 to May 31).
The results suggest that the fractally modified reconstructions explain less
variance by removing noise between the series. Additionally, fractal analysis provides a complementary tool (with ARMA) to examine and model
persistence.
GRUNER, DAN S.* University of Hawaii at Manoa, dgruner@hawaii.edu,
Honolulu, HI, USA. Influences of resources and bird predation on species richness and diversity of Hawaiian arboreal arthropods.
Predator (top-down) and resource (bottom-up) influences in food webs are
strong and pervasive. Both resource enrichment and predation, in isolation,
are predicted to increase community diversity overall, but few studies have
investigated their interactive effects on diversity in real terrestrial ecosystems. This study focuses on arthropods associated with the dominant spe-

cies in young successional systems, Metrosideros polymorpha (Myrtaceae),

also the dominant tree in the Hawaiian Islands. In August and September
of 1998 on a 120-year-old flow, severe nutrient limitation was removed by
fertilization and combined with bird predator removal cages in a largescale, well-replicated, randomized block design. Arthropods were taken
from foliage clipping samples at the outset and conclusion of the 2.75 year
experiment. I sorted and enumerated all arthropods, well over 100 species
from 14 orders, to the species level. Previous reports documented increased
plant growth with fertilization and positive and interactive effects of treatments on arthropod densities. Fertilization also increased species richness
of arthropods, but diversity declined due to disproportionate increases in
dominant species. Bird exclusion increased species richness to an intermediate level, and strongly influenced species composition and densities
of several restricted groups, such as spiders. Ordination revealed distinct
compositional differences among treatments. Among trophic groups, parasitoids and predators represented the most species, but were generally
transient, while detritivores were the most abundant but species poor. Topdown effects in this system are interactive, not purely additive, with bottom-up influences. These results support the prediction that bottom-up constraints set the range of possibilities, while top-down forces determine the
realized community structure.
GUILLETTE, LOUIS J.,1,* CAREN HELBING2 and TAISEN IGUCHI.3
1
University of Florida, Gainesville, FL, USA; 2 University of Victoria, Victoria, BC, Canada; 3 National Institute of Basic Biology, Okasaki, Japan.
Aquatic Species in Ecosystems at Risk: Genomic Approaches for Assessing Normal and Abnormal Physiological Responses.
There has been a dramatic increase in the data available on the genomic
make-up of a number of species in the past decade. With the stimulus of
the human genome project, techniques and technology has provided great
advances in our ability to obtain genetic information from a wide range of
species, and there are extensive databases for members of all vertebrate
classes (e.g., fishes, amphibians, reptiles, birds and mammals), with the
exception of reptiles. Recent advances in our laboratories have begun to
fill the gap in this information for reptiles. Given the amazing improvement
in technologies for obtaining sequence information and the rapidly expanding technologies for their use, we have begun a research program that
focuses on the development of new genomics and proteomics information
and tools for a number of keystone species in subtropical wetland ecosystems. These tools, such as mircoarrays, genomic and proteomic databases,
and bioinformatic approaches for the appropriate use of these tools, will
be used in studies that examine organisms living in ecosystems at risk,
such as most aquatic systems in Florida and the southeastern USA. The
major focus of this project is to take these studies out of the laboratory
and into real world situation examining non traditional species, such as
alligators, mosquitofish, ranid frogs and turtles. The tools developed would
be of a design that would allow worldwide use and collaboration. We currently have functional microarrays for frogs and several small fish species.
Major questions currently being tested include studies of natural and induced metamorphosis in amphibians and natural sex determination and sex
reversal in fishes and reptiles.
GUNAWARDENE, ESHANI U.* and STEVEN A. JULIANO. Illinois
State University, Normal, IL, USA. A tale of two mosquitoes. The effect
of leaf type on competitive interaction of mosquitoes.
Previous experiments have investigated competition between Aedes aegypti
and the recently introduced Aedes albopictus. Often, when leaves are used
as a substrate for the microorganisms on which these Aedes feed, A. albopictus seems to be the better competitor. Although several single species
studies have investigated the effects of leaf type on population performance
of each species, there have been no experiments investigating whether different leaf substrates can alter the outcome of competition between A. aegypti and A. albopictus. We conducted such an experiment. Different leaf
types (scotch pine needles, live oak leaves, foxtail grass clippings) were
placed into water-filled containers with different combinations of A. aegypti+A. albopictus: 10+0; 20+0; 10+10; 20+20; 0+20; 0+10. For each
species in each container, we recorded median development time and mean
dry mass of males and females, and survivorship to adulthood. We incorporated these fitness components into the l index, which is commonly

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133

used as an estimate of finite rate of increase in such cohort studies. There

was no effect of treatment or leaf substrate on survivorship. Development
time for females was significantly affected by the treatment-leaf interaction,
with treatments producing significant differences for oak and pine, but not
for grass. Body mass of females of A. aegypti was significantly affected
by treatments, but body mass of females of A. albopictus was not, suggesting competitive asymmetry, with A. albopictus the superior competitor.
Thus, analysis of these fitness components individually results in heterogeneous conclusions about whether leaf type alters competitive interactions.
Analysis of the synthetic variable l resolves this heterogeneity, and suggests that leaf type does affect the outcome of competition. In order to
understand the interaction of these competitors, we must have further studies of how the interaction varies with important environmental variables.
GUNZBURGER, MARGARET S. Florida State University, gunz@bio.
fsu.edu, Tallahassee, FL. Nonadditive effects of multiple predators on
treefrog tadpoles.
Many studies demonstrate the importance of a single predator on a prey
species, but most prey are subject to predation by more than one predator
species in nature. Experiments evaluating the effects of multiple predators
on prey typically confound predator density with predator identity, thus
prohibiting an assessment of the additivity of predator species. The objective of this research was to implement an experimental design that controls
for total predation rate across treatments to determine the additivity of three
predator types (bass fish, aeshnid and libellulid odonate naiads) on green
treefrog tadpoles (Hyla cinerea). The predation rate of each predator species on the prey in isolation was measured. I then determined the density
of predators necessary in each treatment (all possible combination of the
three predator species) that would produce the same total predation rate if
all predator effects were additive. Each predator contributed equally to the
total predation rate in treatments with combined predators. I performed this
experiment in large mesocosms with a depth gradient and measured survival and habitat use of the tadpoles in response to the 8 different predator
treatments. Survival of tadpoles differed across treatments, indicating that
there are nonadditive effects of predators in this system. Tadpole survival
was highest in the control (no predator), aeshnid and libellulid combination,
and libellulid treatments. Comparison of observed survival of tadpoles to
that expected if all effects were additive indicated that tadpoles had higher
than expected survival in the aeshnid and libellulid combination treatment
and lower than expected survival in the libellulid and bass combination
treatment. Tadpoles did not alter their use of shallow or deep areas of the
mesocosm directionally in response to predator treatment. Predators interacted directly and indirectly to result in the nonadditive effects on tadpole
survival.
GUO, DALI,1 ROBERT J. MITCHELL1 and JOSEPH J. HENDRICKS.2
1
Joseph W. Jones Ecological Research Center, Newton, GA, USA; 2 State
University of West Georgia, Carrollton, GA, USA. Root branching order
and responses of fine roots to altered carbon source and sink strength.
Root branching order, a measure of root structure, can strongly influence
key root functional parameters such as root diameter, specific root length,
carbon (C) and nitrogen (N) concentrations, and root longevity. Yet it is
not clear how the flow of C and N is regulated by root branching order
when fine roots are under concurrent alterations in C source strength and
C sink strength. To answer this question, we manipulated C source strength
via canopy scorching (80% needle biomass reduction), and C sink strength
via N fertilization (50 kg N /ha/yr) in a longleaf pine ecosystem in the
southeastern USA. Our results showed that root starch concentration increased, whereas N concentration decreased with root order almost linearly
in the controlled plots. More importantly, our results demonstrated that root
branching order regulates the flow of C and N in the fine roots in a manner
contrasting with some of the previous predictions. Root starch concentration was significantly (p ,0.05) reduced by canopy scorching as compared
to the control, but only in root orders 3-5 (with the distal roots 5 the firstorder roots, and the fifth-order roots 5 the highest-order roots we sampled).
Root N concentration increased significantly (p ,0.05) with N fertilization
as compared to the control, but also only in root orders 3-5. These results
suggest that stored carbohydrates (e.g., starch) were preferably maintained
in the roots of the lowest-order (1-2), and that N concentration in these

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Abstracts

roots may not be elevated easily by N fertilization (rather, extra N was

exported to the higher-order roots). Because these lowest-order roots are
also the roots with greatest metabolic activity (i.e., highest N concentration
and root respiration) and may potentially consume the majority of the carbon transported to roots, the manner by which C and N are allocated in
these roots may be critical to understanding C and N cycles in forest ecosystems. The results of this study also offer new perspectives for modeling
root longevity based on a cost-benefit approach.
GUREVITCH, JESSICA,1,* TIMOTHY G. HOWARD1,2 and LAURA A.
HYATT.1,3 1 SUNY at Stony Brook, Stony Brook, NY; 2 New York Natural
Heritage Program, Albany, NY; 3 Rider University, Lawrenceville, NJ. Experimental tests of factors promoting forest invasibility.
The purpose of this research was to examine why some forest ecosystems
are vulnerable to biological invasions while others appear to be more resistant. Despite the threat that biological invasions pose for native forest
plants and animals, little previous experimental work has been carried out
in forests to test the importance of hypothesized causes of resistance or
susceptibility of forest communities to plant invasions. We report here on
the first of several field experiments testing the relative roles of specific
factors in determining community vulnerability to invasion by exotic
plants. We compared the responses of a suite of common native plants and
invasive exotics to experimental gaps and nitrogen addition in contrasting
sites: low diversity, low fertility pine barrens and more fertile hardwood
forests. Native and invasive species responded differently to the experimental treatments. Natives grew well in both pine barrens and hardwood
forests, but invasives grew well only in hardwood forests. Surprisingly,
native species grew more rapidly than invasive species overall. Invasive
species had a strong positive response to N in the pine barrens; adding N
to pine barrens soils was the only way that invasives were able to maintain
positive growth rates there. All species responded negatively to the addition
of N in the hardwood forests, probably due to fertilizer lowering soil pH.
Native species responded very little to the addition of N in the pine barrens.
Both native and invasive species had moderate postive responses to the
creation of gaps in the pine barrens. Invasive species were suppressed more
by the canopy in hardwood forests than in pine barrens, and responded
more strongly than did native species to the creation of gaps in the hardwood forest sites.
GURNEE, JULIE E.,* KURT REINHART and RAY CALLOWAY. University of Montana, Missoula, MT. An invasive tree suppresses natives
and facilitates conspecifics by altering the light in the understory.
Facilitation and ecosystem engineering by plants are important processes
that organize natural plant communities, yet the role of these processes in
plant invasions is less clear. Riparian communities invaded by Acer platanoides have understories with 3.5 times fewer plant species (m-2), 23
times less photosynthetically active radiation (PAR), and 57% less red to
far-red light (R:Fr) than natural riparian forests in Montana. Here we describe the effect of these PAR and R:Fr levels on the growth and survival
of seedlings for six native species and two nonnative species, including A.
platanoides. We hypothesized that changes in the understory light environment associated with A. platanoides canopies facilitate conspecifics and
negatively impact natives. We found that A. platanoides seedlings had high
survival (75-100%) in all light treatments. In contrast, 4 of 6 native species
experienced higher mortality at low PAR levels, similar to that beneath
Acer canopies. Low PAR levels reduced the biomass of all species; however the reductions in growth were more dramatic for many native species
than seedlings of A. platanoides. The light quantity beneath Acer trees had
a direct negative effect on the survival and growth of native species, which
corresponds with the general decline in species richness beneath Acer trees.
These results indicate that ecosystem engineering by an invasive tree alters
the light in the understory and negatively impacts native species while
indirectly promoting invasion of non-natives.
M, MIKAEL* and LARS-ANDERS HANSSON. Lund UniGYLLSTRO
versity, Lund, Skane, Sweden. Effects of diapause on zooplankton community dynamics: temporal patterns and the role of environmental
cues.
With the aim of assessing the impact of resting stages on zooplankton
community dynamics we examined production of, and emergence from,

diapausing eggs, together with planktonic community dynamics of three

cladoceran genera (Bosmina, Ceriodaphnia, and Daphnia) for 16 months
in a shallow lake. In a complementary field experiment, we manipulated
abiotic and biotic environmental variables, potentially important as triggering cues for emergence. Ceriodaphnia and Daphnia, but not Bosmina,
were dependent on emergence to initiate planktonic populations in spring.
During the rest of the season, emergence was unimportant for population
and community dynamics. Hatching of Ceriodaphnia and Daphnia occurred when the rate of increase of photoperiod and chlorophyll-a was
highest, but Daphnia hatched before the spring rise in temperature. The
timing of resting-egg production differed between genera, but always coincided with peaks in abundance. No hatching was found in any of the
experimental treatments, despite manipulation of several potentially important environmental variables. Differences in resting egg production were
found between some treatments however. The field study and experiment
together imply that the types of environmental cues involved in diapause
induction are different from the ones stimulating its termination. Abiotic
factors such as photoperiod may be important cues for the hatching of
resting eggs. However, the induction of resting egg production seems less
dependent on season and more on biotic factors such as crowding. This
conclusion is supported by a literature review on the cue used for diapause
induction and termination, which also indicated interesting difference in
cue utilization among taxa with different generation length.
HAAS, CAROLA A.,1,* SHANNON M. KNAPP,2 DOUGLAS N. HARPOLE3 and LORI A. WILLIAMS.1 1 cahaas@vt.edu, Virginia Tech, Blacksburg, VA; 2 Purdue University, West Lafayette, IN; 3 Virginia Cooperative
Extension, Warrenton, VA. Terrestrial salamanders in managed forest
systems: Early results from the Appalachian silviculture and biodiversity study.
In 1994 we began a long-term, stand-level, replicated experiment to address
the potential conflict between demands for increased production of timber
and non-timber forest products versus the desire to maintain native biodiversity and ecosystem processes in the eastern deciduous forest. As public
sentiment against clearcutting has grown, it is important to understand the
ecological and economic effects of alternative harvest and regeneration
methods. This study is designed to compare several forest management
alternatives and their effects on understory plant and animal communities,
processes, and projected economic value. We randomly assigned the following treatments to experimental stands: silvicultural clearcut, commercial
clearcut, leavetree, shelterwood, group selection, understory control using
herbicide, and an unmanaged control. Relative abundance of salamanders
on the control plots fluctuated somewhat annually, probably in response to
weather conditions. Using analysis of covariance with preharvest abundance as a covariate we detected a significant treatment effect on abundance
(p , 0.0001). All treatments with canopy removal had significantly fewer
salamanders than the unmanaged control and the herbicide treatments. Relative abundance of salamanders in plots harvested with alternative methods
were not significantly higher than in the silvicultural clearcut. Because
clearcutting disturbs a smaller area to obtain the same volume of wood,
clearcutting may have the lowest impact on salamander populations. We
examined two species and found that the percent of females gravid did not
differ between animals captured in uncut vs. cut treatments (57% vs. 55%
of Desmognathus ocrophaeus females, and 31 vs. 32% of Plethodon cinereus females were gravid). We discuss sampling issues, additional demographic data, and habitat characteristics in the context of identifying
species or populations at risk from varying management practices.
HAINES, BRUCE,1,* IVELISSE RUIZ-BERNARD,2 CHRIS PETERSON1
and DARRYL COLE-CHRISTENSEN.3 1 University of Georgia,
bhaines@plantbio.uga.edu, cpeterson@plantbio.uga.edu, Athens, GA.,
USA, USA; 2 University of Puerto Rico, iruiz@fs.fed.us, San Juan, PR.,
USA, USA; 3 Finca Loma Linda, fcallcb@sol.racsa.co.cr, Agua Buena,
Coto Brus, Costa Rica. Pre-montane forest succession reconstructed
from a chronosequence in Costa Rica.
Secondary succession was investigated to determine the time required to
naturally regenerate forest on abandoned agricultural lands near Agua Buena, Costa Rica (Lat 8.7 N, Long 82.9 W, elev. 1400 m). Trees of . 10 cm
diameter at breast height were inventoried in 7 conterminous patches of

forest 11, 12, 29, 38, 40, and 50 years of age and in old growth forest
(approximately 500 years of age). Preceding agricultural uses of these
patches were beans and corn, coffee, beans, pasture, pasture, unknown, and
corn and cassava respectively. Tree species richness values were 9, 2, 4,
17, 17, 20 and 22 respectively, while basal areas were 7, 6, 8, 20, 36, 39
and 30 m2/ha respectively. Densities were 500, 110, 300, 750, 410, 752
and 620 stems/ha. Tree species richness and basal areas differed little between the 50-year old and the old growth patches, suggesting very little
accumulation between these ages. Forest regeneration on isolated abandoned patches will likely be slower because of seed dispersal limitations.
These findings have important implications for landscape management.
HALE, BRACK W.* and ESTHER M. ALSUM. University of WisconsinMadison, Madison, WI, USA. Fifty years of change in the floodplain
forests of the lower Wisconsin River.
Recent studies of several Midwestern floodplain forests revealed these forests are shifting to Acer saccharinum dominated communities. This trend
is troubling, as the diversity of the overstory vegetation drives the diversity
of the forest ecosystem. In this study, we analyze the changes that have
occurred in the floodplain forests of one of the Midwest9s longest freeflowing stretches of river, the Lower Wisconsin, over the last 50 years and
investigate the impact of several anthropogenic factors on forest composition and structure: river regulation, overabundant herbivore populations,
and timber harvests. We resurveyed five locations surveyed in the 19509s,
as well as five other representative sites along the Lower Wisconsin. Preliminary analysis indicates the forests today have a lower diversity than 50
years ago (H9 51.38 and 1.54 respectively), although sites where timber
harvests had occurred showed increased diversity. At a species level, A.
saccharinum has greatly increased in abundance. Further, this trend appears
to be transitory, as A. saccharinum is almost completely absent from the
understory. Early-successional species and Ulmus americana have decreased in importance, while two relatively minor species in the 19509s
have gained importance, Celtis occidentalis and Carya cordiformis. The
understory has seen dramatic increases in two thorny shrubs, Rhamnus
cathartica and Zanthoxylum americana, indicating that species possessing
herbivore defenses may possess a strong competitive advantage in this
system. Overall, the results reveal the forests of the Lower Wisconsin are
at a later stage of succession than they were 50 years ago, an indication of
the reduced flooding regime of the river. Timber harvests appear to play a
role in maintaining some disturbance and encouraging the growth of certain
species. However, they do not completely replace the importance of floods
as the major form of disturbance.
HALITSCHKE, RAYKO.* Max Planck Institute of Chemical Ecology,
Jena, Germany. Ecological consequences of altered jasmonate signaling
capacity in Nicotiana attenuata.
The involvement of jasmonic acid (JA) and related compounds, collectively
called jasmonates, in the activation of wound- and herbivore-induced responses has been shown in several plant species. In the wild tobacco plant
Nicotiana attenuata the wound-induced accumulation of JA is specifically
amplified by fatty acid amino acid conjugates in the oral secretions of
the main natural herbivores of N. attenuata, Manduca sexta and Manduca
quinquemaculata. The activation of several direct and indirect defense responses in N. attenuata is mediated by the jasmonate signaling cascade.
We isolated genes involved in jasmonate biosynthesis and manipulated the
signaling capacity of N. attenuata by antisense expression of these biosynthetic genes. We used transformed N. attenuata plants as model ecological
expression system to investigate the signal function of jamonates in N.
attenuata9s defense response activation. Analysis of development of M.
sexta larvae feeding on transformed plants revealed reduced resistance in
plants impaired in JA biosynthesis which is mediated by reduced expression of direct (nicotine accumulation and protease inhibitor activity) and
indirect (terpenoid volatile emission) defense traits.
HALL, MYRA C.,1,* PETER STILING,2 DANIEL C. MOON2 and MARK
D. HUNTER.1 1 University of Georgia, Athens, GA; 2 University of South
Florida, Tampa, FL. Foliar quality and herbivory as predictors of litter
quality under elevated CO2 conditions in a scrub-oak community.
Atmospheric CO2 concentrations have increased over the last century and
continuing increases are expected to have significant effects on ecosystems.

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135

We studied interactions among atmospheric CO2, foliar quality, litter quality, and herbivores within a scrub oak community. Sixteen plots of opentop chambers were followed; eight of which were exposed to ambient levels
of CO2 (350 ppm), and eight of which were exposed to elevated levels of
CO2 (700 ppm). We focused on three oak species, Quercus geminata, Quercus myrtifolia, Quercus chapmanii, and one nitrogen fixing legume, Galactia elliottii. Our results thus far indicate that the links between the chemistry of green foliage and leaf litter are very weak. We also looked at six
insect groups on green foliage and found that damage decreased under
elevated CO2 conditions for four of the groups. In leaf litter we examined
undamaged, chewed and mined leaves. Undamaged litter was lower in nitrogen and higher in C:N ratios than was litter damaged by leaf chewers
and leaf miners. The overall increase in litter quality due to herbivore
damage and the decrease in herbivore damage under elevated CO2 treatments suggests that herbivory is a strong predictor of litter quality under
elevated CO2 conditions. Given the current rate of CO2 increase, it is important to determine general predictions for modeling ecosystem processes
under such environmental changes. We hope to be able to assess whether
herbivore-mediated changes in litter quality under elevated CO2 influence
essential ecosystem processes.
HALL, SHARON J.* The Colorado College, Colorado Springs, CO. Factors controlling nitrification in nitrogen-fertilized tropical forest soils.
Atmospheric N deposition has been shown to alter biogeochemical processes in terrestrial ecosystems. Rates of nitrification in soil are known to
increase following long-term N additions in many temperate systems, but
little is known about the response of soil microbial communities and nitrification in tropical forests. I measured laboratory rates of nitrification (potential and net/gross nitrification) in soils collected from N-fertilized, Nlimited and P-limited tropical forests in the Hawaiian Islands. Nitrification
was significantly higher from soils in the P-limited forest compared to the
N-limited forest, and long-term N fertilization increased nitrification in soils
from both sites. Although soil pH at both sites is naturally low in the field
(pH 3-4), potential nitrification (shaken-slurry) was significantly inhibited
at pH 4 compared to pH 7. Heterotrophic nitrification (measured w/acetylene) comprised approximately 25% of total nitrification at both sites (and
in N-fertilized plots) when measured at pH 4 (close to field pH), but contributed less than 10% of nitrate produced at pH 7. Two-month laboratory
incubations of soil with a range of N-additions (dose response w/ equivalent
of 0-125 kg N/ha) did not significantly increase rates of nitrification in
control or long-term N fertilized plots in the N-limited site. Based on prior
dose-response experiments in the field, it is expected that nitrification in
soils from the P-limited site (both control and N-fertilized) will show high
sensitivity to increasing N additions. Preliminary molecular analyses of
microbial community structure failed to locate nitrifying microorganisms
in either the N-limited or P-limited forest soil using known DNA primers
specific to nitrifiers and their enzymes.
HALL, SONIA A.,1,* INGRID C. BURKE,1 DAVID O. BOX,2 MERRILL
R. KAUFMANN3 and JASON M. STOKER.4 1 Colorado State University,
Fort Collins, CO; 2 3Di Technology Inc., Eagle Scan Remote Sensing,
Boulder, CO; 3 Rocky Mountain Research Station, U.S. Forest Service, Fort
Collins, CO; 4 EROS Data Center, U.S. Geologic Survey, Sioux Falls, SD.
Estimating stand structural attributes of ponderosa pine forests with
discrete return lidar.
Stand and biomass structure are dynamic attributes of forests. Reliable
spatially explicit estimates of these characteristics are needed periodically
for many objectives, such as ecosystem modeling, carbon budget estimation, timber production, biodiversity studies and habitat mapping. Of growing importance is the estimation of stand-level (i.e. per hectare) live and
dead fuels, to determine fire hazard and for fire behavior modeling. Optical
remote sensing has provided estimates of stand attributes, but saturating
relationships limit the usefulness of two-dimensional data. Continuous return lidar has been used to estimate three-dimensional stand structural variables. We used discrete, multiple return lidar data to estimate stand structural variables in ponderosa pine forests in Colorado. We used an information-theoretic approach to select the best simple models to estimate
mean and maximum heights, tree biomass and its components, tree and
biomass densities, basal area and bole volume. Most predicted vs. observed

136

Abstracts

values were highly correlated, with r2 ranging from 30% to 86%. The mean
height estimates had the lowest r2, but it increased substantially when measured tree heights were weighted by basal area (r271%). We combined
the model selection techniques with validation with new data and a comparison of regressions at two spatial scales to select a subset of the 45
initial metrics calculated from the lidar data. This subset is sufficient to
predict forest and biomass characteristics with reasonable accuracy. Fusion
of lidar and Landsat data will help provide periodic, spatially explicit estimates of these variables at low cost/ha, providing necessary inputs for
fire behavior models. Modeling efforts will provide insight needed to determine the critical scale or scales at which fuels influence fire behavior.
HALL, SPENCER R.* University of Chicago, Chicago, IL. Competition
between grazers along light:nutrient gradients.
Ecologists remain challenged to mechanistically explain changes in species
composition along environmental gradients. Recent developments in stoichiometric theory suggest that elemental body composition of grazers may
be a key trait explaining responses of grazer community composition to
changes in elemental food quality (driven by the relative supply of light
and nutrients to ecosystems). In a semi-natural pond mesocosm experiment,
the phosphorus (P)-rich, planktonic grazer Daphnia responded as predicted
by verbal stoichiometric theory: it dominated grazer communities in environments with P-rich food (at high nutrients), and was virtually absent
in environments with P-poor food (at low nutrients). Other P-rich species,
however, did not respond similarly to the light:nutrient-driven gradient in
food quality. In order to understand this result, I considered theoretical
competition between two grazers for a single plant resource using analysis
of nullclines. The plant varies in P-content, and its production depends
upon nutrients and light. Grazers can be carbon- or P-limited, and feed
according to linear functional responses. When elemental food quality is
good for both grazers, the competitive dominant at equilibrium maintains
zero growth rate at a lower level of plant carbon density, N* (a familiar
result of resource competition theory). When food quality is bad for both
grazers, the winner of competition maintains zero growth rate at a lower
level of plant nutrient density, QN* (a new result). This QN* rule can
explain response of Daphnia in the experiment, but it also reveals why
other P-rich grazers can respond oppositely: grazer body composition is
only one of several grazer traits comprising QN*. Therefore, this theory
emphasizes that body composition should not be considered separately
from a grazers trait suite. Furthermore, a N*-QN* tradeoff among grazers
can permit species coexistence at intermediate nutrient supply, but it also
determines shifts in grazer composition along nutrient and light gradients.
HALVERSON, MARK ANDERS.* Yale University, New Haven, CT. Examining the evolutionary ecology of wood frogs (Rana sylvatica) using
microsatellite analysis.
Amphibians have commonly been used as a model system in ecology and
evolution. However, because of the difficulty of assessing relatedness between individuals and tracking individuals over time, most of these studies
have been conducted at the population level or in artificial environments.
I used multilocus microsatellite analysis to uniquely identify wood frogs
(Rana sylvatica) and their offspring in the wild and thereby address several
questions about the ecology and evolution of this species at the individual
level. I found that the weight of the males and females was significantly
related to breeding success, but not always to the number of metamorphs
produced. There was no evidence for multiple paternity in any egg mass.
There was a significant difference in the weight of the metamorphs and the
larval period of different sibships but this was not significantly related to
the weight of the parents. We also found evidence that the larvae of different sibships were nonrandomly distributed in the ponds. Ongoing study
of this system using these techniques will make it possible to address similar questions at longer time scales.
HALVORSON, JONATHAN J.,* DAVID P. BELESKY and HARRY W.
GODWIN. Appalachian Farming Systems Research Center, USDA-ARS,
Beaver, WV. Inhibition of seed germination and early development by
leaf litter extracts.
Silvopastoral management strategies in hill-land Appalachia seek to expand
spatial and temporal boundaries of forage production and promote ecosys-

tem integrity through a combination of tree thinning and understory pastures. Little is known about forage establishment in the understory, which
may be impacted by competition with trees for light or water, or by the
physical and chemical effects of deciduous tree litter. To gain understanding
of forage establishment, we determined the effects of water extracts of leaf
litter from common Appalachian tree species; yellow poplar, Liriodendron
tulipifera, red maple, Acer rubrum, white oak, Quercus alba, and eastern
white pine, Pinus strobus, on germination and seedling growth of forage
species. Leaf extracts added to soil inhibited seedling root elongation of
white clover, Trifolium repens (cv Huia), with greatest inhibition caused
by Liriodendron and Acer, species with highest rates of decomposition.
Conversely, extracts from tree species with relatively slow decomposition
rates, Pinus and Quercus, had little impact on seedling root elongation. The
effects of Liriodendron leaf extract persisted and decreased dry mass of
white clover after a 4-week growth period relative to other extracts. Seedling growth might be diminished by immobilization of soil nutrients by
microorganisms, stimulated by litter extracts, or by allelopathy. Liriodendron litter extract delayed seed germination for several forage species, especially legumes, but germination resumed immediately after washing the
extract from the seeds. Water soluble compounds that inhibit seed germination and impair seedling growth may improve overall seedling success
by preventing premature growth early in the season but could compromise
forage establishment as an understory crop. Management practices that account for litter impacts are needed to ensure successful sward establishment.
HAMERLYNCK, ERIK P.1 and TRAVIS E. HUXMAN.2 1 Department of
Biological Sciences, Rutgers University, Newark, NJ, USA; 2 Department
of Ecology and Evolutionary Biology, University of Arizona, Tucson, AZ,
USA. Topography/canopy effects on the photochemical performance
of jojoba, a broad-leaved Sonoran Desert evergreen shrub.
We used PSII chlorophyll fluorescence to examine the varying effects of
leaf canopy position and plant topographic exposure on photochemical dynamics at the height of early-summer pre-monsoonal drought and high
temperature stress in jojoba (Simmondsia chinensis), an unusual Sonoran
Desert shrub that possesses an extensive broad-leaved evergreen canopy.
Predawn water potentials were lowest in plants at ridgetop and eastern
exposures (-9.5 and -9.1 MPa, respectively), and highest in western exposure plants (-8.0 MPa). Mean predawn levels of optimal PSII photochemical yield (Fv/Fm) were 0.63, 0.54 and 0.74 for ridge, eastern- and
western-exposure plants, respectively, with minor differences between inner- and outer-canopy leaves. During the day, canopy-position effects on
Fv/Fm and NPQ, an index of engagement of thermal-dissipative photoprotection, were topographic-specific. In general, Fv/Fm and NPQ were higher
in shaded inner-canopy leaves, while light-adapted PSII yield (FPSII)was
lower compared to outer-canopy leaves. However, in eastern-exposure
plants Fv/Fm increased and NPQ remained constant in inner-canopy leaves,
while Fv/Fm and NPQ in outer-canopy leaves declined through the day.
This latter pattern was apparent at higher parameter levels in outer- and
inner-canopy leaves at ridgetop and western-exposure locations. These findings show that the dense canopy of jojoba might serve to ameliorate exposure to high early-morning photon flux, allowing for sustained photosynthetic activity deeper in the canopy during limiting soil-water conditions, thereby facilitating jojobas persistence across topographically complex desert landforms.
HAMILTON, E WILLIAM1,* and SCOTT A. HECKATHORN.2 1 Washington and Lee University, Department of Biology, Lexington, VA; 2 Syracuse University, Department of Biology, Syracuse, NY. Stress reponses
of Chenopodium album and Amaranthus retroflexus to heat stress
grown in elevated CO2 and temperature.
The predicted increase in global carbon dioxide concentrations and temperatures with associated increased frequency of temperature extremes will
have a significant impact on plant populations. This experiment quantified
the responses of Chenopodium album (C3) and Amaranthus retroflexus (C4)
to elevated CO2 and temperature and chronic heat stress (HS). The experiment consisted of 2-CO2 concentrations (350 and 700 ppm) x 2 growth
temperatures (16:24 and 22:30 8C night:day) x 2 temp treatments (Control
and HS at 428C after one month of growth in treatments). We quantified

net photosynthesis and a set of stress responses related to heat and oxidative
stress tolerance. The stress responses of interest were small heat-shock
proteins, ascorbate (Vitamin C), Cu/Zn superoxide dismutase and catalase.
Photosynthesis was stimulated by elevated CO2 in Chenopodium at low
growth temperatures but not at elevated growth temperatures and CO2 and
there were no significant effects in Amaranthus. Production of stress responses increased in Chenopodium grown in 350 ppm CO2 when grown at
low temperatures and heat stressed, but Chenopodium grown at elevated
temperatures and elevated CO2 had reduced production. Amaranthus had
increased production of stress proteins in both growth temperature treatments when heat stresses and production was highest at 700 ppm CO2. The
results suggest that the interaction of elevated CO2 and temperatures and
temperature extremes may negatively impact Chenopodium populations
and potentially other C3 herbaceous perennials and that Amaranthus will
not be negatively effected and potentially other C4 species.
HAMILTON, JASON G.,1,* ORLA DERMODY,2 MIHAI ALDEA,1 ARTHUR R. ZANGERL,2 MAY R. BERENBAUM,2 ALISTAIR ROGERS3
and EVAN H. DELUCIA.2 1 Ithaca College, Ithaca, NY jhamilton@
ithaca.edu; 2 University of Illinois at Urbana-Champaign, Urbana, IL;
3
Brookhaven National Laboratory, Upton, NY. Elevated atmospheric
carbon dioxide affects crop herbivory by insects.
Human activity is rapidly altering the chemistry of the troposphere in ways
that may have important effects on chemical composition of leaves. These
changes in leaf chemistry will potentially affect the interactions between
plants and insect herbivores. At the University of Illinois SoyFACE (FreeAir Concentration Enrichment) facility, experimental plots of soybean (Glycine max) were exposed to ambient air, elevated CO2 (550 ppm), or elevated
ozone (1.2 X ambient). In July and August 2002, we measured levels of
insect herbivory, insect populations and changes in leaf chemistry. In July,
elevated CO2 more than doubled the amount of leaf tissue consumed by
herbivores (ambient losses 4.5%; elevated losses 10.6%), whereas elevated
ozone had no measurable effect (5.9%). In August, overall levels of herbivory were much lower with no significant difference between ambient
and elevated CO2 plots. In July, the main insect causing this damage was
the introduced Japanese beetle (Popillia japonica), with significantly larger
populations in the elevated CO2 plots compared to control. Measurements
of leaf tissue chemical composition showed large increases in three known
feeding stimulants of Japanese beetle (sucrose, glucose and fructose). Feeding trials confirmed that Japanese beetles consume more tissue from soybean leaves grown under elevated CO2 conditions. These results suggest
that changes in tropospheric chemistry caused by human activities may
increase crop damage by insect herbivores in the future.
HAMMAN, SARAH T.* and INGRID C. BURKE. Colorado State University, Ft. Collins, CO. Prescribed fire effects on the soil biogeochemistry of mixed-conifer forests in Sequoia National Park, California.
Prescribed burning of forests may have significant effects on water and
nutrient availability, and understanding these will aid in determining the
most efficient and practical restoration activities. We collected data on soil
moisture, N mineralization and soil respiration during the summer of 2002
in control and burned sites of an old-growth mixed-conifer forest of Sequoia National Park, California. We measured nitrogen mineralization using
the soil core incubation method and the resin strip method, and measured
soil respiration using a PP Systems soil respirometer. Soil moisture decreased by 2.9% with prescribed burning due to the increased soil temperature and possible formation of a water-repellant layer. Soil respiration rate
decreased by 0.44g CO2/m2/hr, most likely from the lack of autrotrophic
contribution to total soil respiration. There was a significant increase in
nitrogen mineralization rate with prescribed burning (0.83 mg N/resin strip/
day) that was likely due to a shift in microbial activity from an increase
in labile SOM (carbon source), an increase in soil temperature and the
decreased plant nutrient uptake. All results were significant at p,0.0001.
The information gathered from this project will be used, along with data
collected for the Fire and Fire-Surrogate Study, to determine overall effects
of prescribed burning on the physical and chemical properties of soils in
the mixed-conifer zone of the Sierra Nevada.

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137

HAMPTON, STEPHANIE E.* and DANIEL E. SCHINDLER. University

of Washington, Department of Zoology, Seattle, WA. Relative effects of
climate and biotic interactions on plankton dynamics.
Aquatic ecosystems throughout the world are subject to changing climate,
human alterations of food webs and nutrient cycles, and introductions of
exotic species. Understanding the ecological effects of these human perturbations is somewhat limited because they are usually confounded in
time. The probability that population dynamics are affected through both
direct and indirect pathways further complicates the prediction of response
to anthropogenic disturbance. Development of new multivariate autoregressive models (MAR) provides a new set of tools for understanding complex ecological dynamics in time series data. The long-term limnological
data set from Lake Washington provides the opportunity to study plankton
community responses to several anthropogenic disturbances as well as
long-term climatic changes. Over the last four decades, lake temperature
has clearly increased, nutrient loading has dramatically decreased, and the
food web has been altered. To discern the effects of multiple factors on
multiple interacting species, we used MARs to determine the relative importance of abiotic drivers and biotic relationships in altering species abundance. Nutrient loading reduction directly altered phytoplankton characteristics by reducing total abundance of cyanobacteria. At the same time,
anthropogenic alteration of the fish community reduced predation on zooplankton grazers by Neomysis, a voracious invertebrate predator. Together
these abiotic and biotic perturbations overrode climatic effects in shifting
dominance of the grazer community to Daphnia, which produced further
changes in phytoplankton characteristics as well as phenological changes
for competing grazers such as Diaphanosoma, Bosmina, and Conochilus.
HAMRICK, JAMES L.* University of Georgia, Athens, Georgia, U.S.A.
Trees and herbaceous plants: genetic implications for the conservation
of old-growth communities.
One of the goals of any conservation effort is to maintain native levels of
genetic diversity. To accomplish this goal, detailed information is usually
needed concerning the distribution of genetic diversity within and among
populations of the target species. But strategies for the conservation of
genetic diversity may differ among species within a plant community with
different life history traits (i.e. breeding systems, seed dispersal mechanisms, etc.) and spatial distributions. For example, most dominant forest
tree species maintain the majority of their genetic diversity within their
populations while many herbaceous species growing beneath their canopy
may have a much higher proportion of their genetic diversity among their
populations. These divergent patterns of genetic diversity are magnified if
the tree species has a continuous distribution while the herbaceous species
have disjunct distributions. How then can efficient strategies of genetic
conservation be developed for the community? A further complication is
that both theory and empirical data demonstrate that plant species with
higher levels of genetic differentiation among populations also have more
population-to-population variation in the amount of diversity contained
within their populations, increasing the difficulty to design effective conservation plans. Case studies of southeastern plant species will be presented
to illustrate these points. Suggestions regarding the decision-making process and the information needed to develop efficient and effective conservation strategies will be discussed.
HANAN, NIALL P. and MAHESH SANKARAN. Colorado State University, Fort Collins, CO, USA. Structure and function in African Savannas: a continental scale analysis of patterns.
Many explanations for the persistence of tree-grass mixtures in savannas
have been advanced thus far. In general these concentrate on equilibrium
mechanisms, where niche separation and competition for water lead to
coexistence, or non-equilibrium mechanisms, where coexistence is promoted by disturbances related to rainfall patterns, fire, herbivory, or other
forms of spatial and temporal variability. All of these explanations and
processes may be active to different degrees in the diverse savannas of the
World where rainfall, fire frequency, herbivory and human management
vary, and a comprehensive conceptual or numerical model that explains
both coexistence and the relative productivity and cover of the tree and
grass components is yet to emerge. The savannas of East, West-Central

138

Abstracts

and Southern Africa share many characteristics with each other and with
other savanna regions of the world, but are also distinct in their combinations of soil, climate, herbivory and resulting savanna dynamics. Here
we report on analysis of data from a large number of savanna research
sites in Africa, investigate the range of structural states that are realized
within the potential multi-dimensional environmental space, explore patterns in savanna characteristics that emerge from complex interactions and
feedbacks between interacting components, and discuss the underlying processes that contribute to these patterns.
HANCOCK, THOMAS E.* and WILLIAM K. SMITH. Wake Forest University, Winston-Salem, NC. Ecophysiology and life history effects on
plant survival in barrier beach sand dunes.
Barrier beach sand dunes are harsh, highly dynamic environments that are
physically controlled. Daily abiotic factors plus severe episodic salt-water
overwash during storm events may require specific characteristics of plant
phenology and ecophysiology for species survival. The present study is
evaluating the importance of ecophysiological characteristics influencing
carbon gain versus life history and phenology characteristics in four species
inhabiting the foredune, primary dune, and swale on Topsail Island, North
Carolina (USA). Amaranthus pumilus and Cakile edentula typically grow
on the foredune while Iva imbricata grows from the foredune into the
primary dune. Hydrocotyle bonariensis is occasionally found on the foredune but most often grows on the primary dune and swale. Permanent
quadrats were established and monitored at monthly intervals for number
of individuals, leaf stage, flowering and fruiting. Annual carbon gain was
estimated based upon daily carbon gain values measured throughout the
year for each species. The four species exhibited differences in phenology
including germination, length of growth season, and timing of reproduction.
Hydrocotyle bonariensis showed differences in reproduction based on microsite, with 6.5%, 17%, and 35% of production allocated to sexually reproductive structures in foredunes, primary dunes, and swale, respectively.
Foredune and primary dune leaf production was 28% and 50% of swale
leaf production, respectively. Amaranthus pumilus and I. imbricata had
similar light response curves with no indication of light saturation (maximum values recorded were 25 umol m-2 s-1) while C. edentula and H.
bonariensis reached light saturation at 1300 and 1000 umol m-2 s-1 PAR,
respectively. Total estimated carbon gain was 46.2, 44.4, 45.3, and 69.0
mol m-2 yr-1 for A. pumilus, C. edentula, H. bonariensis, and I. imbricata,
respectively. Although notable differences in microsite selection, phenology, and physiology are apparent among the four species, ecophysiological
and life history characteristics appear integrated to provide the adaptive
advantages necessary for survival in a community subjected to severe episodic events.
HANKS, JOSEPH H.* and KENT A. HATCH. Brigham Young University,
Provo, UT. Genetic variation vs. phenotypic plasticity in growth rates
of side-blotch lizards.
Variation in growth rates has been hypothesized to be an important adaptive
response of many species of lizards. Common garden experiments have
been used to distinguish genetic and environmental components of variation in growth among populations of several of these species. However, the
interaction between genetically based physiological differences and behavioral thermoregulation has never been separated. This study demonstrates
that observed differences in the growth rates of five populations of sideblotch lizards (Uta stansburiana) are primarily genetically based and have
a strong physiological component. We compared the growth rates of 100
adult individuals, collected from 5 different altitudes and latitudes (20 from
each population), and held under different environmental temperature regimes. Individuals from each population were randomly assigned to one of
two treatments 1) exposure to constant temperature (268C, 328C, 358C,
or 388C), and 2) thermoregulation in a temperature gradient of 268C-388C.
Snout-vent-length and weight were measured weekly. We found that populations at lower latitudes and altitudes grew faster at all fixed temperatures.
However, when allowed to behaviorally thermoregulate, growth rates of all
populations became more similar.

HANNON, SUSAN J.* Dept of Biological Sciences, University of Alberta,

Edmonton, AB, Canada. Source sink dynamics of American redstarts
in an agricultural landscape.
The major goal of this project was to complete the seventh year of a longterm program that monitors the reproductive success of American redstarts
in 21 clusters in an agricultural area of Alberta. Clusters that had the highest
probability of being reproductive sources were in larger woodlots that were
more isolated from other woodlots and from farms and had fewer alternative cowbird hosts. We determined whether redstarts can recognize
whether a patch has a high probability of being a reproductive source or
sink. To test this we monitored settlement times of males and females in
spring and found that redstarts were not settling in source clusters first,
suggesting that the cues they use to settle are not related to the ultimate
reproductive success in the cluster. Second, we wished to determine if a
redstart settled in a sink, would it switch to a source the next year. We
found that birds that switched clusters between years were equally likely
to have bred in a source or a sink the previous year, but that birds that had
failed to produce young the previous year were more likely to switch clusters. However, they had an equal probability of switching to a source or
sink, again suggesting that they cannot recognize a source patch. Several
males made out-of-cluster movements, and some had nests in more than
one cluster. We are currently analysing DNA from blood samples taken
from 26 complete families to assess the amount of extra-cluster paternity.
HANOWSKI, JOANN,1,* ROBERT HOWE,2 CHARLES SMITH,3 DAVID
MARKS2 and STEVE PRICE.2 1 University of Minnesota-Duluth, Duluth,
MN; 2 University of Wisconsin-Green Bay, Green Bay, WI; 3 Cornell University, Ithaca, NY. What can birds and amphibians indicate about the
ecological condition of coastal ecosystems?
Birds have a long and successful history as being useful indicators of environmental condition in many ecosystems. In contrast, we are just beginning to understand amphibian ecology and their potential response to environmental disturbances. Individually, or together, these two animal groups
likely respond to anthropogenic disturbance type and severity differently
from each other. Birds are sensitive to habitat alteration within small and
large areas, while amphibians may be better indicators of chemical stressors
in the environment. Our objective was to develop a suite of indicators
derived from monitoring amphibian and bird populations in wetland and
upland systems that could be used to document the condition of coastal
ecosystems of the Great Lakes. We selected wetlands and adjacent upland
sampling areas along a disturbance gradient defined by physical, chemical,
and biological stressor data from the study area. Surveys for calling anurans
and breeding birds were conducted in over 200 coastal wetlands throughout
the Great Lakes Basin in 2002 and 2003. In addition, coastal upland breeding birds surveys were completed in 180 shoreline segments during the
same time period. Breeding bird and amphibian indicators were developed
and their response to the disturbance gradient was used to assess their
usefulness. Preliminary results suggest that breeding birds (upland and wetland) are excellent indicators of physical land-use stressors in the coastal
region. It was more difficult to develop indicators for calling anurans because few species occur in the coastal wetlands surveyed. However, combining wetland bird and amphibian parameters shows promise for their
usefulness in indicating condition of coastal ecosystems.
HANSON, PAUL J.1 and 17 OTHERS.2 1 Environmental Sciences Division,
Oak Ridge National Laboratory, Oak Ridge, TN, USA; 2 Various Institutions, USA, Canada, and UK. Carbon and water cycle simulations for
an upland oak forest from 1993 to 2000 using 13 stand-level models:
Intermodel comparisons and evaluations against independent measurements.
Models represent our primary method for integration of small-scale, process-level phenomenon into a cohesive description of forest-stand or ecosystem function. They also represent our only viable means for testing
hypotheses about the response of forest ecosystems to changing environmental conditions. This poster summarizes the evaluation of 13 stand-level
models varying in their spatial, mechanistic, and temporal complexity for
their ability to capture intra- and inter-annual components of the water and
carbon cycle for an upland, oak-dominated forest of eastern Tennessee.

Comparisons between model simulations and observations were done for

hourly, daily and annual time steps. Response surfaces of carbon and water
flux as a function of environmental drivers, and a variety of goodness-offit statistics were used to judge model performance. A single model did
not consistently perform the best at all time steps. Not surprisingly, those
models missing key forest components or processes (e.g., soil water content) were unable to provide accurate predictions of ecosystem responses
to short-term drought phenomenon. An inability to correctly capture physiological processes under drought, however, was not necessarily an indicator of poor long-term water and carbon budget simulations. This is possible because droughts in the subject ecosystem were of short duration over
the 1993 to 2000 period, and therefore had a small cumulative impact.
Models using hourly time steps, based on mechanistic processes, and having a realistic spatial representation of the forest ecosystem provided the
best predictions of observed data. Models designed for wide application to
a range of ecosystems performed poorly in this comparison, suggesting the
need for continued evaluation and improvement of ecosystem models targeted for extrapolation to broad geographic regions. Co-authors: J.S. Amthor, S.D. Wullschleger, K.B. Wilson, R.F. Grant, A. Hartley, D. Hui, E.R.
Hunt Jr., D.W. Johnson, J. Kimball, A.W. King, Y. Luo, S.G McNulty, G.
Sun, P.E. Thornton, S. Wang, M. Williams, and R.M. Cushman.
HARAMOTO, ERIN R.,* ERIC R. GALLANDT and THOMAS MOLLOY. University of Maine, Orono, ME, USA. Effects of Brassica cover
crops on weed dynamics.
Allelopathic cover crop residues may contribute to weed management by
reducing weed establishment and suppressing the growth of weed seedlings. Members of the Brassicaceae contain glucosinolates (GSLs), which
degrade to compounds that are toxic to a variety of organisms. Brassicas
used as cover crops include, but are not limited to, canola (Brassica napus),
rapeseed (also Brassica napus), and yellow mustard (Sinapis alba). Glucosinolate content of these cover crops varies widely, with canola having
very low GSL concentrations, yellow mustard typically having very high
concentrations, and rapeseed being intermediate between the two. During
the summer of 2002, two experiments were conducted in Stillwater, ME,
to examine the effect of different cover crop residues on weed and crop
emergence and the effect of these residues on growth and reproduction of
redroot pigweed (Amaranthus retroflexus) and green bean (Phaseolus vulgaris). Weed and crop emergence was measured following incorporation
of six cover cropsthree brassicas (canola, rapeseed, and yellow mustard)
and three non-brassicas (buckwheat (Fagopyrum esculentum), crimson clover (Trifolium incarnatum), and oats (Avena sativa)). Incorporated cover
crop residues reduced weed and crop emergence by 33% (p50.036). However, emergence following brassicas and non-brassicas was similar. In the
second experiment, monocultures of redroot pigweed, green bean, and a
mixture of the two were seeded into incorporated residues of canola and
yellow mustard. Redroot pigweed grown with green bean was smaller than
redroot pigweed grown alone; the same was true for green bean. However,
no residue effect on growth or yield of redroot pigweed or green bean was
found. Despite the presence of glucosinolates in the incorporated residues,
the residue-mediated effects of brassicas on weed dynamics appear to be
similar to other commonly grown cover crops. Brassica cover crops may
not provide distinct advantages for weed control, but may suppress other
soil-borne pests.
HARDESTY, BRITTA DENISE.* Smithsonian Tropical Research Institute,
Panama City, Panama. Linking dispersers and seed fate: Seed arrival
and survival in a neotropical tree.
Seed dispersal is a fundamental process that shapes plant communities at
local and large scales in both space and time. From the animal perspective,
researchers assess disperser movement patterns, generally addressing seed
deposition without empirical evidence of seed fate. From the plant perspective, ecologists have traditionally relied on seed traps and other indirect
measures of determining seed shadows and dispersal curves. Recent advances in molecular techniques now permit analyses of gene flow via seed
dispersal using maternal tissue from seeds, and enable us to determine
genetic relationships among large numbers of individuals and populations
of plants at widespread geographic scales in contemporary and historical
time. Using microsatellite genetic markers developed for the vertebrate-

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139

dispersed dioecious tropical tree, Simarouba amara, I inferred the parentage of seeds and seedlings in the Barro Colorado Island 50 ha plot and
related this to fine-scale spatial genetic structure and successful seed arrival
and survivorship near and far from the parent plant. The spatial distribution
of parents and their offspring, coupled with field measures of floral and
fruit production, crown area and tree height, was used to calculate relative
reproductive success and the frequency of long-distance seed and pollen
movement. The dispersal dynamics examined here provide insight into a
demographic filtering stage that is critical for generating forest structure.
This study assesses the relative importance of gene flow via pollen versus
seed with respect to distance, thus gaining insight to the role of frugivores
in structuring plant communities at multiple forest sites characterized by
high species richness and tree density, but with different ecological histories.
HARE, J. DANIEL1,* and ELIZABETH ELLE.1,2 1 University of California, Riverside, California, United States; 2 Simon Fraser University, Burnaby, British Columbia, Canada. Survival and seed production of sticky
and velvety Datura wrightii in the field.
Leaves of Datura wrightii in California are covered either with glandular
trichomes (sticky plants) or non-glandular trichomes (velvety plants), and
sticky plants are resistant to many of D. wrightii9s insect herbivores. Theoretical models suggest that variation in levels of resistance to herbivores
may persist if resistance is costly and herbivore damage is variable. If so,
then natural selection should favor resistant plants when herbivore damage
is high and disfavor it when damage is low. However, without long-term
equivalence between costs and benefits, then natural selection either should
eventually drive the trait to fixation if the trait has a net benefit or eliminate
it if the trait has a net cost. In previous short-term studies, we found that
the production of glandular trichomes by D. wrightii had a net fitness cost
even in the presence of herbivores. If always true, then natural selection
should eliminate this expensive resistance trait. We monitored survival and
seed production of sticky and velvety D. wrightii in the presence of herbivores in 11 natural populations over four or five years. In all populations
where both types occurred, the finite rate of increase for velvety plants was
from 60% to 274% greater than for sticky plants. Plant survival averaged
between two and three years and did not differ significantly between plant
types. Because seed production consistently favored velvety plants, we predict that the frequency of sticky plants should decline, and this prediction
was met within five years in three of eight monitored populations where
both types occur. Results strengthen our prediction that, in the absence of
additional benefits to the suite of characters associated with the production
of glandular trichomes, sticky plants should be eliminated from D. wrightii
populations.
HARLEY, CHRISTOPHER D.G.,1,* LUKE J.H. HUNT1 and ROBERT T.
PAINE.2 1 Hopkins Marine Station, Oceanview Blvd., Pacific Grove, CA,
USA; 2 University of Washington, Department of Zoology, Box 351800,
Seattle, WA, USA. Seemingly gradual changes in climate provoke punctuated biological responses.
Biotic response to changing climate can occur gradually in small increments, or suddenly in large jumps. Because limited sampling through time
may detect change in the former case, but not in the latter, distinguishing
between these two patterns is central to our ability interpret available data
and forecast future biological change. Unfortunately, the high resolution,
long-term datasets required to make such distinctions are rare. Here, we
use photographic reconstruction and direct measurements to describe
changes in the vertical distributional limits of two intertidal algae in response to changing sea level. In Pacific Grove, CA, sea level has been
rising 1-2 mm/yr for the past several decades. In approximately 1960, the
upper limit of Endocladia muricata shifted upshore by 30 cm within a few
years. Following this rapid shift, Endocladias upper limit has remained
remarkably constant to the present. On Tatoosh Island, WA, where tectonic
uplift outpaces sea level rise by 1.4 mm/yr, the upper limit of Mazzaella
parksii was relatively stable for at least 15 years. In the 1990s, the upper
limit of Mazzaella shifted downwards by 25 cm within three years to a
new level where it has remained since 1995. In both cases, distributional
changes were sudden and could not have been predicted by prior trends in
the data. These results caution against the use of stasis in available data as
an indicator of future stability.

140

Abstracts

HARMON, JASON P.1,2,* and DAVID A. ANDOW.1 1 University of Minnesota, St. Paul, MN, USA; 2 University of California-Davis, Davis, CA,
USA. The ecological mechanisms of indirect interactions among natural enemies and multiple prey.
Despite their simple veneer, agroecosystems harbor complex and diverse
arthropod communities. Applied ecologists have historically dealt with this
complexity by focusing on just individual organisms or the direct interactions between two focal species. More recently, we have begun to understand the crucial role of indirect interactions, interactions between two organisms that depend on the presence of a third. The traditional model of
arthropod biological control is itself an indirect interaction: crop plants
benefit from natural enemies when that natural enemy controls a pest herbivore that would otherwise harm the plant. Simplified community modules
have provided important foundations for looking at indirect interactions
and their effects on arthropod populations and community processes. However, some theoretical studies have used these community modules to produce sweeping generalizations about how indirect interactions should function. While these simple predictions are attractive, empirical studies have
not necessarily supported them. I propose that we need a better understanding of the ecological mechanisms that generate these indirect interactions
to resolve this apparent conflict and improve our ability to predict their
consequences. I will illustrate this mechanistic framework and its potential
use with two community modules: shared predation where a shared natural
enemy attacks two prey species, and intraguild predation where two predators attack each other and a single, shared prey. Using this type of framework, we may be able to make better a priori predictions of community
interactions and the magnitude of their effects in different systems; ultimately helping us best manage agroecosystems for their ecological services.
HARPOLE, W S.* and D TILMAN. University of Minnesota, St. Paul,
MN. Non-neutral patterns of species abundance explained by species
tradeoffs.
The neutral theory of biodiversity has been proposed as a null model
against which to test tradeoff-based theories of the maintenance of species
diversity. Patterns of plant species abundances in late-successional grasslands in MN, USA deviate from assumptions of the neutral theory in two
important aspects. First, the relative abundances of species among independently assembled late-successional old fields are not random: some
plants are always dominant whereas others always rare. Second, species
traits are not identical and the observed differences explain which species
are abundant or rare. Data from 28 late-successional perennial grasses and
forbs grown in monoculture for 5 years show that species differ in three
correlated traits indicative of competitive ability for nitrogen: 1. R*, i.e.
the level to which a species can reduce limiting soil nitrogen pools, 2.
tissue nitrogen concentrations, and 3. biomass in roots, leaves, and reproduction. The species with greater abundance in these nitrogen-limited habitats have significantly lower R* values, have a greater proportion of their
biomass in roots and less in reproduction, and have lower tissue N content.
Rare species are expected to be poorer competitors for N, but do have
greater allocation to reproductive biomass, suggesting they may persist via
a competition-colonization tradeoff. Our results refute several major predictions of neutral theory. In particular, communities assemble in a nonneutral manner because the same species become dominant or rare for a
wide range of starting conditions and the identities of rare and dominant
species are predominantly determined by species traits rather than by past
history or drift.
HARRELL, WADE C.,1,* SAMUEL D. FUHLENDORF,1 DAVID M. ENGLE1 and ROBERT HAMILTON.2 1 Oklahoma State University, Stillwater,
OK; 2 The Nature Conservancy, Pawhuska, OK. Increasing grassland heterogeneity with fire and grazing: Implications for grassland birds.
Disturbance-driven heterogeneity has been suggested as a critical process
in the maintenance of species in many ecosystems. For example, in grasslands, fire and grazing interact to generate variably scaled disturbance
patches across the landscape, contributing to a shifting mosaic that presumably enhances biodiversity. Grassland birds, a declining group within grasslands, have species-level affinities for certain structural characteristics of

vegetation, thus changes in spatial heterogeneity may alter bird community

dynamics. Our objective was to investigate grassland bird community dynamics in relation to changes in spatial heterogeneity. Our study site was
located on the Tallgrass Prairie Preserve in Northeast Oklahoma, USA. We
varied the spatial and temporal extent of fire and introduced livestock within management units in an attempt to increase spatial heterogeneity (patch
burn treatment, n52) and contrasted this disturbance regime with a relatively uniform application of fire and grazing (control, n52). We monitored
bird abundance during the spring of 2001-2003 (May-June) using point
counts to determine the influence of our patch burn treatment on the bird
community. Our patch burn treatment resulted in more spatially heterogeneous management units than our controls. Increased heterogeneity in the
patch burn treatment led to greater coexistence of grassland bird species
with one species, Henslow9s sparrow, only occurring within the unburned
portion of the patch burn treatment. Species-level abundance varied among
the patch types that occurred within the patch burn treatment, with some
species exhibiting greater abundance in unburned patches and other species
occurring in the greatest abundance in the most recently burned and grazed
areas. The more uniform application of disturbance in our controls resulted
in less spatial heterogeneity and a subsequent reduction in the abundance
of some bird species. Our study demonstrates that increased spatial heterogeneity can alter grassland bird community dynamics, and may assist in
the conservation of declining species within grassland ecosystems.
HARRIS, REID N.1 and PATRICE M. LUDWIG.2,* 1 James Madison University, Harrisonburg, VA, USA; 2 University of Virginia, Charlottesville,
VA, USA. Resource level and reproductive frequency in female fourtoed salamanders, Hemidactylium scutatum.
Skipping a reproductive event may be selectively advantageous if the reduced reproductive output is balanced by gains in survival and lifetime
fecundity. Alternatively, skipping a reproductive event may be due to the
effects of resource limitation. Data from a 10-year field study showed that
females that did skip reproduction had higher growth, which can translate
into higher fecundity, and tended to have higher survival. However, these
benefits did not compensate for the absence of reproduction in a year
caused by skipping. We experimentally examined whether female four-toed
salamanders (Hemidactylium scutatum) skip a reproductive event because
of a lack of adequate resources after nesting and before the onset of winter.
A manipulation of food level after the nesting period revealed that females
fed a high food ration were twice as likely to return the next year to oviposit
as females fed a maintenance food level. Females from the maintenance
food level treatment that did return were in similar condition to females
from the high food ration treatment. This result suggests that a threshold
body condition is necessary before reproduction is attempted. We conclude
that skipping reproduction in this species is due to the proximate effects
of resource limitation.
HARRIS, WYLIE N.,* THOMAS W. BOUTTON and R. J. ANSLEY. Texas A&M University, College Station, Texas. Soil carbon and nitrogen
cycling in subhumid temperate grasslands: Effects of seasonal fire and
simulated grazing.
Soil carbon (C) and nitrogen (N) dynamics in grassland ecosystems depend
on interactions between climate, plant productivity, soil fertility, and disturbance. We tested the hypotheses that turnover of C and N are 1) accelerated by simulated grazing, and 2) reduced by fire, in southern Great
Plains native grasslands. Seven combinations of fire and simulated grazing
(clipping) treatments were implemented annually from 1999-2002 on replicated experimental plots. Prescribed fires were conducted in spring (Feb/
Mar) or fall (Oct/Nov). Simulated grazing treatments consisted of light
(annual) or continuous (monthly during growing season) clipping. Response variables, measured monthly, included plant cover and biomass by
species, soil moisture and temperature, soil microbial biomass (SMB) C
and N, soil inorganic N, soil respiration, and net N mineralization. All
response variables displayed pronounced seasonal variation, particularly in
relation to pronounced summer drought in 2000 and 2001. Within that
climatically-determined context, season of fire and clipping affected plant
and soil responses relative to controls. Both fire and clipping reduced plant
cover and biomass, with spring treatment favoring C4 grasses. Both fire
and clipping reduced soil moisture and increased soil temperature. SMB-

C and -N showed transient (1-3 month) declines after spring and fall fire,
while clipping reduced soil inorganic N. Soil respiration was controlled by
soil moisture and temperature, with a lack of treatment effects stemming
from the small magnitude of treatment differences in the latter variables.
Contrary to our hypothesis (1), these results suggest that fire and simulated
grazing are similar in their effects on C and N cycling in this ecosystem.
Continued removal of aboveground biomass may retard C and N cycling
by limiting inputs of aboveground plant material to the soil. The mechanism and intensity of biomass removal (i.e. fire or clipping) appear to be
of secondary importance, and timing of removal may be more critical.
HARRISON, PHILIP M. University of Northern Colorado, Earth Sciences
Dept., Greeley, CO, USA. How much water do trout streams need?
Instream flow water allocations became common after the passage of the
National Environmental Policy Act of 1969. Two contemporary models
used by the Biological Resources Division are Physical habitat simulation
System and SALMOD. The Physical habitat simulation System models the
physical properties of a stream habitat (hydrology and substrate) and then
conceptually represents the fish habitat requirements by weighted useable
area (WUA) or habitats theoretically suitable for fish. The SALMOD model
is an extension of the Physical HABitat SIMulation System model. SALMOD adds and incorporates fish cohorts throughout their life cycle. As
part of monitoring the effects of a hydropower dam on the Tule river in
California, the CompMech model was developed. This research will use a
common dataset as input to compare the three stream models of Physical
HABitat SIMulation System, SALMOD and CompMech. Model comparison will include advantages and disadvantages, the amount of data required
to run the models (input files), the model output (output files), and how
difficult it is to learn and use the models. The processes used by each
model will be compared. CompMech and SALMOD will be used to study
two variables, discharge and trout population. Population limiting events
will be determined by simulating hydrographs that cause a 50 percent decline in trout population (number or biomass) over ten years or less.
HARRISON, SANDY P.1,* and COLIN I. PRENTICE.2 1 Max Planck Institute for Biogeochemistry, Jena, Thuringia, Germany; 2 Max Planck Institute for Biogeochemistry, Jena, Thuringia, Germany. Global patterns
of plant functional diversity and palaeovegetation.
Global analysis and modelling of vegetation changes in response to past
and future climates requires some way to classify plants, irrespective of
phylogeny, based on the biophysical principles of environmental adaptation
and tolerance. This idea lies behind the concept of the plant functional type
(PFT). A global scheme of PFTs, based on life form, leaf form, leaf phenology and mechanisms for tolerance of climatic extremes, is currently
under development. A prototype has been applied to map vegetation patterns today, during the mid-Holocene and at the last glacial maximum
(LGM) based on pollen and plant macrofossil data. Abundances and distributions of PFTs have varied substantially during the period since the
LGM. Associations of PFTs (vegetation types or biomes) have changed
between glacial and interglacial regimes giving rise to biomes that have no
analog under modern conditions. These changes can be explained by a
combination of changes in climate and atmospheric CO2 concentration. CO2
concentration is expected to influence vegetation composition through effects on plant water-use efficiency and C3-C4 plant competition. In the
tropics, these effects appear to have been as important as climate changes
in determining the extent of forest. Further efforts are underway to improve
the PFT scheme, especially by including a wider range of traits related to
drought and cold tolerance, and through quantitative analysis of trait-environment relationships.
HART, MIRANDA M.* University of Guelph, mhart@uoguelph.ca,
Guelph, ON, Canada. The effect of pesticides on soil biodiversity of
urban ecosystems.
It is well known that the application of herbicides (spraying) on urban
landscapes has a negative impact on the diversity of plants. However, the
indirect effects of spraying on the diversity and functioning of soil microorganisms is not known. To determine this, I compared front yards in
Guelph, Ontario that were regularly sprayed with herbicide to those that

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141

were not sprayed at all. I found that microbial biomass was significantly
lower in sprayed lawns, and fungi were more greatly affected than bacteria.
Infectivity potential by arbuscular mycorrhizal fungi was also reduced. Effects were also oberved on higher trophic groups. The number and diversity
of protozoa, nematodes, collembola, and mites were also reduced in response to herbicide application. In another controlled microcosm study, I
then investigated the consequences of such reduced microbial activity on
ecosystem functioning (plant productivity and stability to drought). I found
that microbial communities extracted from sprayed yards were less able to
promote plant growth, especially under drought conditions. These results
indicate that herbicide application can have negative indirect effects on soil
organisms with strong feedback responses on ecosystem productivity and
stability.
HART, ZACHARY H.,1,* ANGELA C. HALFACRE1 and MARIANNE K.
BURKE.2 1 College of Charleston, Charleston, SC, 29424; 2 USDA Forest
Service Center for Forested Wetlands, Charleston, SC, USA. Assessing
stakeholder views of sweetgrass (Muhlenbergia filipes) management in
the South Carolina lowcountry.
Sweetgrass (Muhlenbergia filipes) is a non-timber forest resource found in
coastal areas from North Carolina to Texas. The grass has special cultural
and economic importance in coastal South Carolina where it is used by the
local Gullah community in a specialized form of coiled basketry. Sales of
these baskets provide a crucial source of income for many basket makers
and are a major component of the Charleston Areas historical and cultural
identity. The sweetgrass on which these basket makers depend, however,
is becoming increasingly unavailable through habitat destruction, private
property rights, and range limitation. Many basket makers must now buy
the raw material from men who are able to travel outside the Charleston
Area, sometimes as far as Georgia and Florida, to harvest the grass. Unless
local sweetgrass supplies are restored, the art form may soon disappear.
This study examines stakeholder opinions and perceptions of past, current,
and future sweetgrass management. Twenty-three interviews were conducted with Charleston Area basket makers, and interview transcripts were
analyzed for emergent themes using content analysis (a technique to objectively analyze text). Survey respondents indicated that buying grass rather than collecting it has become standard practice and that development is
the primary reason for the difficulty in accessing the resource. Further,
respondents indicated several potential solutions to the problem and expressed their willingness to contribute their time to restoration efforts. This
study provides essential input for preserving the centuries-old art of sweetgrass basketry and offers valuable lessons for incorporating public and
stakeholder participation in environmental decision-making processes.
HARTLE, R. TODD.* University of Nevada, Reno, Reno, Nevada, USA.
Larrea tridentata defense of root territory in the Mojave Desert.
Larrea tridentata roots have been shown to exclude the roots of other
Larrea or of another Mojave Desert shrub, Ambrosia dumosa, from a soil
zone surrounding the Larrea roots (Mahall and Callaway 1991, 1992). The
existence of root zone territory defense could explain field observations of
unequal distribution and community structure in desert shrubs (Fonteyn &
Mahall 1978, 1981; Brisson & Reynolds 1994), as well as add supporting
evidence to the theory of allelopathy. In the spring of 2002, I developed a
field method based on a phenomenon called hydraulic redistribution (formerly hydraulic lift) (Richards & Caldwell 1987; Yoder & Nowak 1999)
In this method, I looked at paired shrubs, including Larrea with Larrea,
Ambrosia with Ambrosia, and Larrea with Ambrosia, and tested whether
the second shrub of the pair (receiver shrub) had access to the soil immediately surrounding the roots of the first shrub (donor shrub). Using
hydraulic redistribution to place deuterated water (D2O) into the soil immediately surrounding the roots of the donor shrubs, I sampled the tissue
of the receiver shrub and tested it for the presence of the deuterated water.
The results of this study not only add to our knowledge of belowground
interactions among Mojave Desert shrubs, but also add significantly to the
evidence on the concept of territory defense in the plant kingdom, i.e. that
plants actively defend soil resources for their exclusive use.

142

Abstracts

HARTLEY, ANNE E.,* HEATHER L. REYNOLDS, JAMES D. BEVER

and PEGGY A. SCHULTZ. Indiana University, Bloomington, IN. Nitrogen acquisition by old field perennials: The role of arbuscular mycorrhizal fungi.
The fundamental role of arbuscular mycorrhizal fungi (AMF) in plant P
acquisition has long been appreciated, but we know comparatively little
about the role of AMF in N acquisition. We asked whether AMF improved
the ability of old field perennials to grow on a range of inorganic and
organic N sources, and whether AMF facilitated differential performance
of plant species on different N sources (N niche partitioning). We manipulated form of N (control versus NH4, NO3, urea, glycine, and chitin) and
AM fungal species (control versus 4 AMF species) for 5 old field perennials grown in a greenhouse under N-limiting conditions. Based on shoot
biomass after 5 months of growth, we found that AMF never enhanced
plant growth and that some AMF species depressed growth for some plant
species (significant plant x fungus interaction). We found no evidence for
AMF-facilitated N niche partitioning, although groups of plant species did
respond differently to form of N (significant plant x N interaction).
HARTLEY, LAUREL M. and JAMES K. DETLING. Department of Biology, Colorado State University, Fort Collins, Colorado, USA. Effects of
Black-tailed prairie dogs on plant community composition and nutrient
cycling in the shortgrass steppe.
Black-tailed prairie dogs (Cynomys ludovicianus) physically influence their
environment by intensely grazing, by actively removing taller vegetation,
and by creating extensive burrow systems. These activities can have profound impacts on plant communities and nutrient cycling. In the mixedgrass prairie of South Dakota and the savanna of Texas, grazing by prairie
dogs results in an overall lower canopy height, decreases in litter, standing
biomass, and graminoids relative to forbs, and an increase in crude protein
content of some plant species. Compared to effects on the mixed grass
prairie, the effects of prairie dogs on the shortgrass steppe may be qualitatively or quantitatively different for several reasons. The shortgrass prairie
is drier and is dominated by drought resistant species such as blue grama
(Bouteloua gracilis) that are also resistant to grazing. Unlike on the mixedgrass prairie, prairie dog colonies on the shortgrass steppe are subject to
periodic extinction and recolonization events due to plague. Extinction
events may serve to temporarily alleviate grazing pressure from prairie
dogs. This poster presents the initial results from a comprehensive study
of the effects of prairie dogs on plant community composition and nutrient
cycling on the shortgrass steppe of Northeastern Colorado. Plant community composition, species richness, biomass, and nitrogen to carbon ratios
were measured on 3 recently colonized towns, 3 recently abandoned towns,
and 3 towns that had been occupied consistently for over 15 years and
compared to off-town control sites in similar habitats.
HARTMAN, WYATT H.,* PALLAOOR V. SUNDARESHWAR, RYTAS
VILGALYS, GREGORY L. BRULAND and CURTIS J. RICHARDSON.
Duke University, Durham, North Carolina. Linking microbial diversity
and ecosystem functioning across land use patterns.
Our present work attempts to draw links between soil microbial diversity,
nutrient speciation, and land use to illuminate the relationship between
ecosystem form and function. We will compare soil phosphorus speciation
from NMR spectra to microbial biomass and microbial diversity metrics
such as operational taxonomic unit (OTUs) accumulation curves, community phylograms, and presence of microbial species possessing distinctive phosphorus metabolic pathways (e.g. polyphosphate and phosphonate
accumulation). We will explore functional relationships of microbial diversity across pristine, disturbed and restored wetland ecosystems. Our network of sites encompass a range of impacts from nutrient enrichment to
conversion to agriculture and forestry. Preliminary P NMR data suggests
that sites impacted by nutrient enrichment and/or agricultural activity exhibit reduced diversity in chemical speciation of this important nutrient.
Specifically, impacted sites are marked by the absence of organophosphates
and polyphosphates as compared to reference sites. Importantly, restored
sites exhibit partial recovery of nutrient species diversity, potentially indicating functional restoration. We anticipate microbial genomic analyses
will show differences in diversity and composition across land use patterns.

Collectively, these data will help establish links between changes in microbial communities and biogeochemical cycling of nutrients. Our study
has important implications for evaluating the progress of ecosystem restoration efforts.
HARTVIGSEN, GREGG.* SUNY Geneseo, Geneseo, NY. Quasi-chaotic
differences among families of logistic curves.
Chaotic dynamics are characterized by populations diverging exponentially
when initial conditions differ only slightly, yielding a positive Lyapunov
exponent. The logistic equation y 5 ax(1 - x) yields chaotic behavior when
a exceeds about 3.57. This divergence, however, cannot increase indefinitely due to the range constraint of x in the above equation (0 # x # 1).
Therefore, the long-term average differences among runs with differing
initial conditions must be finite. To explore this accumulated variance computer simulations were conducted for populations starting at x 5 0.5 and
with values for a known to generated positive Lyapunov exponents. Ten
thousand additional runs were made for each setting of a using new initial
populations determined by adding a small value (10-7) to each prior initial
population. Each of these was run for 10,000 iterations and the last 5,000
iterations were analyzed. The dynamics of the average differences among
these simulated populations were found to be quasi-chaotic, lying between
chaos and deterministic outcomes. These results suggest the average dynamics of a deterministic chaotic system lie at the edge of chaos.
HARTWAY, CYNTHIA.* Department of Ecology and Evolutionary Biology, University of California, Santa Cruz, CA, USA. Is one strategy good
enough? How spatial and temporal variability may mitigate fitness
costs associated with aggregative behavior in a willow leaf beetle.
The herbivorous beetle, Chrysomela falsa, is typically found feeding in
aggregations within stands of its preferred hosts (Salicaceae). Predation
studies have shown that individual beetle fitness (measured as long-term
reproductive success) is a parabolic function of beetle density. At low conspecific densities, C. falsa experiences high mortality due to intense predation pressure, whereas at high densities beetles experience decreased fecundity and survival due to increased resource competition and disease
incidence. Despite this trend, studies of adult movement and oviposition
behavior indicate that, in the absence of host quality differences, beetles
consistently prefer to remain at and oviposit within areas of conspecific
density high enough to have a significant negative impact on their longterm reproductive success. In other words, C. falsa appears to follow a rule
of thumb: always seek out and remain within areas of high beetle density.
I use a dynamic state variable model, incorporating the use of a spatially
explicit landscape, to compare the fitness consequences of the observed
beetle behavior with behavior that is predicted to optimize individual beetle
fitness. I then assess how the interaction between the cost of dispersal and
the configuration of beetle densities and disease across space and time
mitigates the apparent fitness cost of C. falsas behavioral strategies.
HARVEY, CHAD T.1,2,* and MICKY D. EUBANKS.1 1 Auburn University,
Auburn, AL; 2 University of Wisconsin-Madison, Madison, WI. The effect
of habitat complexity on interactions within a terrestrial food web.
Habitat complexity and species diversity have long been believed to affect
interactions within food webs. It has even longer been believed that species
diversity and the strength and number of interactions among species are
related. Although appealing to many ecologists, and highly significant to
biological control, there are few studies that have experimentally tested
these ideas in conjunction. The focus of this study was to experimentally
determine the effect of habitat complexity on the number and strength of
natural enemy (NE)- herbivore interactions, the number of interactions
among NE, and the effect of interactions among NE on herbivore suppression in collards (Brassica oleracea). Based on interaction models for the
11 most abundant herbivore, we found that habitat complexity did not
affect the number of NE-herbivore interactions, nor did it affect the strength
of NE-herbivore interactions (paired t 5 2.31, df 5 10; p . 0.05). Furthermore, there was no relationship between the number of NE attacking
a herbivore and the mean strength of NE-herbivore interactions (top-down
control) (R 5 0.41, df 5 10, p . 0.05). Habitat complexity significantly
affected the number of interactions among NE associated with two herbi-

vores, however, there was no correlation between the number of interactions among NE and the strength of NE-herbivore interactions (R 5 0.29,
df 5 10, p . 0.05, simple habitat; R 5 -0.53, df 5 10, p . 0.05, complex
habitat). The results of this study suggest that interactions among NE and
herbivores and interactions among NE do not respond to changes in habitat
complexity as predicted by classical community ecology theory. This study
highlights that the basis of our knowledge of interactions within food webs
is highly theoretical and more empirical studies need to be performed before we can identify the ecosystem characteristics affecting NE assemblages and top-down control.
HASSELQUIST, NILES J.,1 MATT J. GERMINO,1 TERENCE P. MCGONIGLE1 and WILLIAM K. SMITH.2 1 Idaho State University, Pocatello,
ID; 2 Wake Forest University, Winston-Salem, NC. Differences in ectomycorrhizal infection of conifer seedlings across an alpine-treeline ecotone: Relative importance of variation in soil properties and microclimate.
The importance of mycorrhizae to plants in stressful environments, such
as for conifer seedlings above the elevation limits of forest, is not well
known. Lower ectomycorrhizal infection rates could contribute to lower
seedling survival in alpine soils. Experiments were designed to test whether
seedlings were less likely to form an ectomycorrhizal symbiosis at higher
versus lower altitude sites, and whether differences in microclimate or soil
properties among these sites were more likely to explain variation in infection rates. Ectomycorrhizal infection of Engelmann spruce (Picea engelmannii) seedlings was measured across an altitudinal gradient from subalpine forest to alpine meadow. The percent of root tips infected was determined for seedlings that germinated in soils transplanted in a reciprocal
fashion among the three sites. Three seedlings from each combination of
soil and site were harvested biweekly throughout the growing season, and
the percent of root tips infected were determined microscopically. Mean
infection rates at the end of the growing season were 30, 13, and 0% for
seedlings in the forest, treeline-ecotone, and alpine soils, respectively, and
were not affected by transfer to another site. These results indicate that
conifer seedlings may be less likely to establish ectomycorrhizal symbioses
in treeless, alpine meadows compared to more forested sites. Moreover, site
differences in infection appeared more attributable to variation in soil properties, possibly inoculum availability, than to microclimate and corresponding effects on seedling physiology.
HATCH, KENT A.* and JESSICA L. MAKIN. Brigham Young University,
Provo, UT, USA. 15N enrichment of lizard uric acid: Can 15N be used
as a measure of body condition in the wild?
Stable isotopes are used by animal physiological ecologists to determine
such things as dietary sources, migration, and trophic levels. Attempts have
been made to use d15N values as indicators of body condition as well. The
assumption is that proteins are 15N enriched during anabolism whereas nitrogenous wastes are 15N depleted relative to diet. Catabolism then only
further enriches the tissues as 15N depleted urates are excreted. However,
this model is based, in part, on only a single study of 8 cattle demonstrating
15
N depletion of urea relative to diet. Because of their small size and ease
with which uric acid can be collected, lizards are a logical system to use
to test the hypothesis that, across taxa, nitrogenous wastes are 15N depleted
relative to diet. Based on a study of twenty side-blotched lizards, we show
that nitrogenous wastes are 15N depleted across taxa. We evaluate the usefulness of this method for determining body condition on the wild.
HATTON, ELIZABETH S.,1,* RUDY BUENO1,2 and ROBERT PARMENTER.1 1 University of New Mexico, Albuquerque, New Mexico; 2 Albuquerque Environmental Health Department, Albuquerque, New Mexico.
Ecology of larval mosquitoes in New Mexico and transovarial transmission of West Nile virus.
The state of New Mexico has implemented a comprehensive surveillance
program in anticipation of possible outbreaks of West Nile virus. This program not only provides detection of WNV if present but also provides
valuable information on the ecology of mosquito species. Surveillance for
WNV is conducted in various ways. Adult mosquitoes are collected using
CDC light traps and gravid traps and specimens are forwarded for WNV

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143

testing. In 2003, mosquito larvae will also be collected and tested for WNV.
If WNV is found in larval specimens, this will provide further evidence
that WNV can be transovarially transmitted, i.e. transmission from the female mosquito to the offspring. Preliminary data collected in 2002 show
that mosquito species identified and collected from adult and larval surveillance in the same area are different. More data will help to demonstrate
the efficacy of various trapping methods for various mosquito species. This
information will also aid in determining the typical distances female mosquitoes in this specific environment travel between breeding site, blood
meal, and egg deposition, and can ultimately help to predict whether occurrence of mosquito-borne disease will be localized or dispersed and the
impact that specific ecological parameters of New Mexico have on disease
transmission. These data can then be used to maximize the effectiveness
of integrated pest management in the event of a disease outbreak, and to
gain improved understanding of the ecology of the regions mosquitoes,
specifically as it relates to the epidemiology of the West Nile virus.
HAWKES, CHRISTINE V.,1,2,3,* MARY K. FIRESTONE1 and CARLA
DANTONIO.2 1 Dept. Environmental Studies, Policy, & Management,
Ecosystem Sciences Division, Univ. California, Berkeley, CA, US; 2 Dept.
Integrative Biology, Univ. California, Berkeley, CA, US; 3 Dept. Biology,
Univ. York, York, UK. Interactions of exotic species with rhizosphere
microbial communities and consequences for nitrogen cycling.
The success of exotic plant invasions and their subsequent impact on ecosystems may be mediated by interactions with belowground microbial and
mycorrhizal communities. We used a combination of DNA-based characterization and 15N pool dilution to link changes in plant and mycorrhizal
communities with changes in nitrogen cycling. In a California grassland,
small alterations of mycorrhizal community composition with exotic grass
invasion (Avena barbata, Bromus hordeaceus) affected nitrogen cycling,
with depressed rates of gross mineralization and nitrification when mycorrhizae of exotic grasses were present. In a semi-arid grassland in Utah,
mycorrhizal communities of native grass roots shifted dramatically after
cheatgrass (Bromus tectorum) invasion and were correlated with known
differences in nitrogen cycling. Mycorrhizae can alter nitrogen cycling either directly or through effects on bacterial community composition and
function; we are in the process of quantifying changes to functional genes
involved in nitrification to explore the latter at the California site. We have
demonstrated that plant invasions can transform belowground microbial
and mycorrhizal communities with consequences for ecosystem processes.
The success of exotic plant species in novel habitats may be related to their
ability to use or manipulate belowground associations and alter the ecosystem to the disadvantage of natives. A better understanding of plantmicrobe interactions may facilitate strategies for both invasion prevention
and site restoration.
HAYES, JACK.* Paine College, hayesjt@yahoo.com, Augusta, GA. Phenology of adult phantom crane flies from southeastern states.
As part of a larger study of the Phantom Crane Fly, Bittacomorpha clavipes
Fabr. (Diptera: Ptychopteridae), adult specimens of this species were examined in selected state university and related insect collections in the
southeastern United States during 2002. Data were recorded for each specimen, as to the date and place collected, using information on the label for
each specimen. Insect collections visited were at the University of Alabama
in Tuscaloosa, AL; Auburn University in Auburn, AL; the University of
Florida in Gainesville, FL; Virginia Tech in Blacksburg, VA, and the University of Virginias Mountain Lake Biological Station (MLBS) in Pembroke, VA; and the University of Tennessee, in Knoxville, TN. These collections included a total of 64 adult specimens from these four states. An
additional four adult specimens were collected on 21 JUN 02, at the Pandapas Pond Recreation Area, off Highway 460 west of Blacksburg, VA.
Data from all the insects reviewed showed a range of collection dates from
January through November. The phenology of the Phantom Crane Fly in
the southeastern United States seems to involve at least two peak periods
annually, with a low point in the summer. This contrasts with data obtained
earlier from northern (WI) and northeastern (ME, NH, VT, MA, NY) states,
in which there is more typically a single "summer" generation from April
through September.

144

Abstracts

HAYWARD, APRIL J.* and JUREK KOLASA. McMaster University,

Hamilton, Ontario, Canada. Size matters: 3/4-power scaling of community metabolic rate.
Metabolic rate is known to scale as a function of body mass to the power
of 3/4 in nearly all organisms. Recently, it has been mathematically demonstrated that this relationship may be the result of fractal-like exchange
surfaces, which maximize resource acquisition and distribution efficiency.
Because fractal-like networks maximize efficiency of matter and energy
transfers in living systems, they are favoured in the process of natural
selection. We hypothesize that the 3/4-power scaling law will apply wherever system structure and processes are likely to be selected for, including
integrated ecological systems above the organismal level. We tested if 3/
4-power scaling of metabolic rate applies at the community level. Oxygen
measurements were used to estimate respiration rate in natural aquatic rock
pool microcosm communities at Discovery Bay, Jamaica in January and
June of 1999. Total community respiration rate was slower in larger pools.
Specifically, we found community respiration to scale as a function of rock
pool volume to the power of 0.72 (n536, r2 5 0.631, p,0.0000) in January, 0.77 (n537, r2 5 0.708, p,0.0000) in June, and 0.75 (n573, r25
0.662, p,0.0000) for both dates combined. The results of this study provide strong evidence that 3/4-power scaling applies at the community level.
Furthermore, if the aforementioned mathematical derivation provides a true
account of the mechanism underlying 3/4-power scaling, the results of this
study suggest that fractal-like resource networks exist not only within organisms, but also within communities. In the latter case such networks may
be expressed differently than in organisms, as virtual pathways of material
and energy transfers.
HE, FANGLIANG.* Canadian Forest Service, Victoria, BC, Canada. Local-regional diversity relationships: Immigration, extinction and scale.
While local processes (e.g., competition, predation and disturbance) presumably cause species exclusion and thus limit diversity in a community,
regional processes (e.g., historical events, immigration and speciation) are
assumed to provide source species for colonizing and thus enrich a local
community. Ecologists desire to distinguish between these two sets of processes using evidence for local assemblage saturation, but such efforts have
been controversial and are antithetical to the fact that local diversity bears
an imprint of both. Here we examined the local-regional species richness
relationship from the perspective of the theory of island biogeography and
developed a model that unified assemblages ranging from saturated to unsaturated. The model shows that there is no dichotomous division between
saturated and unsaturated assemblages, the degree of saturation of an assemblage is determined by the magnitude of the difference between colonization and extinction rates and sampling scale. Saturation will result if
extinction rate is large relative to colonization rate, while a lack of saturation will result if colonization rate is relatively large. Because sampling
scale is also predicted to have a similar effect on the shape of the localregional relationship to that of immigration and extinction, the relationship
must be interpreted with caution and in order for it to be useful for inferring
ecological processes, it is necessary to collect data on sampling scale in
any such study.
HE, XINHUA,2 WILLIAM R. HORWATH,1 CAROLINE S. BLEDSOE,1
ROBERT J. ZASOSKI1 and TRAVIS W. IDOL.1 1 University of California
at Davis, Davis, CA; 2 University of California at Davis, Davis, CA. Belowground N transfer among plants in a California oak woodland.
Movement of soil resources among plants through common mycorrhizal
networks (CMNs) in the field has received less attention. We examined
nitrogen (N) movement among different species of trees and grasses that
had either ectomycorrhizae or arbuscular mycorrhizae in a California oak
woodland. The experiment consisted of 3 tree pairs with foothill pine (Pinus sabaniana) being a N donor and receiver plants consisting of foothill
pine, blue oak (Quercus douglasii) or buck brush (Ceanothus cuneatus).
We foliarly applied an 15-NNO3- solution to isotopically label the pine donor. We collected leaf samples from all trees and grasses between the trees
weekly for one month following 15NO3- labeling, and tree root samples at
4 weeks. Samples were analyzed for 15N content. Leaf d15N values at weekzero were 2.37, 2.22, -0.12 and 2.51%0 (per mil) in pine, oak, ceanothus

and grasses, respectively. After 4 weeks, leaf d15N values increased to

45.69%0 in the donor pine and 5.72%0 in receiver pine, to 6.16, 3.70 and
8.12%0 in the oak and ceanothus receivers and grasses, respectively. Root
d15N values were 10.36, 14.05 and -0.13 %0 for the control of pine, oak
and ceanothus, and increased to 29.52%0 in the donor pines, and to 20.84,
19.11 and 8.04%0 for the receiver pine, oak, ceanothus, respectively. Transfer of 15N from the donor tree roots to the receiver tree roots and grasses
occurred at an appreciable rate and amount regardless of the tree pair. Since
some these tree pairs and grasses had different ectomycorrhizae or arbuscular mycorrhizae, direct transfer of N probably did not occur. The enrichment of grasses suggests the root N of the donor tree leaked into the
rhizosphere before being translocated among the receiver trees and grasses.
The similar d15N increase of the receiver tree roots compared to their respective controls suggests that CMNs play an important role in competing
and distributing root N among plants.
HEARD, STEPHEN B.,1,* JOHN D. NASON2 and JOHN O. STIREMAN.2
University of New Brunswick, Fredericton, NB, Canada; 2 Iowa State University, Ames, IA, USA. Parallel host-race formation among goldenrod
herbivores and their parasitoids.
1

Evidence is mounting that among phytophagous insects, new biodiversity

can arise via host-race formation and subsequent speciation - even in sympatry. A few examples of recent host-race formation have been well documented, but it remains unknown whether host-race formation is rampant
or is a rare and special process. An ideal system in which to assay the
frequency of host-race formation is the community of herbivorous insects
and their parasitoids associated with the goldenrods Solidago altissima and
S. gigantea. About 100 species of insect herbivore attack these goldenrods,
and one of them (the tephritid fly Eurosta solidaginis) has well-known host
races (work of Abrahamson et al.). We have used allozyme and mtDNA
markers to test for host-associated genetic differentiation in several goldenrod herbivores and their parasitoids. Host-race formation has been common among gallmaking herbivores: genetically distinct host forms attack
S. altissima and S. gigantea in E. solidaginis, the gelechiid moth Gnorimoschema gallaesolidaginis, and the cecidomyid fly Rhopalomyia solidaginis, and may also exist in the tortricid moth Epiblema scudderiana. Parasitoids of gallmakers can also show differentiation: distinct forms attacking hosts on S. altissima and S. gigantea appear to exist for an encyrtid
parasitoid of Gnorimoschema, Copidosoma gelechiae, and for a torymid
parasitoid of Rhopalomyia, Torymus sp. Overall, our results suggest that
host-race formation is extremely common, at least for goldenrod gallmakers
and their parasitoids; in fact, what is missing to date is much evidence of
goldenrod insects that have NOT evolved host races. Cross-lineage surveys
such as ours will eventually document phylogenetic and ecological factors
that control the likelihood of host-race formation, and therefore shape the
origin of new biodiversity, among insects.
HECKMAN, KATHERINE A.,1,* WENDY B. ANDERSON1 and ALEXANDER WAIT.2 1 Drury University, Springfield, MO, USA; 2 Southwest
Missouri State University, Springfield, MO, USA. Ecological contributions of hypolithic cryptobiotic soil crusts in hyperarid ecosystems.
The important role cryptobiotic soil crusts play in terrestrial ecosystems
has only recently been recognized. In arid ecosystems, this role is amplified
by the fact that soil crusts represent a large percentage of overall productivity. In such environments, where conditions at the soil surface are exceedingly harsh, crusts can be found living underneath semi-translucent
rocks such as quartz. Very little is known about the ecological role of these
hypolithic crusts in arid ecosystems, e.g., their impacts on soil, C-fixation,
N-fixation, and contributions to other taxa, including plants and animals.
We addressed these questions with a study of hypolithic crust communities
on three hyperarid islands in the midriff region of the Gulf of California.
Crusts were found in areas with inherently low soil N content, but were
absent in soils with high N content (i.e. those with seabird guano enrichment). When present, crusts were found on 41% of quartz rocks, covering
approximately 1.0% of the sampling area. Soils underneath encrusted rocks
exhibited higher NO3 and organic content than soils unassociated with
hypolithic crusts. Crusts were also found to alter soil properties such as
pH, element concentrations, moisture content, and salinity. The hypolithic
crusts in these communities also represent an integral part of the nitrogen

cycle. The crusts were found to have 15N isotope ratios between 0 and 1.5,
indicating a high level of N fixation. We hypothesized that this fixed N
would be released into the surrounding soils where it would be taken up
by annual plants growing within a certain proximity. Stable isotopes confirmed soil crusts as the dominant source of N for surrounding plants and
soil. Though hypolithic soil crusts remain a somewhat obscure and unapparent part of the landscape, further exploration of their relative importance
will inspire appreciation of the fundamental part they play in hyperarid
ecosystems.
HEDIN, LARS O. Princeton University, Princeton, NJ. Global signatures
in plant-nutrient interactions: Implications for terrestrial ecosystems.
Alfred Redfields observation of global-scale imprints of biology on stoichiometric ratios between carbon, nitrogen, phosphorus and other nutrients
has been fundamentally important for our understanding marine ecosystems. In contrast, our knowledge of terrestrial plant-nutrient interactions
has historically been more locally focused, resulting in only limited understanding of whether analogous large-scale stoichiometric imprints
emerge at scales of ecosystems, biomes, or even globally. I will here discuss evidence and possibilities for such large-scale rules in how terrestrial
biological processes act to link carbon, nitrogen and phosphorus cycles over
ecological and evolutionary time. I will discuss evidence (and lack thereof)
of biological imprints on nutrient ratios in biomass, soils and nutrient losses
across ecosystems, and whether such patterns may reflect a form of "diffuse" coevolution between nutrient cycles and plant physiological strategies. I will contrast two different, and in some ways alternative, approaches
for further progress: the traditional ecosystem "black-box" approach vs. a
scaling-based approach rooted in physiological constraints at the level of
individuals and/or functional groups.
HEDRICK, VICKI J.* and PAUL BRUNKOW. Southern Illinois University-Edwardsville, Edwardsville, Illinois, USA. Effect of human intrusion
on bird behavior in an Illinois nature preserve.
Increasing outdoor recreation in forested landscapes can affect forest-dwelling birds. It can lead to or increase existing forest fragmentation as camping
facilities, picnic facilities, and parking areas subdivide forested habitat.
Less accessible locations become reachable as trails, designed to channel
human activity, are chiseled into forests. Trail use by humans may cause
birds to alter foraging and nesting behavior by fleeing. This effect of trails
can also extend beyond the immediate trail edge as birds located away
from the edge still flee or seek a more secure position. This study seeks to
evaluate the area of influence of trails in a nature preserve in central Illinois. Visibility of a bird engaged in normal activity was the criteria used
to select a bird for this analysis. An intrusion consisted of one or several
humans walking along the trail. Initial trail distance (the closest perpendicular distance of the bird to the trail prior to disturbance) and its initial
reaction distance (the distance from the intruder at which the bird first
reacted as noted by a sudden change in behavior) were recorded. We computed the ratio of initial reaction distance divided by initial trail distance
(an index of response sensitivity) and correlated that with initial trail distance. Results to date show the relationship between response sensitivity
and initial trail distance to be significantly negative. This relationship was
also highly curvilinear with response sensitivity declining very rapidly with
initial trail distance. The farther the bird from the trail edge, the closer it
allowed a human to approach along the trail before reacting. Our technique
can be applied to quantifying the area of influence of a trail with respect
to species-specific responses, habitat types, and seasonal sensitivity to human disturbance. As such, it may be an important conservation tool for
trail system design and for preserve or park management
HEIMAN, KIMBERLY W.* and FIORENZA MICHELI. Hopkins Marine
Station of, Stanford University, Pacific Grove, CA, USA. The impacts of
the habitat-forming invasive tubeworm, Ficopomatus enigmaticus, in a
heavily-invaded central California estuary.
The number of invasive species in coastal marine environments is on the
rise. Many of these invasives can threaten biodiversity and alter ecosystem
function through competition, predation, habitat modification, or by changing the availability of limited resources. Elkhorn Slough is a central Cali-

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145

fornia estuary with very little natural hard substrate, its native intertidal
habitat being mudflats. In 1994 the reef-building tubeworm, Ficopomatus
enigmaticus, was discovered in the system. Over the past 8 years, F. enigmaticus has spread to several sites within Elkhorn Slough. At one site it
occupies nearly 100% of the available substrate forming reefs that grow
out from dock pilings and spread over the surrounding mudflats. The reefs
greatly increase the amount of complex hard substrate and create a new
unique habitat type. We explored the role of the new complex structure, F.
enigmaticus reefs, on the invertebrate community composition in surrounding mudflats. Replicate reefs were removed in an experiment designed to
identify their impacts on infaunal communities under the reefs, as well as
those 5cm and 100cm away from the edge of the reefs. Initial multivariate
and univariate analyses showed that there is a significant effect of reef
removal on infaunal mudflat communities 6 months after the removal. Differences in the abundances of several invasive species including the amphipods Monocorophium insidiosum, Grandidierella japonica, and the
cryptogenic polychaete Capitella capitata drive the observed community
effects of reef removal.
HEIN, CATHERINE L., M J. VANDER ZANDEN, JOHN J. MAGNUSON and BRIAN M. ROTH. Center for Limnology, University of Wisconsin-Madison, Madison, WI, USA. The removal of rusty crayfish from
Sparkling Lake, WI.
The rusty crayfish (Orconectes rusticus), native to the Ohio River Valley,
has become a problematic invader throughout many areas of the U.S. and
Canada, including northern Wisconsin. It replaces congeners, reduces the
abundance of macroinvertebrates and macrophytes, and has negative effects
on the recruitment of sport fishes. While prevention of further dispersal is
most effective, rusty crayfish have already become established in many
lakes. We are attempting to eradicate rusty crayfish from Sparkling Lake,
Vilas County, Wisconsin through intensive removal with traps and the implementation of trophy fishing regulations on important crayfish predators.
This talk will focus on the trapping aspect of the removal effort. In August
of 2001, we removed approximately 11,000 crayfish over two weeks. We
further reduced the population by removing 38,500 crayfish over 40 days
in the summer of 2002. Catch rates declined with removal, but also changed
due to temporal differences in behavior during the summer. The ratio of
females to males in traps increased from 2001 to 2002, perhaps due to
competitive interactions occurring outside traps. We found no change in
the average carapace length of crayfish caught during the removal. Few
attempts at removing crayfish from natural systems without using biocides
have been made, and prior attempts have failed. By combining an increase
in predation with intensive removal, we may be able to push the population
to collapse through depensatory dynamics.
HEISLER, JANA L.* and JOHN M. BRIGGS. Arizona State University,
Tempe, AZ 85287-1601, USA. An ecosystem in transition: What are
the mechanisms driving shrub expansion in temperate grasslands?
Fire is an integral component of the disturbance regime in temperate grasslands and impacts the native grass and shrub species through both its direct
and indirect effects. During the last century, fire suppression in these grasslands has been accompanied by an increase in shrub cover. While this shift
in growth form dominance has been well documented, short-term mechanistic studies are integral in understanding the mechanism driving this transition from grassland to shrubland. In 2001, an experimental study was
initiated at Konza Prairie to assess the ways in which fire acts as a constraint to and/or facilitator of shrub expansion. Briefly, the experimental
design included fire, nitrogen addition, and litter as main effects and pairwise comparisons between shrub islands allowed us to uncouple and subsequently quantify the impact of the fire event itself from that of the postfire microclimate. In shrub islands (species Cornus drummondii) exposed
to spring fire, 100% aboveground mortality was observed, but by ca. 60
days post-fire, a pulse of sprouting enabled stem density to return to preburn levels. Light availability, and subsequently warmer soil temperatures,
further stimulated this sprouting response with burned islands and burned
+ litter islands increasing in stem density by six-fold and four-fold, respectively. By comparison, unburned islands (controls) increased by only
two-fold. While ANPP for C. drummondii in all islands was similar, resource allocation differed between burned and unburned islands, with new

146

Abstracts

sprouts investing in stem tissue to rapidly regain height and pre-burn stems
producing greater foliar tissue. These results, when paired with a recent
long-term study of fire frequency and patterns of shrub expansion, suggest
that fire events stimulate rapid increases in C. drummondii stem density,
which may lead to faster rates of shrub expansion, especially when followed by fire-free intervals during which shrubs may increase in total cover.
HELLQUIST, C. E.,* S. NELSON, B. RHODES, R. W. LEE and R. A.
BLACK. Washington State University, Pullman, WA. Is the invasive
cordgrass, Spartina anglica, becoming integrated within invertebrate
food webs in north Puget Sound, WA, USA?
Spartina anglica is an invasive estuarine cordgrass that has colonized approximately 400 ha of intertidal habitats in north Puget Sound, WA. This
invasion has resulted in the extensive colonization of some mudflats while
other similar mudflats may have little or no Spartina present. This distribution pattern has created opportunities to examine trophic relationships
following colonization of a non-native species. We examined estuarine trophic relationships by sampling the stable isotopic composition (d13C, d15N,
and d34S) of phytoplankton, zooplankton, benthic invertebrates, macroalgae,
and vascular plants. Initial data using d15N and d13C indicate that invertebrate isotopic ratios vary between sites and species. Macoma balthica (Bivalvia) collected within roots of S. anglica has a C signature (d13C -16)
that is relatively similar to S. anglica (d13C -13). In bivariate isotopic
scatterplots with d15N, Macoma balthica is placed closer to Spartina than
it is to other bivalves. This placement may indicate the use of some Spartina-derived carbon. However, Macoma nasuta (Bivalvia) and Mytilus edulis (Bivalvia) collected among Spartina have coinciding isotopic signatures (d13C -21; d15N 9-10) that are different from those of Spartina (d13C
-13; d15N). Isotopic signatures of Macoma nasuta and Mytilus for d15N
more closely resemble published values for phytoplankton (ca. d13C -21;
d15N 9-10). These data suggest that despite living among the roots of
small Spartina clones (Macoma nasuta) or living attached to the base of
Spartina stems (Mytilus), these bivalves are not using Spartina productivity
for nutrition.
HELMS, BRIAN S.,1,* JOHN W. FEMINELLA1 and PHILIP L. CHANEY.2 1 Department of Biological Sciences, Auburn University, Auburn,
Alabama, USA; 2 Department of Geology and Geography, Auburn University, Auburn, Alabama, USA. Land use and stream biotic signatures:
The effects of urbanization on periphyton, macroinvertebrates, and fish
in watersheds of west Georgia.
In 2002 we began a long-term study designed to examine the relationship
between changes in land use associated with urbanization and stream communities. Twenty small watersheds (500-2500 ha) were selected along an
urbanization gradient in the piedmont ecoregion north of Columbus, Georgia. Based on GIS data from 2002 Landsat 7 TM imagery, study streams
drained watersheds that included largely urban, developing (suburban), agricultural (pasture), managed pine forest, and unmanaged mixed forest land
uses. A representative 100-m reach including 3 separate run-pool segments
was sampled seasonally for periphyton (attached algae and diatoms, quantified with artificial and natural substrates), benthic macroinvertebrates
(Surber samplers), and fish (electrofishing and seining) in each watershed.
Preliminary data indicated that abundance of tolerant fish species (e.g.,
Lepomis cyanellus, Gambusia affinis, Semotilus atromaculatus, and Ameirus spp.) were higher in urban and or developing watersheds than in forested watersheds. In contrast, numbers of macroinvertebrate species in the
aquatic insect orders Ephemeroptera, Plecoptera, and Trichoptera (EPT) and
algal biomass (as chlorophyll a) were higher in mixed-forested watersheds
than in urban watersheds. Taken together, these data suggest that particular
biological signatures may be reliable indicators of human-induced disturbance at the landscape scale, which in turn may be useful in forecasting
the magnitude of whole-system changes in stream structure and function
associated with the conversion of forests to urban/suburban land in the
Southeast.

HELMS, SANDRA E.,* SCOTT J. CONNELLY and MARK D. HUNTER.

Institute of Ecology, University of Georgia, Athens, GA, USA. Effects of
variation in plant quality on the interaction between an herbivore and
its parasitoid.
Previous studies have demonstrated that phenotypic traits of plants have
the potential to affect interactions between herbivores and their natural
enemies. Consequently, the impact of natural enemies on herbivore vital
rates and population dynamics may vary among plant species. Our study
was designed to investigate the potential for density-dependent parasitism
of an aphid herbivore feeding on six different host plant species. Population
densities of the aphid Aphis nerii and its parasitoid Lysiphlebus testaceipes
were recorded within a single growing season on six different species of
milkweed in the genus Asclepias. Asclepias species are known to vary in
their quality as food for herbivores. Although data on plant quality were
not available in this study, population data were analyzed to determine the
effects of different Asclepias species on rates of parasitism and aphid population growth. Parasitism rates of A. nerii varied among Asclepias species
but were temporally density dependent over at least some range of aphid
density on all plant species. Aphid population growth rates also varied
among Asclepias species, and declined with an increase in the maximum
parasitism rates among plant species. However, in no case was densitydependent parasitism sufficient to prevent exponential population growth
of aphids within the growing season. The results serve to emphasize that,
if natural enemies are to regulate herbivore populations, density-dependent
mortality is a necessary, but not sufficient, condition for regulation.
HELMUS, MATTHEW R.1,* and DAVID E. DUSSOURD.2 1 University of
Wisconsin-Madison, Madison, WI; 2 University of Central Arkansas, Conway, AR. The vein-cutting monarch: a bioassay for vein-cutting stimulants in milkweed latex.
Larvae of the Monarch butterfly (Danaus plexippus) eat milkweeds (Asclepiadaceae), plants that when punctured exude a sticky latex. Monarchs
sequester milkweed cardenolides, which are toxic to most animals and occur at high concentration in milkweed latex. Paradoxically, monarchs have
evolved behaviors that circumvent latex and thus diminish their defensive
potential. Before eating, final instar larvae cut the latex canals of a leaf
and feed distal to the cuts where latex no longer flows. Although vein
cutting clearly functions to reduce larval contact with latex, it is not known
what characteristic of latex stimulates this behavior. We have developed a
novel bioassay to test for vein-cutting stimulants found in milkweed latex.
When given pure latex or a solution containing a vein-cutting stimulant, a
feeding monarch will turn and rechew its previous vein-cuts. All larvae
tested with fresh latex rechewed their vein cuts, while larvae tested with a
sticky polyethylene glycol solution or a solution of cardenolides did not.
The supernatant and pellet of centrifuged latex were tested, and the supernatant found to be active. Latex supernatant was dried and washed with
methylene chloride (a solvent for cardenolides) and methanol, with the
residual dissolved in water. The methanol fraction only, elicited rechewing.
Presence for cardenolides was tested in the methanol fraction using a
2,2,4,4-tetranitrodiphenyl base catalyzed reaction in two methanol fractions. One fraction tested negative while the second showed slight traces(;0.05mg/ml). Activity was also maintained in a methanol fraction inoculated with a general proteinase suggesting the stimulant(s) is not proteinaceous. These initial results suggest a chemical cue(s) for vein cutting
in monarch larvae. Furthermore, since over 100 species of Lepidoptera,
Coleoptera and Orthoptera also cut veins on host plants with secretory
canals; these test methods (i.e., vein cutting, solution application and rechewing) could be used to characterize vein-cutting stimulants for other
taxon.
HELMUTH, BRIAN* and JENNIFER JOST. University of South Carolina,
Columbia, SC. Living on the edge: Biophysical and biomechanical consequences of body size to intertidal invertebrates.
Coastal environments are among the most physically challenging ecosystems on earth. Forces exerted by crashing waves regularly exceed those
imparted by hurricane force winds on land, and fluctuations in body temperature of 208C or more are not at all uncommon. While researchers have
made significant inroads into understanding the consequences of body size

and morphology to the frequency of dislodgement and breakage of intertidal invertebrates and algae, only recently have biophysical models been
used to unravel similar impacts of body size and shape on intertidal heat
flux. We therefore have only a rudimentary understanding of what spatial
and temporal patterns of body temperature are in nature. Compared to work
in terrestrial habitats, the exploration of how biomechanical and biophysical
responses of individuals to environmental forcing scales to the level of
populations and communities is still relatively unexplored in intertidal environments. Nonetheless, the unique and often severe environmental conditions in this habitat provide important lessons for how the consequences
of body size and morphology change in counterintuitive and often highly
nonlinear ways in both space and time. We present a series of models and
measurements which explore the consequences of body size to thermal
stress of intertidal mussels (Mytilus spp. and Geukensia demissa). Size has
little effect on mussel temperature in soft sediment environments, due to
the coupling between organism and sediment temperatures. In contrast, the
body temperatures of rocky intertidal mussels vary markedly with size, but
the relative effect changes with the duration and timing of exposure at low
tide. As a result, the impact of body size likely varies with intertidal zonation height and with latitude. Similarly, latitudinal-scale patterns of body
temperatures (and hence levels of thermal stress) are species-dependent,
and we discuss the implications of these differences to Environmental
Stress Models.
HEMBRE, LEIF K.,1,2,* LAURA A. PETERSON,1 EMILY WALSH,1 REBECCA FORMAN2 and DAN R. ENGSTROM.3 1 Hamline University, St.
Paul, MN, USA; 2 University of Minnesota, St. Paul, MN, USA; 3 St. Croix
Watershed Research Station, Science Museum of Minnesota, Marine on St.
Croix, MN, USA. Evolution of negative phototaxis in a Daphnia population revealed by reanimation of resting eggs.
The diapausing sexual eggs (ephippia) of cladocerans (e.g., Daphnia) deposited in sediments represent biological time capsules that provide a record of the evolutionary history of populations. This study examines the
egg bank of a D. pulicaria population in a lake that has been stocked
annually with rainbow trout (a predator of Daphnia) for 40 years. Ephippia
from sediments dating from the early 1900s to 2001 were hatched, and
clonal cultures were established from the hatchlings. To evaluate whether
the population had evolved over the past 40 years in response to elevated
predation levels, we examined the phototactic behavior (a proxy for diel
vertical migration (DVM) behavior) of clones in the presence and absence
of fish kairomones. DVM is widely accepted to be a predator-avoidance
behavior. Individuals migrate to depth during the daytime to avoid predation by visual predators (e.g., fish), and then typically ascend into surface
waters at night when there is a lower risk of predation. Chemicals exuded
from predators (kairomones) provide a proximate cue that influences the
DVM behavior of individuals. Therefore, we expected that 1) the high
levels of predation over the past 40 years would have selected for more
negatively phototactic clones, and 2) kairomones would influence the
strength of the behavioral response. Our results from assays using kairomone-free water indicate that clones hatched from ephippia produced after
the trout stocking program began (1961) were indeed significantly more
negatively phototactic than those produced before 1961. Interestingly, in
the presence of kairomones, clones from both the pre-and post-stocking
eras became more negatively phototactic, but the response was only significant for clones from the pre-stocking era. This suggests that before trout
were introduced to this lake clones in the population had considerable
plasticity in their DVM behavior. Their ancestors, however, appear to have
evolved more conservative DVM strategies.
HEMMER, MICHAEL J.,1 NANCY D. DENSLOW,2 PATRICK LARKIN,2,3 REBECCA L. HEMMER,1 IRIS KNOEBL2 and PEGGY HARRIS.1
1
U.S. Environmental Protection Agency, NHEERL, Gulf Ecology Division, Gulf Breeze, Florida, USA; 2 University of Florida, Gainesville, Florida, USA; 3 EcoArray LLC, Alachua, Florida, USA. Field application of
a sheepshead minnow estrogen responsive cDNA macroarray.
Preliminary experiments with the sheepshead minnow (Cyprinodon variegatus) have revealed at least 30 genes which are up-regulated by estrogen
treatments. Identical patterns of gene up-regulation have been observed for
the native ligand estradiol and the pharmaceutical estrogens ethynyl estra-

Abstracts

147

diol and diethystilbestrol as well as the xenoestrogens, methoxychlor and

nonylphenol. Sheepshead minnow cDNA libraries were prepared and
probed to identify full length transcripts which were used to develop a
estrogen responsive cDNA macroarray. An initial validation of the membrane macroarray was accomplished by measuring the hepatic expression
of these genes in male sheepshead minnows using fish previously exposed
in laboratory studies to the above native, pharmaceutical and xenoestrogens. In a second study, adult male sheepshead minnows were collected
from six sites throughout the Pensacola (Florida) Bay system. Collections
occurred over the course of two years,corresponding to the normal reproductive season for this species. These sites receive a variety of urban and
industrial chemicals and definitive results have shown that male, sheepshead minnows captured at several of these locations contained high plasma
levels of vitellogenin indicating exposure to either natural and /or xenoestrogenic chemicals. The sensitivity and reproducibility of the cDNA membrane array was assessed by correlating results of the macroarray with
previously measured plasma vitellogenin expression for individual fish
sampled from field sites. The degree to which alterations in gene regulation
in field-collected organisms correspond to those observed in animals exposed to known estrogenic substances in the laboratory was used as a basis
to further assess the utility of gene arrays as indicators of exposure and
effects to environmental contaminants. This is the first step in developing
a suite of specific macroarrays to monitor the status of aquatic environments.
HENEBRY, GEOFFREY M.1,* and DOUGLAS G. GOODIN.2 1 University
of Nebraska, Lincoln, NE; 2 Kansas State University, Manhattan, KS. Land
surface phenology, ecoregional expectations, and change analysis: Toward ecological forecasting with image time series.
To make ecological forecasting an operational possibility, we need the capability to establish and to update complex spatio-temporal baselines that
will enable prediction of the usual and identification, quantification, and
assessment of the unusual. For the analysis of image time series, it is
necessary to move beyond viewpoints that center on pixels or patches and
characterize the dynamic of pattern and process that sequences portray. The
phenology of the land surface provides a basic ecological phenomenon
fruitful for comparative analyses. The USGS CONUS AVHRR maximum
NDVI biweekly composites for 1990-2000 were partitioned by Omerniks
84 Level III ecoregions. For each ecoregion at each composite date, a
random walk resampling procedure was used to estimate measures of spatial dependence, spatial heterogeneity, and aspatial intensity. Expectations
of land surface phenology were calculated for each ecoregion. Comparisons
of ecoregional expectations reveal broad patterns in the relationships between spatial dependence, spatial heterogeneity, and aspatial intensity.
Analysis of seasonal trajectories in spatial structure reveals time periods
(watchpoints) with high convergence or divergence in interannual variation.
Patterns of these watchpoints provide a basis for forecasting anomalous
land surface behavior.
HENNE, PAUL D.* and FENG SHENG HU. University of Illinois at Urbana-Champaign, 265 Morrill Hall, 505 S Goodwin Ave, Urbana, IL,
60801. Holocene ecosystem change in the Great Lakes snow belt: Initial results from Huffman Lake.
Lake-effect snow increases annual moisture in a consistent geographic pattern to areas downwind from the Great Lakes. Today, vegetation, fire frequency, and soil weathering in the Great Lakes region exhibit patterns
related to the average snowfall. We are studying pollen, macroscopic charcoal, and oxygen isotopes of carbonate in sediment cores from lakes located
within and outside the snowbelt in northwestern Lower Michigan to examine the influence of lake-effect snow on Holocene ecosystem development. Preliminary pollen and charcoal data from Huffman Lake (45808 N,
84846 W) show that during the early Holocene, surrounding forests were
dominated by Picea, Pinus, and Betula, and had short fire return intervals.
Forests dominated by Pinus and Quercus were prevalent during the middle
Holocene. Fire frequency was highest near the early to middle-Holocene
transition, and decreased markedly within the middle Holocene. Around
6000 years BP, mesic species including Acer, Fagus, Tsuga, Betula, Ulmus,
and Tilia became abundant, and the sedimentary organic content rose greatly. Fire return intervals remained similar to those of the middle Holocene

148

Abstracts

until a late-Holocene decline. The onset of mesic species dominance probably resulted from increased lake-effect snow, as postulated by previous
studies. These preliminary results will be verified with oxygen-isotopic data
from the same site, and through comparison with paleoecological and oxygen-isotopic data from a site outside the Great Lakes snow belt.
HENRY, HUGH A.L.* Department of Botany, University of Toronto, Toronto, Ontario, Canada. Herbivores as opportunists: migratory connectivity and large scale disturbance in Arctic coastal ecosystems.
In recent decades, migratory geese that breed in the Arctic, but overwinter
in temperate regions of Northwestern Europe and North America, have
shown dramatic increases in numbers that appear to be linked to their
increasing use of agricultural crops as a food resource. The foraging behaviour of the mid-continent population of lesser snow geese is an example
of this change in resource acquisition that has led to a relaxation of the
density-dependent regulation which occurred earlier in their traditional wintering area, the coastal marshes of the Gulf States. The agricultural nutrient
subsidy has led to strong top-down effects in the coastal marshes of the
Hudson Bay Lowlands and at other locations where the increased numbers
of birds breed - a consequence of migratory connectivity. Direct and indirect biotic and abiotic processes mediated by feedback mechanisms and
initiated by goose foraging have led to sustained change resulting in the
development of alternative stable states. These non-linear changes, that
display threshold responses, include loss of vegetation, irreversible changes
in soil properties and a decline in microbial activity, and decreases in invertebrate and passerine species. The coastal systems display low resilience, and the coalesence of local disturbed areas has led to a highly fragmented landscape where there is a marked temporal asymmetry between a
possible decline in goose numbers and revegetation of coastal areas. The
effects of increased numbers of geese on nitrogen dynamics in particular
may alter drastically the productivity and stability of these systems. In a
model of the flows of nitrogen of a grazed lawn in an Arctic coastal marsh,
the alternative stable states of the system are obtained with values for the
state variables that are consistent with field data. The determining factor in
the transition to an alternative stable state is the loss of the input of nitrogen
from fixation when lawn area is reduced by geese.
HERENDEEN, ROBERT A.* and WALTER R. HILL. Illinois Natural History Survey, Champaign, IL. Growth dilution in multilevel food chains.
Microalgae absorb toxicants from the aqueous environment, passing them
along to higher trophic level. Rapid growth of microalgae (stimulated by
increased light, for example) results in lowered tissue concentration of toxicant. Reductions in autotroph toxicant concentration have been observed
to propagate to herbivores. Here we investigate (with simulation and analytically) the propagation of growth dilution to all levels of a food chain.
We are concerned with concentration as well as overall mass of toxicant
in each level, for different functional relationships between levels. We find
a wide variation in how growth dilution progresses up the chain, depending
on functional relationship (e.g., ratio vs. prey dependence). For example,
transient growth dilution can be followed by steady-state growth concentration. These results, which have implications for pollution control, call
for experimental testing.
HERRICK, JEFFREY E.,1,* DAVID A. PYKE,2 MIKE PELLANT3 and
PATRICK SHAVER.4 1 jherrick@nmsu.edu, USDA-ARS Jornada Experimental Range, Las Cruces, NM; 2 USGS Forest & Rangeland Ecosystem
Science Center, Corvallis, OR; 3 BLM Idaho State Office, Boise, ID;
4
USDA-NRCS Grazing Lands Technology Institute, Corvallis, OR. A new
standard for qualitative rangeland health assessments.
The term rangeland health is increasingly used by land managers to refer
to the maintenance of complex ecological processes on rangelands. In an
attempt to provide a rapid, moment-in-time assessment of rangeland health,
we developed the Interpreting Indicators of Rangeland Health protocol.
This protocol uses 17 observable indicators to assess 3 attributes of rangeland health: soil and site stability, hydrologic functioning and biotic integrity. It is now being widely applied throughout the United States and Mexico. In the initial published version (3.0), the standard for assessing an
evaluation area was the NRCS Ecological Site Description, a description

of physical and biotic characteristics of land with similar soils and climate
and that respond similarly to management. Multiple reference areas provided visual representations of the ecological site and were used to supplement Ecological Site Descriptions or as the standard when Ecological
Site Descriptions were not available. This approach failed to integrate all
of the diverse sources of information about rangeland ecological processes.
In response, we developed an ecological reference sheet for version 4.0. A
reference sheet is developed for each ecological site based on the integration of ecological site descriptions, expert knowledge, scientific literature
and reference sites. Each sheet should be developed by a group of experts
familiar with the range of variation in the soil, hydrologic and vegetation
indicators. When possible, data similar to those found in ecological site
descriptions should be used to generate a quantitative range of expected
variation. We believe this change will increase the quality and consistency
of evaluations completed with this protocol.
HERRON, PATRICK M.,1,* DANIEL J. GAGE2 and ZOE G. CARDON.1
1
Department of Ecology and Evolutionary Biology, University of Connecticut, Storrs, CT; 2 Department of Molecular and Cell Biology, University of Connecticut, Storrs, CT. Divining Rods: Pseudomonas putida as
a microbiosensor of fine-scale osmotic potentials in soil.
The physical distribution and availability of water in soil influences plant
growth, the mineralization of organic matter, the diffusion of dissolved
nutrients and microbial dynamics. Current tools commonly used to measure
water availability in soil, such as psychrometers, tensiometers and time
domain reflectometry, integrate water availability on a gross scale but do
not provide information at microscopic scales where microbes are operating. We have inserted an osmotically controlled proU-GFP transcriptional
fusion developed by Axtell and Beattie (Appl. Env. Microbiol. Vol. 68:9,
pp 4604-4612. 2002) into the soil bacterium Pseudomonas putida KT2440.
The resulting soil microbial biosensor produces green fluorescent protein
(GFP) as a function of osmotic potential around the bacterium. GFP fluorescence from the microbiosensors can be detected in nonsterile soils nondestructively using epifluorescence microscopy, and thus provides fine
scale information on an important determinant of water potential in the soil
microbial environment. These microbiosensors promise to provide a novel
portrait of dynamics of rhizosphere osmotic potential associated with root
water uptake.
HIAASEN, BARBARA A.,* JOHN C. VOLIN and STEVEN P. SIMMONS. Florida Atlantic University, Davie, FL. The effect of water velocity on the periphyton taxonomic composition of mesocosms in the
Everglades.
Numerous studies conducted within lotic systems have shown that the
structure and composition of freshwater periphyton communities are sensitive to many variables, including phosphorus availability and water velocity. Phosphorus uptake by periphyton is influenced by the metabolic
characteristics of the component algal species. Differences in phosphorus
uptake at different water velocities could thus be mediated by differences
in periphyton composition. The primary objective of this research was to
investigate the effects of water velocity on taxonomic composition of periphyton communities growing within artificial mesocosms at flow rates
representative of Everglades marsh communities. These mesocosms received water from Stormwater Treatment Areas (STAs) constructed in the
Everglades to serve as filter marshes for wastewater with high nutrient
loads. A unique mesocosm design was utilized to control velocity, while
maintaining equal hydraulic loading rates, residence times and colonizing
surface between velocity treatments. The slow and fast water velocity treatments were 0.22 cm s-1, and 2.0 cm s-1, respectively. A comparative taxonomic analysis of genera using DIC and epi-fluorescence light microscopy
was performed to examine periphyton relative abundance, generic diversity,
and taxonomic differences within and between treatments. Periphyton tissue phosphorus and nitrogen levels, as well as numerous other metrics
including dry weight biomass and water column phosphorus, nitrogen, pH
and temperature, were determined. Analysis of initial samples has revealed
clear differences in periphyton composition, particularly with regard to the
relative proportion of diatomic and blue-green algal species present. Studying the effects of water velocity on periphyton communities provides data
that can be used to optimize the design of STAs and increase the efficiency
of nutrient removal technologies used in Everglades restoration.

HIGGINS, PAUL A.1,2 1 Stanford University, Stanford, CA; 2 Global

Change Education Program, Department of Energy, Oak Ridge, TN. Potential biological diversity loss due to abrupt climate change.
Occurrence of abrupt climate change, such as could occur under thermohaline circulation (THC) weakening, is increasingly evident in the paleoclimate record and model projections of future climate. The climate changes that accompany forced weakening of THC, as projected by HadCM3,
cause large changes in potential ecosystem structure and function in northern and eastern South America. Precipitation increases in eastern South
America cause a shift from drier habitat types characterized by relatively
low levels of biological diversity to wetter habitat types characterized by
high levels of biological diversity. In contrast, vegetation in northern South
America shifts from wetter habitat types with higher diversity to drier habitat types with lower diversity. These opposing biotic responses to the climate change occur in regions characterized by vastly different levels of
human population density and land use. Population density in eastern South
America is high with a concomitant degradation in habitat quality. In contrast, the vegetation shift in northern South America occurs where the human population density is low and where habitat quality is high. As a
consequence, the high diversity ecosystem types move from areas of high
habitat quality to areas characterized by heavy disturbance and large biodiversity losses occur.
HILDREW, ALAN1,* and GUY WOODWARD.1,2 1 School of Biological
Sciences, Queen Mary, University of London, Mile End Road, London,
UK, United Kingdom; 2 Department of Zoology and Animal Ecology, University College Cork, Lee Maltings, Prospect Row, Cork, Ireland, Ireland.
Strong density dependence in a stream insect: large scale manipulations.
Recruitment of the abundant, predatory alderfly Sialis fuliginosa was manipulated over three years in a well known southern English stream. Egg
masses, that are laid on the leaves of trees overhanging the stream, were
removed (.90% efficiency), left as a control or added to 150m stretches
of stream. The effects were marked at first but transitory. Strong density
dependence had homogenised densities among the treatments within 4 - 12
months of the manipulation in each year of the experiment. The source of
this density dependence was starvation and/or intraguild predation (including cannibalism) acting on the very earliest stages. Strong density dependence is rarely reported in stream communities. Broadstone Stream has a
highly reticulate, donor-controlled food web and density dependence
among the top predators, such as Sialis, could be crucial for the stability
of the web.
HILL, JUDSON P.,1,* MATTHEW J. GERMINO,1 JON M. WRAITH2 and
BRET E. OLSON.3 1 Idaho State University, Department of Biological Sciences, Pocatello, ID; 2 Montana State University, Department of Land Resources & Environmental Sciences, Bozeman, MT; 3 Montana State University, Department of Animal & Range Sciences, Bozeman, MT. Differences in water use and photosynthetic gas exchange in Centaurea maculosa and competitors.
Whether interspecific, ecophysiological differences in water uptake and water use influence community invasibility is not well known for semiarid
rangelands of the Western US. Previous research showed that water contents were lower in soils of SW Montana grassland communities containing
spotted knapweed (Centaurea maculosa; CEMA) one of the most problematic invasive species of the northwestern region. The purpose of this research was to determine if interspecific differences in water status and
photosynthetic gas exchange exist between CEMA and several co-occurring, established species: Pseudoregneria spicata (PSSP), Agropyron smithii, and Bromus inermis (BRIN). Microclimate, plant water potential (C),
and photosynthesis were measured for naturally occurring plants, and for
plants grown under varying soil water availabilities in a greenhouse. No
differences in photosynthesis or water use efficiency (WUE) were detected
between CEMA and natives in the greenhouse during 4 days of progressive
drying. However, photosynthesis of CEMA was 40-60% greater than BRIN
and PSSP in the field, but it had lower WUE, which agreed with previous
estimates based on carbon isotopes. CEMA also maintained 85% greater
C than BRIN during early and mid summer, and moreover had persistent

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149

photosynthesis in August and September when nearly all other grasses and
forbs had become dormant. These ecophysiological differences may reflect
greater soil water extraction in CEMA, possibly through dual use of shallow roots and taproots, which could contribute to its competitiveness under
water-limited conditions.
HILL, RONALD L.* NOAA/NMFS/Southeast Fisheries Science Center,
ron.hill@noaa.gov, Galveston, TX. The influence of conspecific and heterospecific juveniles on benthic recruitment in a reef-fish.
Although newly-settled reef fish juveniles demonstrate heterogeneous distributions, little progress has been made in discerning interactions or cues
that create these differences. Previous hypotheses and studies have assumed
or demonstrated negative interactions, such as competition, between residents and newly settling juveniles, with a few notable exceptions. Habitat
selection by newly-settled white grunts, Haemulon plumieri, was tested
using experimental manipulation of similar settlement sites: a) Acropora
cervicornis with small juveniles, b) unoccupied A. cervicornis colonies, c)
A. cervicornis from which residents had been recently removed, and d)
coral colonies occupied by older juveniles. Out of 1496 white grunts, significantly more (85.7%) recruited to coral colonies with early stage juveniles than to (b) 4.5%, (c) 8.2%, or (d) 0% (ANOVA, P,0.0001). Recruitment of all newly settled haemulids (4610 total) followed the same pattern:
87.4% to A. cervicornis with young resident juveniles. These results support the hypothesis that settlement of white grunts and other haemulids is
facilitated rather than inhibited by the presence of earlier settlers. The conspecific- and heterospecific-attraction hypotheses predict advantages to
schooling fish recruiting together based on mutually beneficial behaviors
and habitat cues.
HILLE RIS LAMBERS, J.,* W.S. HARPOLE and D. TILMAN. University
of Minnesota, St. Paul, MN. The diversity-productivity relationship:
Putting species back into the equation.
The mechanisms causing greater productivity at higher diversity have been
hotly debated. We use data from the longest running biodiversity experiment to test two hypotheses. First, species-specific resource use (niches) or
positive interactions among species may result in greater productivity at
higher diversity (a complementarity effect). Alternatively, diverse mixtures
could be more likely to include productive species, thus causing greater
productivity (a sampling effect). Studies of the productivity-diversity relationship have focused on the response of communities, not species, to
increasing diversity, making it difficult to distinguish between these two
hypotheses. We develop analyses to 1. identify species that overyield, and
2. determine which species characteristics are associated with overyielding.
We found that 6 of 17 species significantly overyield, i.e. have greater
productivity when grown in mixture than expected from their monoculture
productivity. Overyielding species are not the most productive species in
monoculture, implying that a positive sampling effect does not explain
increased productivity in diverse plots. The degree to which species overyield is correlated with their ability to compete for nitrogen in these nitrogen-limited grasslands, suggesting that resource use plays a role in determining whether species overyield in diverse mixtures. Greater than expected productivity of the six overyielding species far outweighs the lower
than expected productivity of four underyielding species, thus, resulting in
the positive relationship between diversity and productivity.
HILLHOUSE, HEIDI L.* and JOHANNES M.H. KNOPS. University of
Nebraska-Lincoln, Lincoln, Ne. Tissue fraction and N content controls
on leaf litter decomposition.
N and lignin content have long been studied as predictors of mass loss in
decomposing plant litter, and N content has been analyzed as a predictor
for changes in N concentration during decomposition. Previous research
has demonstrated that both of these factors are critical to understand N
release and cycling, but most studies have focused on either tissue fractions
or N content, rather than a combination of the two. To address this issue,
we analyzed newly senesced and 1 yr old leaf litter from 25 species of
grassland and woodland plants in an attempt to better understand the dynamics between initial tissue distribution, initial N concentration, % mass
loss, 1 year N concentration, and 1 year change in N. Each tissue sample

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was separately analyzed for mass loss, tissue fraction distribution, and total
N content. Species varied widely in initial tissue fraction distribution, initial
N content (0.2%-2.5%), mass lost after 1 year (37%-80%), and 1-year
change in total N (42%-137%). Note that, although we refer to the tissue
fractions by their predominant biochemical components (soluble, hemicellulose, cellulose and lignin), these fractions are heterogeneous and not well
defined from a biochemical point of view. Multiple regression analysis
showed that, while lignin and cellulose fractions explain much of the variation in mass loss (R2 5 .63), initial N content of the tissue is a better
predictor of both 1yr N content (R2 5 .81) and total change in N (R2 5
.40). These results suggest that, instead of focusing on N or tissue fraction,
it is important to consider both tissue fraction distribution and the initial
N content of the litter.
HINES, JESSICA E.,1,* JAMES P. MEGONIGAL2 and ROBERT F. DENNO.1 1 University of Maryland, Department of Entomology, College Park,
MD, USA; 2 Smithsonian Environmental Research Center, Edgewater, MD,
21037. Linking aboveground and belowground dynamics: Indirect effects of microbe decomposers on plant-herbivore interactions.
Because plants take up nutrients from the soil and allocate them to the
production of leaf tissue, they provide an obvious link between belowground soil nutrient cycling and aboveground interactions between plants
and their associated community of insect herbivores. Despite this logical
connection, very few studies have examined how plants, and the decomposition of their leaf litter, influence soil microbial processes that in turn
alter plant nutrition and the success of aboveground herbivores. To elucidate the indirect effects of leaf litter, as a carbon source for microbes a
possible nitrogen sink, on plant-herbivore interactions, we performed a resource subsidy experiment on an Atlantic coast Spartina marsh and measured the consequences for plant quality and aboveground herbivores,
namely Prokelisia planthoppers. Treatments were achieved by adding carbon (sucrose), nitrogen (NH4NO3), and leaf litter in a completely randomized factorial field plot experiment for two years. We found that nitrogen
addition treatment plots (low C:N) exhibited higher soil nutrient availability, higher plant quality (N content), and higher herbivore abundance. In
contrast carbon addition treatment plots (high C:N- designed to enhance
the microbial community) showed increased dissolved organic nitrogen,
decreased plant quality (plant biomass), and decreased herbivore abundance
compared to non-manipulated controls. Thus, leaf litter has the potential to
contribute to herbivore dynamics on this salt marsh by indirectly altering
the quality of their Spartina host plant by altering microbial nitrogen mineralization. This study emphasizes the importance of linking belowground
ecosystem processes such as nutrient cycling and decomposition with
aboveground food web interactions.
HITOSHI, MIYASAKA,1,* DZYUBA V. YELENA,2 SHUBENKOV G.
SERGEY,3 KHANAYEV V. IGOR,4 NELNIK G. NATALIA,5 GENKAIKATO MOTOMI6 and WADA EITARO.7 1 Center for Marine Environmental stadies, Ehime Univ., Matsuyama, Ehime; 2 Limnological Institute,
Irkutsk, Irkutsk, Russia; 3 Limnological Institute, Irkutsk, Irkutsk, Russia;
4
Limnological Institute, Irkutsk, Irkutsk, Russia; 5 Limnological Institute,
Irkutsk, Irkutsk, Russia; 6 Research Institute for Humanity and Nature, Kyoto, Kyoto, Japan; 7 Research Institute for Humanity and Nature, Kyoto,
Kyoto, Japan. Food habit divergence between two pelagic sculpin species, Comephorus baicalensis and C. dybowski in Lake Baikal.
Stable isotope and diet composition analyses were used to show the comparisons of food habits between two species of common pelagic sculpin,
Comephorus baicalensis and C. dybowski, in the northern and southern
basins of Lake Baikal. The two pelagic sculpins are one of the important
members for shaping the pelagic food web. Carbon and nitrogen stable
isotopes could be assessed that the pelagic food web has an ideal, isotopically ordered structure in past studies. The isotopically simple structure
of pelagic food web, however, has a swelling part in the middle trophic
level which constructed by some pelagic sculpin species, an amphopod
(Macrohectopus branickii) and a zooplankton (Epischura baicalensis). This
middle part of pelagic food web was examined under the two sides, diet
of two pelagic sculpins and food resource of two basins. Both nitrogen and
carbon isotope data indicated that trophic positions of two pelagic sculpins
differed between the species, but not between the basins. In both two ba-

sins, the delta15N value of C. baicalensis was higher than C. dybowski,

because adult of C. baicalensis ate more pelagic sculpin juveniles than that
of C. dybowski. In our results, we suggest that differences in diet and
isotope values between the two pelagic sculpins was shown in both basins,
having each different food resource environment and food choice.
HLADIK, CHRISTINE* and JOHN SCHALLES. Department of Environmental and Atmoshperic Sciences, Creighton University, Omaha, NE. Phytoplankton signals in southeastern estuaries: Calibration of airborne
imagery with close range measures.
Our study examined the phytoplankton component of water column reflectance at and in the vicinity of three National Estuarine Research Reserves
(Apalachicola Bay, Sapelo Island, and ACE Basin). Close-range water reflectance and optically active water constituents were measured with a dualheaded Ocean Optics USB2000 system at stations selected to capture longitudinal gradients in coastal rivers and embayments. The stations included
a wide range of algal, total seston, and CDOM concentrations. In October,
2002 and June, 2003, an airborne imaging spectrometer (AISA system)
was flown at Apalachicola and ACE Basin, respectively, coincident with
the close range measures. The close range data were used to parameterize
and compare several chlorophyll prediction algorithms developed for turbid, Case II waters. These algorithms rely primarily on pigment absorption
and particle scattering features in the red and lower NIR wavelengths. In
turn, these algorithms were applied to pixel classifications of the imaging
spectrometry data to produce chlorophyll distribution maps. Our findings
have relevance to monitoring efforts and HAB detection in the optically
complex coastal environments of the Southeastern U.S.
HO, CHUAN-KAI,* CRISTIANO S. SALGADO,* NILAM B. DAVE*
and STEVEN C. PENNINGS. University of Houston, Houston, TX. Latitudinal variation in herbivore densities and damage to plants in coastal salt marshes.
Biogeographic theory predicts that plant-herbivore interactions are more
intense at low versus high latitudes. Salt marshes along the Atlantic coast
of the U.S. provide an ideal system to examine this theory because similar
plant and animal communities occur across a wide latitudinal range from
Maine to Florida. We tested two hypotheses: 1) herbivores are more abundant at low versus high latitudes, and 2) damage to plants by herbivores
is greater at low versus high latitudes. Sampling was conducted monthly
from June to September, 2002, at ten high-latitude sites (ME, MA, RI, CT)
and ten low-latitude sites (SC, GA, FL). We measured herbivore damage
to four plant species (Iva, Juncus, Solidago, Spartina) that occurred in both
geographic regions, and measured herbivore densities (aphids, beetles,
crabs, acridid grasshoppers, tettigoniid grasshoppers, hemiptera, snails) on
each plant species. Results showed that 1) as predicted, herbivores were
more abundant at low versus high latitudes for most plant species (Solidago
was the exception), and 2) as predicted, damage by herbivores to plants
was more intense at low versus high latitudes, for all four plant species.
These results support the biogeographic theory that plant-herbivore interactions are stronger at low versus high latitudes.
HOBBS, N. THOMPSON1,* and MICHAEL W. MILLER.2 1 Natural Resource Ecology Laboratory, Colorado State University, Fort Collins, Colorado; 2 Colorado Division of Wildlife, 317 W. Prospect Road, Fort Collins,
CO. Dynamics of chronic wasting disease in captive Mule Deer.
Chronic wasting disease (CWD) of the deer family is a transmissible spongiform encephalopathy, a member of a group of infectious diseases affecting animals and people known as prion diseases. Similar diseases include
scrapie in sheep and goats, bovine spongiform encephalopathy in cattle and
Creutzfeldt-Jacob disease in humans. These diseases are associated with
proteinase-resistant prion protein that accumulates in the brain of affected
individuals, causing neural degeneration and eventually death. An epidemic
of CWD has been ongoing in northeastern Colorado and southeastern Wyoming for at least two decades. The ability to model the dynamics of the
disease is hampered by the absence of estimates of transmissions rates,
which are exceedingly difficult to measure in free-ranging populations. We
used likelihood techniques and information theoretics to select models portraying two epidemics of CWD in captive populations. Candidate models

included a simple two state variable model of susceptibles and infecteds,

a model including susceptibles, latents and infecteds, and a model that
included transmission from environmental residues. Transmission was represented as mass action and pseudo-mass action. Models with support in
the data (wr . 0.9) suggested that transmission occurs at the rate of 1-2
infections per infected individual per year.
HOCHSTRASSER, TAMARA,1 DEBRA P.C. PETERS1 and JEFFREY S.
FEHMI.2,* 1 USDA-ARS Jornada Experimental Range, New Mexico State
Unversity, Las Cruces, NM, USA; 2 US Army ERDC-CERL, Ecological
Processes Branch, Champaign, IL, USA. Sustainable management of
arid lands: Recovery time and threshold dynamics in desert grasslands
and shrublands.
For managed disturbances, such as military training activities or recreational use, a sustainable disturbance regime allows ecosystems to recover
to pre-disturbance conditions between disturbance events. In arid environments, recovery from disturbance typically requires long time periods. Furthermore, disturbance may alter vegetation dynamics such that thresholds
are crossed and a shift in dominance from grasses to shrubs is observed.
Such a shift in dominance is considered irreversible under current climatic
conditions because of positive feedback mechanisms between the biotic
and abiotic environment. Disturbance management in arid lands depends
on knowledge about recovery time and thresholds in system dynamics. The
objective of this study was to investigate how recovery time and susceptibility to a change in dominance vary between vegetation types in Southern
New Mexico. We used an individual-based simulation model (ECOTONE)
to simulate disturbances with different timing with respect to climate
events, different sizes, and different intensities in desert grasslands and
shrublands. Both vegetation types recovered easily from low intensity disturbances. Recovery time on large disturbance was determined by the seed
dispersal distance of the dominant species. Disturbances with high intensity
in grasslands caused a shift in dominance from grasses to shrubs. The
probability of such a shift in dominance depended on the timing of the
disturbance with regard to climatic events and soil type. These results correspond to observations on vegetation dynamics after disturbance in southern New Mexico. Using this model, we were able to refine disturbance
management recommendations for desert grasslands and shrublands.
HOFFMAN, KATHERINE E.,* JASON R. TAYLOR* and MELISSA
JURGENSEN-ARMSTRONG.* Ecological Society of America, Silver
Spring, MD. SEEDS: Promoting ecology opportunities for minority
students.
The Strategies for Ecology Education, Development and Sustainability
(SEEDS) program aims to increase the number of underrepresented minorities in the field of ecology by promoting ecology opportunities for
minority students and their faculty. Started in 1996, the program expanded
in 2002 to serve a greater number and diversity of students, faculty, and
schools. The expanded SEEDS program has two main components: (1)
Program Activity Grants partner the Ecological Society of America with
the United Negro College Fund to award institutional grants to five UNCF
schools for faculty development, curriculum development and recruitment
efforts and (2) Project-Wide Activities expand and extend the impact of
the program to a broader spectrum of faculty and students by supporting
Student Undergraduate Research Fellowships, Student Field Trips, Campus
Ecology Chapters, and ESA Annual Meeting Travel Awards.
HOFMOCKEL, KIRSTEN S.* and WILLIAM H. SCHLESINGER. Nicholas School of the Environment and Earth Sciences, Duke University, Durham, NC, 27705. Heterotrophic N fixation in a Loblolly pine FACE
site.
Although forests can be large terrestrial carbon sinks, soil fertility may
limit the sustainability of carbon sequestration in response to increased
atmospheric carbon dioxide. At the Duke Free-Air CO2 Enrichment (FACE)
site, 6 years of carbon dioxide fertilization (ambient +200 ppm) has significantly increased forest net primary production (+25%) and total nitrogen
in the foliar canopy (+16%), indicating an additional 13 kg N/ha/yr has
been assimilated from soils under elevated carbon dioxide. Mechanisms
supporting increased nitrogen acquisition have not been identified. Here we

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151

report on biological N-fixation rates determined using the acetylene reduction assay, in litter and mineral soil during three years of the CO2 enrichment experiment. Rates of N-fixation averaged 0.69 and 0.34 kg N/ha/yr
for litter and mineral soil. Nutrient addition experiments (+glucose, Fe, Mo,
P) indicate biological N fixation at this site is primarily limited by labile
carbon availability. Despite significant increases in forest floor carbon
(+60gC/m2/yr) under CO2 enrichment, no significant difference in N-fixation between control and CO2 fertilized plots was found in litter or mineral
soil (p50.95 and 0.20). These results suggest that (1) the carbon associated
with CO2 enrichment is unavailable for diazotroph metabolism and (2)
biological N-fixation is not an incremental source of N supporting increased
primary productivity in CO2 enriched plots.
HOFSTETTER, RICHARD W.,1,* MATTHEW AYRES,1 KIER KLEPZIG2
and JOHN MOSER.1 1 Dartmouth College, Department of Biological Sciences, Hanover, NH, USA; 2 USDA Forest Service, 2500 Shreveport Hwy,
Pineville, LA, USA. Evaluating the importance of multiple drivers in
outbreak population dynamics.
Understanding how and why populations fluctuate has remained a central
focus of ecology. Several bark beetle species exhibit dramatic fluctuations
in abundance and are capable of causing massive mortality within pine
forests during outbreaks. Bark beetle population dynamics are often attributed to variations in natural enemies, climate, or host tree resistance, and
multivoltine bark beetle species are likely differentially impacted by various mortality factors throughout the year. The relative impact of each of
these factors in driving oscillations within years and across many years is
not well understood. Annual growth rates of multivoltine species are the
collective result of mortality and reproduction of each generation throughout the year. Without accurately sampling mortality factors at each generation, the biological mechanisms that drive prey population dynamics can
be misleading. For example, how do univoltine predators affect the withinyear dynamics of multivoltine prey? Here, we present results that suggest
that within-year dynamics play an important role in the long-term (years)
dynamics of the southern pine beetle, and model the important biological
mechanisms (predators, competitors, climate) that drive within-year and
multi-year fluctuations in southern pine beetle populations. Specifically we
demonstrate that seasonal variation in mortality associated with tree defenses (spring), major beetle predators (spring, fall), and antagonistic fungi
(late spring-fall) can result in complex, non-linear dynamics in beetle population dynamics.
HOHMAN, DOUGLAS S.* and STEVE BREWER. University of Mississippi, University, MS. An investigation of bird species of concern in
three climax communities in northern Mississippi: Implications for fire
management.
Declines in bird populations are of great concern to biologists and wildlife
managers. Efforts to restore natural plant communities would benefit from
considering effects on bird species of concern. In this study, we contrasted
bird and plant communities among three climax communities in northern
Mississippi: 1) mature beech-poplar-sweetgum forests on alluvial terraces,
2) a rare fire-maintained upland oak-pine savanna, and 3) a rare fire-maintained calcareous prairie. Bird communities were identified during the
breeding season in the closed-canopy forests and throughout the year in
the prairie and adjacent oak-pine savanna. As expected, the mesic closedcanopy forests were well represented by forest-obligate Neotropical migrant
species of concern, and the relic calcareous prairie appeared to be beneficial
habitat for grassland and savanna species of concern such as Bachman9s
Sparrow, LeConte9s Sparrow and Henslow9s Sparrow. In contrast, we detected no bird species of concern in the oak-pine savanna. The latter community was unique in other respects, however, in that we found successful
oak regeneration and a rich herbaceous groundcover that contained some
uncommon savanna plant species. The characteristics of this upland oakpine savanna contrasted sharply with those of closed-canopy upland oakpine-sweetgum forests, which contained many of the forest-obligate migrants we found in alluvial-terrace forests, but lacked successful oak regeneration or flowering by relic savanna plants. We conclude that maintenance of natural disturbance regimes favors characteristic bird and plant
species in prairies and mesic forests. Restoration of fire-maintained oakpine savannas from oak-pine-sweetgum forests will likely adversely affect

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some species (forest-obligate birds) and benefit others (upland oaks, savanna wildflowers). Hence, oak-pine savanna restoration efforts need to be
integrated with efforts to preserve and restore refugia for forest-obligate
migrants in mesic forests of alluvial terraces and floodplains.
HOLDO, RICARDO M.1,* and JONATHAN TIMBERLAKE.2 1 Department of Ecology and Evolutionary Biology, Princeton University, Princeton, NJ, USA; 2 Biodiversity Foundation for Africa, Bulawayo, Matabeleland North, Zimbabwe. Rooting profiles along a Kalahari sand woodland catena.
The below-ground structure of plant communities has been largely unexplored, despite the fact that below-ground processes may be the primary
determinants of above-ground community structure in ecosystems where
water is a limiting resource. In a study conducted in northwestern Zimbabwe, we analyzed rooting patterns of the woody plant community along
an edaphic gradient on a Kalahari sand woodland catena. The catena extended for 200 m from open grassland on fine-textured sand in the valley,
to deep, unstructured, medium-textured sand on the heavily wooded plateau. The root systems of 45 trees and shrubs were excavated, mapped and
digitized to quantify root biomass as a function of depth. Rooting depth
increased (50% rooting depth: R250.49, p , 0.0001; 95% rooting depth:
R250.27, p , 0.005), and the center of mass of the profile shifted towards
the maximum rooting depth (p , 0.005) as a function of distance from the
valley. The valley margin was dominated by shallow-rooted Terminalia
sericea, the mid-slope by a mixture of shallow and deep-rooted species
(e.g., Combretum zeyheri and Erythrophleum africanum), while on the plateau woodland dominants such as Brachystegia spiciformis and Baikiaea
plurijuga had massive and deep roots with a more even distribution of
lateral roots along the taproot. The below-ground changes were associated
with above-ground increases in tree basal area and species richness. We
hypothesize that increasing differentiation of below-ground niche space allows greater coexistence of alternative water-use strategies, resulting in the
observed increase in species richness up the catena.
HOLDSWORTH, ANDREW R.,* LEE E. FRELICH and PETER B.
REICH. University of Minnesota, hold0094@umn.edu, Saint Paul, MN.
Patterns of earthworm invasion and understory plant composition in
lakeside northern hardwood forests.
Current studies show that exotic earthworm invasion can significantly affect
the understory plant communities and nutrient cycling of northern hardwood forests. However, little is known about the extent of earthworm invasion. We conducted a survey of understory plant communities, earthworms, soils, and tree composition in 314 plots located in 40 mature hardwood stands in the Chippewa and Chequamegon National Forests (Minnesota and Wisconsin, respectively). Stands were comparable in overstory
composition, biomass, geology, and management history. In both regions
we found a correlation between the presence of exotic earthworms and the
presence of cabins, resorts, boat landings, roads, and campsites. The earthworm species most associated with duff loss (Lumbricus rubellus) was
found in over two-thirds of the Chippewa and Chequamegon plots. The
nightcrawler, L. terrestris, inhabited nearly half of the Chequamegon plots
while only occupying 16% of Chippewa plots. However, earthworm-free
plots were much more common in the Chequamegon, 19% vs. only 3% in
the Chippewa. Plots with all three earthworm ecological groups (litter
dwelling, upper soil dwelling, deep burrowing) had an average of 50% less
sugar maple seedling cover and between 10 and 20% less plant species
richness than plots with few or no earthworms. The abundance of the sedge
Carex pensylvanica was significantly higher in plots with abundant earthworms. Our results further support other studies showing that exotic earthworms significantly alter the structure and composition of northern hardwood forest understory plant communities and that the invasion is in an
advanced but incomplete stage.
HOLLAND, MARJORIE M.,* N.A. N.A., N.A. N.A., N.A. N.A., N.A.
N.A., N.A. N.A., N.A. N.A. and N.A. N.A. P.O. Box 1848, The University
of Mississippi, mholland@olemiss.edu, University, Mississippi, USA.
Common themes of a holistic approach to sustainable water resources.
Successful examples of holistic approaches to water resource management
and restoration have recently become more widespread, and thus have pro-

vided opportunities for reflection on common characteristics. While

grounding in sound science continues to be absolutely critical in formulating appropriate management strategies, a greater appreciation of the intrinsic links between human infrastructure and natural areas is allowing for
further constructive discussions of ways to foster sustainable freshwater
systems. Managers and the lay public understand the need to look both
upstream and downstream of a project in order to truly appreciate the possible impacts of what is being proposed. Especially in the last couple of
years the emphasis on "partnerships" has expanded, whether at the local,
state, regional, national, or international level. These partnerships can include collaborations between industry and government, between government and academia, between academia and nongovernmental organizations,
or in some cases among them all. Partnerships allow us to share the burden,
the responsibility, the credit, and the pleasure of watching something develop, grow, and work well. Several successful regional and national partnerships are presented and discussed throughout this special session. Leadership in promoting sustainable water resources is applauded on all levels.
HOLT, ROBERT D. University of Florida, Gainesville, Florida, USA. Synthesis and conclusions: Community modules and infectious disease epidemiology.
This talk provides an overview of common themes that emerge from case
studies presented by other speakers in the symposium. I will suggest that
one approach to analyzing the interface of classical community ecology
and epidemiology is the close analysis of community modules: small sets
of species interacting in a specified pattern of interactions. Systems with
infectious disease can both mimic familiar modules (e.g. food chains,
shared predation leading to apparent competition, and intraguild predation)
and lead to interesting effects when woven into modules that exist independent of the disease. Infectious disease impacts in a community context
often involve an interesting blend of trait-mediated and density-mediated
indirect interactions, with important and at times even surprising dynamical
consequences. This will be illustrated using several models, including models with a mixture of predator-prey and host-pathogen interactions.
HOLZMUELLER, ERIC J.,1,* SHIBU JOSE2 and MICHAEL A. JENKINS.3 1 University of Florida, Gainesville, FL, United States; 2 University
of Florida, Milton, FL, United States; 3 Great Smoky Mountains National
Park, Gatlinburg, TN, United States. Survival and health of dogwood
(Cornus florida L.) in response to burning.
Survival of flowering dogwood (Cornus florida L.), one of the most common understory species in eastern forests, is threatened by a fungus, Discula destructiva, the causal agent of dogwood anthracnose. Where anthracnose is present, mortality among Cornus florida has been quite high. Currently there are no management plans to protect populations of dogwood
in eastern forests. This study looks at Cornus florida in burned and unburned areas in the Great Smoky Mountain National Park (GSMNP). Stand
composition, structure, microenvironment, and soil conditions of burned
and unburned areas are being evaluated to see how Cornus florida is affected by fire. Preliminary results indicate that heavy Cornus florida mortality has occurred in the western GSMNP over the past two decades (between 1979 and 2000). However, Cornus florida density doubled in burned
areas compared to unburned areas, likely a result of stump sprouting and
reduced shading in burned stands. This information will help decide how
prescribed burning should be applied to reduce dogwood anthracnose in
the GSMNP as well as other areas along the East Coast of the US.
HONDORP, DARRYL W.1,2,* and STEPHEN B. BRANDT.2 1 Cooperative
Institute For Limnology & Ecosystems Research (CILER)-University of
Michigan, Ann Arbor, MI, USA; 2 NOAA-Great Lakes Environmental Research Laboratory (GLERL), Ann Arbor, MI, USA. Comparative foraging behaviors of slimy sculpin and deepwater sculpin.
Sympatric populations of ecologically similar fish often exhibit variation
in food and habitat use that is related to interspecific differences in morphology, feeding behavior and the elusiveness of potential prey. In this
paper, we examined how these factors interact to influence the prey and
habitat selection patterns of slimy sculpin (Cottus cognatus) and deepwater
sculpin (Myoxocephalus thompsoni) in the Laurentian Great Lakes. Con-

clusions were based on results from a series of feeding experiments where

slimy and deepwater sculpin were videotaped while foraging on the benthic
amphipod Diporeia hoyi and the freshwater shrimp Mysis relicta. Both
Diporeia and Mysis are key constituents of the diet of these fish. Preliminary results indicate that both sculpin have significantly greater capture
success rates with Diporeia than with the more elusive Mysis. Mysis have
a keen ability to detect and evade foraging sculpins. When attacked, strong
thrusts of the telson accelerate mysids rapidly away from the predator, a
behavior that greatly increases the probability of escape. Diporeia lack any
notable predator defenses and are therefore more easily captured. Results
also indicated that deepwater sculpin capture Mysis with greater success
than do slimy sculpin. This difference in feeding performance suggests
interspecific variation in sculpin prey detection capabilities and strike mechanics and may help explain the consistent pattern of depth segregation
exhibited by these sculpin species in the Laurentian Great Lakes.
HONG, BONGGHI,1,* DENNIS SWANEY,1 PETER WOODBURY2 and
DAVID WEINSTEIN.1 1 Boyce Thompson Institute for Plant Research,
Tower Road, Cornell University, Ithaca, NY, USA; 2 US Forest Service,
Northeastern Research Station, Durham, NH, USA. Long-term nitrate
export from Hubbard Brook Watershed 6 driven by climatic variation.
The long-term pattern of nitrate export from Watershed 6 at Hubbard Brook
Experimental Forest (HBEF), a long-term ecological research (LTER) site,
during 1964 to 1994 can be characterized as ten years of high export (19681977) followed by twelve years of low export (1978-1989), including three
"spikes" in 1970, 1973, and 1976, and a subsequent "spike" in 1990. Disruptions in N cycling by events such as soil freezing, insect defoliation,
and drought have been suggested to explain this pattern. We developed a
nitrogen cycle model to demonstrate that this long-term pattern can be
reproduced largely without consideration of these events. Comparisons of
simulated N fluxes between high and low export years suggested that inorganic N input to the soil, both as atmospheric N deposition and N mineralization, was significantly higher in periods of high streamflow nitrate
flux than in low periods. As a result, simulated inorganic N pools (ammonium and nitrate) and fluxes (nitrification, plant uptake, denitrification,
and ammonia volatilization) were also significantly higher in periods of
high nitrate streamflow losses. By swapping the time sequences of inorganic N input between high and low export years, it was shown that N
mineralization, not atmospheric N deposition, was the actual driver for
creating the observed long-term pattern. Although simulated nitrification
showed a stronger relationship with measured streamflow nitrate flux than
N mineralization did, the rate of nitrification was heavily dependent upon
availability of soil ammonium supplied from the N mineralization process.
Because N mineralization in the model is a function of soil temperature
and moisture only, we conclude that shifts in the interaction of these two
variables over time produced the shifts in nitrate stream exports.
HOOK, JAMES E.* and ELIZABETH R. BLOOD. How does long-term
drought affect watershed management?
The combination of multi-year droughts in the Southeast and increased
water demand for urban growth and agricultural irrigation raised the awareness of water as a limited resource and has prompted a series of policy and
cultural changes. Between 1980 and 1995, 11 of the 15 most severe agricultural droughts occurred. Irrigation became a mainstay of production.
Meanwhile, Florida and Georgia became the 3rd and 4th fastest growing
states. Demand grew for dependable water supplies to drought-proof municipalities and industries. Georgia reservoir construction and requests to
alter management of federal reservoirs prompted neighboring states to challenge the US Army Corps of Engineers and Georgia in court. Settlement
of these interstate water disputes is still underway, but early in negotiations,
it became clear that Georgia would have to develop a culture of water
consciousness and carefully plan its municipal and agricultural growth to
minimize impacts on natural systems and users in downstream states. Planning centered upon demands during drought years. Data and modeling
raised concerns whether Georgia could protect streams and needs of neighbors during severe drought. Regulators opened dialog with farmers concerning mandatory cutoff during severe droughts to protect streams and
rivers. Compromise legislation was passed to protect both investments in
irrigated farms and flow in streams. Meanwhile, local groups in the health,

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153

business, and agriculture began education and action to assure a place in

decision making. Georgia also began work on its first comprehensive
drought plan. Tri-state negotiations, agricultural compromise legislation,
and drought planning quickly rose to crisis level as a drought that began
in summer of 1998 stretched through 2002, becoming the worst in its history. Water supplies reached precariously low levels in several communities, private wells went dry, and mandatory bans on many outdoor water
uses occurred statewide. With public awareness and planning already underway in Georgia, it became both possible and necessary for the state to
initiate comprehensive statewide water planning. As this abstract is submitted, the Georgia legislature is debating water legislation drafted following a bipartisan planning study committee, sweeping legislation prompted
by recent droughts.
HOOKER, BETH A.,* THOMAS F. MORRIS and ZOE G. CARDON.
University of Connecticut, Storrs, CT. Long-term effects of tillage and
corn stalk return on the dynamics of old and new carbon pools.
The ability to increase pools of soil organic carbon (SOC) in agricultural
ecosystems is of interest both for sequestering atmospheric CO2 in relatively long-lived soil pools, and for restoring organic matter pools that are
important to soil "health." It has been well established that tillage and
harvest management regimes can influence SOC in cropland, but long-term,
continuous experiments are rare. Such experiments are essential for exploring processes whose results accrue slowly in soils. We are investigating
the dynamics of old and new soil carbon pools using d13C analysis in corn
fields established in 1972 at the UConn Research Farm. The plots have
been under no-till (NT) or conventional till (CT) management, with silage
corn (no residue returned annually) or grain corn (shoot residue returned
annually), with three replicates. These plots are very unusual in long-term
(.20 year) agricultural experiments because both tillage differences and
differential aboveground C input practices are in place. Our data indicate
that the relative half life of the relic C3-C (g C kg-1) is not influenced by
tillage or C input levels in the upper 5 cm of the soil, though significantly
more relic C3-C has been lost from the tilled, silage treatment in the 5-15
cm depth than from other treatments. After 30+ years of corn shoot return
to the grain plots, the total amount of corn-derived C remaining in grain
or silage treatments under NT or CT is similar, even though nearly four
times as much C was applied to the grain plots. Our results indicate that
there is a rapid cycling of the aboveground corn-derived C back to the
atmosphere. This calls into question the potential for the annual return of
biomass to substantially increase soil C storage over the long term.
HOOKER, TOBY D.,* JOHN M. STARK and URSZULA NORTON. Department of Biology / Ecology Center, Utah State University, Logan, UT,
USA. Plant phenology effects on soil microbial C and N cycling in a
semiarid ecosystem.
Invasion and dominance of winter-annual cheatgrass (Bromus tectorum)
into perennial dominated shrub-steppe ecosystems may alter seasonal patterns of microbial C and N cycling via shifts in timing of plant detrital C
inputs to soil. Previous work in Rush Valley, Utah, found large increases
in dry-season soil NO3- concentration under cheatgrass over three consecutive years, occurring 6-8 weeks after plant senescence, with no large NO3accumulation observed under perennial dominated vegetation types. Both
plants and soil microbes respond to infrequent and unpredictable summertime pulses of rainfall and inorganic N, though little evidence exists that
this uptake can be related back to net C gain for plants. A herbicide study
was initiated to examine whether plant N uptake maintained low soil NO3levels under perennial vegetation. Glyphosate was added to plots in annual
cheatgrass and perennial crested wheatgrass vegetation types in late May
2001, the time of cheatgrass senescence. Measurements of field soil inorganic N concentrations were collected monthly, with laboratory potential
C and net N mineralization rates, and organic C and N contents from soil
organic matter density fractions. Soil moisture was slightly higher under
cheatgrass, and herbicide treatment differences were small for the first year
of measurements. Cheatgrass had greater potentially mineralizable C than
crested wheatgrass across the growing season, however there was no effect
of herbicide treatment for either vegetation type. The pattern of field soil
NO3- accumulation was similar among herbicide and control plots for cheatgrass. Soil NO3- concentration increased after four months in crested wheat-

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Abstracts

grass herbicide plots, and remained elevated the next growing season under
continued herbicide treatment, relative to control plots. It remains unclear
whether the observed NO3- accumulation in herbicide treated crested wheatgrass plots, and in the annual grass system, is due to lack of plant N uptake,
or C limitation of microbial biomass due to the reduction in belowground
C inputs.
HOOPER, DAVID U.1,* and JEFFREY S. DUKES.2 1 Western Washington
University, Bellingham, WA; 2 Carnegie Institution of Washington, Stanford, CA. Plant traits, community assembly and ecosystem functioning.
The strengths of different forces that structure communities are particularly
relevant to the debate on diversity effects on ecosystem functioning. Some
argue that complementarity among species or functional groups has a
strong effect on ecosystem processes. Others counter that the traits of dominant species are the primary biotic determinant of process patterns and
rates, and that because ecological communities follow distinct (but unspecified) assembly rules, randomized experiments cannot provide insight
into the effects of diversity on processes. What if complementarity is one
such strong assembly rule? We are investigating the effects on community
composition of seed input, community interactions (complementarity/competition), and historical effects (initial community composition and diversity) on subsequent trajectories of community development in California
serpentine grasslands. We tested these mechanisms in synthetic communities in which we had weeded to maintain plot composition for the previous
nine years. Two years following cessation of weeding, we found that 1)
less diverse plots responded more strongly to input from neighboring seed
sources than did more functionally diverse plots, indicating that random
assembly can operate early in community assembly; and 2) species that
differed in functional characteristics from the intact community were more
likely to invade successfully, indicating that internal community dynamics
(e.g., competition/complementarity) can restrict patterns of community assembly from a functional, if not species, perspective. It is too early to tell
if initially different communities will converge. These results indicate that
both random assembly (within the constraints of environmental conditions)
and complementarity can be strong forces in structuring communities,
though they may have their strongest effects at different points along the
trajectory of community change.
HOPPER, WILLIAM E.* and OLAYENI AKINBOYEWA.* Florida Memorial College, Miami, Florida, USA. Urban tree cover and species richness in three Miami-Dade, Florida neighborhoods.
GIS analysis of historical aerial photographs was used to calculate the
percent tree cover and the number of species present in three urban neighborhoods in Miami-Dade County, Florida. These neighborhoods are located
in Bay Shore (Miami), Opa-locka, and Hialeah. Demographic characteristics of each neighborhood were also calculated and correlations established
to identify areas of concern and to develop strategies to be utilized in
increasing urban tree cover. Preliminary results show that the number of
trees in at least two neighborhoods have decreased significantly between
1968 and 1980 due to factors such as death of trees, natural disturbance,
and disease.
HORNER-DEVINE, M. CLAIRE* and BRENDAN J.M. BOHANNAN.
Stanford University, Stanford, CA. Phylogenetic diversity and structure
of bacterial communities.
In most studies of community ecology, measures of diversity and community structure describe the number and relative abundance of different
species present. Only rarely do these measures include information about
the relatedness of the species. However, the taxonomic structure of communities offers insight into the forces shaping and maintaining community
composition. For example, where negative species interactions, such as
interspecific competition or predation, might cause increased mortality
among ecologically similar species, we expect community members to be
less closely related than expected by chance. In contrast, if co-occurring
community members are more ecologically similar than expected by
chance, species may be partitioning habitats according to their autecology.
We applied phylogenetic tools to a data set of bacterial sequences from
five aquatic communities and found that each community contained lower

taxonomic diversity than expected by chance. This phylogenetic clustering

suggests that closely related taxa in each community share traits related to
habitat requirements and/or dispersal and colonization abilities.
HORSLEY, STEPHEN B.,1,* ROBERT P. LONG,2 RICHARD A. HALLETT3 and SCOTT W. BAILEY.4 1 USDA Forest Service, NERS, Irvine,
PA, USA; 2 USDA Forest Service, NERS, Delaware, OH, USA; 3 USDA
Forest Service, NERS, Durham, NH, USA; 4 USDA Forest Service, NERS,
Campton, NH, USA. Base cation nutrition, stress and sugar maple
health in the northeastern United States.
Sugar maple (Acer saccharum Marsh.) decline disease has occurred sporadically in the northeastern United States and eastern Canada during the
past 50 years. Typically, crown vigor deteriorates, progressing through fine
twig and branch dieback, frequently ending in tree mortality. Sugar maple
decline usually has been associated with excessive stress, particularly from
insect defoliation. The most recent sugar maple decline episode occurred
on the Allegheny Plateau in northern Pennsylvania. We evaluated crown
health, foliar nutrition and stress histories in a total of 84 stands in Pennsylvania, New York, Vermont and New Hampshire that included the range
of soil and geologic sites where sugar maple is found in the northeastern
United States. Crown health was evaluated on three 400 m2 plots per stand
using North American Maple Project protocols. Foliar chemistry was analyzed on 5 asymptomatic trees per stand and stand stress history was
evaluated using a digitized defoliation database and through interviews
with landowners. Foliar chemistry was a good index of site quality. Examination of 43 stands in western Pennsylvania and New York showed that
trees on sites with low foliar Mg and Ca, high foliar Mn and excessive
defoliation stress had poorer crown vigor and higher mortality than those
with better nutrition or those with poor nutrition that lacked excessive defoliation stress. Examination of trees in 41 additional stands with similar
ranges of foliar Mg, Ca and Mn, but lacking defoliation stress, in central
and eastern New York, Vermont and New Hampshire showed that crown
vigor was poorer for sites with low foliar Mg and Ca and high foliar Mn,
but little or no mortality occurred. Overall, our study suggests that sugar
maple is predisposed to poorer crown vigor by imbalanced Mg, Ca and
Mn nutrition and incited to mortality by excessive stress, particularly from
defoliation.
HORVITZ, CAROL C.1,* and DOUGLAS W. SCHEMSKE.2 1 University
of Miami, Coral Gables, FL, USA; 2 Michigan State University, East Lansing, MI, USA. Path analysis of the effects of biotic interactions on
reproduction in a neotropical herb.
We used path analysis in two studies of effects of biotic interactions on
reproduction of the understory herb Calathea ovandensis: 1) to estimate
direct and indirect effects of antguards, an ant-tended herbivore and pollinators on flower and fruit production within a season for two years and
2) to estimate direct effects of plant size, herbivory, local competition and
current fruit production on next years production of inflorescences for five
years. In both studies, natural variation in parameters was quantified in the
field for individual plants. For the first study, pollinator visits to all individuals were observed daily throughout most of their flowering period,
lasting about a month. For both years, we found ants positively affected
flower production directly and fruit production indirectly and the herbivore
of reproductive tissues negatively affected both flower and fruit production
directly. The pollinator community differed between years and so did its
effects. Only in the year with better pollinators were there significant direct
effects on fruit initiation, and these were taxon-dependent. For the second
study, marked individuals (mapped to the nearest cm) were censused for
total leaf area and damage by leaf herbivores each Aug, with follow-up
censuses of reproduction every two weeks until the end of Oct. For each
year of the study and for each life history stage, we estimated path coefficients for the direct effects of predictor variables on demographic rates,
including survival, growth and reproduction, resulting in 55 regression
analyses. We did not find significant direct effects of competition or leaf
herbivory on future reproduction. However, plant size in a given season
had direct positive effects on inflorescence production in the following
season as did, somewhat surprisingly, fruit production in a given season.
Thus, pollinators, ant guards and an herbivore of reproductive tissues may
indirectly affect future inflorescence production.

HOSAKA, NAOMI,1,3,* JOSEF F. STUEFER,2 DENNIS F. WHIGHAM1

and NAOKI KACHI.3 1 Smithsonian Environmental Research Center, Edgewater, Maryland; 2 University of Nijmegen, Toernooiveld, Nijmegen, The
Netherlands; 3 Tokyo Metropolitan University, Hachioji, Tokyo, Japan. Ramet dynamics of the clonal understory tree, Asimina triloba.
Body of Abstract: The ecological feature of clonal growth varies with the
life history of plant species and the habitats where they occur. In temperate
deciduous forests, clonal growth is common to understorey tree species.
We explored three possible features of clonal growth in a natural population
of Asimina triloba: risk spreading, securing space occupancy, and foraging
for light through ramet dynamics. In one study, four annual censuses of
ramet populations in distinct patches were conducted in a shaded understorey to test whether (1) ramet turnover is related to patch size, (2) newly
recruited ramets mainly appear inside or outside of existing patches, and
(3) the survival of new ramets is lower than the survival of established
ramets. In all patches the turnover rates of the ramets were stable and
annual natality exceeded mortality during the three years. New ramets occurred inside and outside of the patches and we found no evidence of intrapatch competition between ramets. The ratio of new ramets to the total
number of ramets in the patches was relatively constant during the census
period. Neither was there a significant relationship between initial ramet
size and mortality. These results demonstrate that clonal growth may play
a crucial role in the long-term survival of Asimina triloba in the forest
understorey. In a second study, annual ramet growth and ramet size structure of patches in an open forest edge site were compared with those of
the understorey patches. In the open site, individual ramets showed more
vigorous shoot growth, suggesting that improvement of light availability
enhances individual ramet growth. Flexible growth of the above ground
part of ramets was suggested by a weak correlation between tree-ring and
basal diameter of the ramets in the open site. The flexible growth may
correspond with heterogeneous light conditions in the forest.
HOSSEINI, PARVIEZ R.,1,* ROGER SLOTHOWER,1 EVAN G.
COOCH,2 WESLEY M. HOCHACHKA,1 ANDY DOBSON3 and ANDRE
A. DHONDT. 1 Cornell Lab of Ornithology, Ithaca, NY; 2 Cornell University, Ithaca, NY; 3 Princeton University, Princeton, NJ. Explaining local
variation in the spread of Mycoplasma gallisepticum in House Finches
(Carpodacus mexicanus) in the Eastern United States.
Understanding the factors that affect the spread of disease can be important
to efforts to control and limit the effects of disease. To date, studies on the
localized and irregular nature of spread have been restricted to terrestrial
mammals, which travel relatively slowly, and for which geographic barriers
and Cordons sanitaires can have prominent effects. In contrast, avian-carried diseases can spread relatively rapidly, and many barriers to terrestrial
spread have little or no impact on spread in avian systems. Thus, we investigate as a case study, the spread of Mycoplasma gallisepticum (MG)
in House Finches, focusing on trying to explain the variation in local arrival
of the disease. We examine the effects of local movement, seasonal migration, forest cover, and elevation in explaining the irregular and directionally
biased spread of the disease. Data for the study comes from the House
Finch Disease Survey, a large network of volunteer observers. We use
multi-nominial maximum-likelihood methods, identical to mark-recapture
analyses, to estimate the directionality, distance, and long range components of spread, as well as estimating the probability of detection of disease,
based on effort. We also investigate the role of isolated, long distance jumps
on the spread of the disease. This case provides an interesting contrast to
prior studies of irregular spread in terrestrial mammals, where spread is
affected by geographic barriers, and long distance dispersal by humans.
Although there are limitations to this approach, it provides a useful diagnostic of spread, and may have predictive value in future applications.
HOULTON, BENJAMIN Z.,* DANIEL M. SIGMAN and LARS O. HEDIN. Princeton University, Princeton, NJ. Stable isotope constraints on
internal nitrogen cycles, input-output balances and N fertility of forests.
While the natural abundance ratio of 15/14 nitrogen (N) offers a promising
tool for understanding nitrogen cycles, the sheer number and complexity
of biological and chemical transformations greatly challenge our ability to

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155

deconvolve natural isotope variations. We here use 18O and 15N of dissolved
inputs and hydrologic outputs along a well-constrained Hawaiian moisture
gradient to determine the isotopic imprint caused by dramatic changes in
the forest N cycle - from a N-rich and nitrate based economy, to a N-poor
dissolved organic nitrogen (DON) based economy. We interpret the empirical patterns with a novel, process-based theoretical model that couples
internal isotope fractionations with external input-output balances of N in
forests. Our data on d15N balances show that simple output-based models
do not explain nitrogen isotope variations in Hawaiian rainforest ecosystems. Across all sites, the d15N of total dissolved nitrogen (TDN) in steamwater losses was significantly enriched relative to the d15N of atmospheric
deposition. The d15N and d18O of hydrologic nitrate losses showed little
variation but were correlated with one another (slope ;1.80; R2 5 0.76)
across the gradient. The d15N of DON in streams became markedly depleted
with decreasing N fertility (from ;3.5 to -0.13 per mil), which led to an
overall depletion of the d15N of TDN losses. Taken together, our theoretical
model predictions and empirical results point to significant gaseous N losses and/or missing N inputs of unknown origin in these forest ecosystems.
We will discuss the implications of such findings for conventional theories
on natural isotope systems and the broader N cycle.
HOUSEMAN, GREGORY R.1,2,* 1 Michigan State University, East Lansing, MI; 2 Kellogg Biological Station, Hickory Corners, MI. Can species
pool size and composition explain different productivity-diversity relationships?
Although the influence of plant diversity on ecosystem function (i.e. productivity) has garnered much attention, understanding how diversity varies
across productivity gradients remains an important way to examine how
underlying differences in environmental conditions influence community
dynamics. Recent reviews have found that, while unimodal patterns are
common, positive, negative or no relationship between diversity and productivity also occur with surprising frequency. If species pools strongly
affect local diversity, then different species pools found among sites may
explain the disparate productivity-diversity results. I tested this idea by
adding 0, 5, 15, 30, or 45 new species to low, medium and high productivity (146, 338, 556 g live biomass/m2 respectively) portions of a natural
productivity gradient in a southwest-Michigan oldfield. I assumed that the
naturally occurring species pool was relatively homogeneous because the
gradient occurred over a relatively small spatial scale (200m). Patterns of
colonization and persistence through one growing season demonstrated that
local diversity increased with increasing species pool size. Consistent with
predictions from Huston (1999), the diversity of colonists was unimodally
related to productivity for larger species pools. Simulations based on the
experimental results demonstrated that randomly selected species pools of
equal size from each portion of the gradient could result in negative, unimodal or no relationship between productivity and diversity. I also determined whether each species could germinate and persist in the absence of
competitors by creating monoculture plots along the gradient. Environmental conditions strongly reduced (filtered) species pools at low productivity and suggested that even when the pool sizes are identical unimodal
productivity-diversity patterns will not always occur because the pattern
also depends upon the particular composition of the pool. Thus, historical
and landscape level factors that influence the size and composition of species pools can explain different productivity-diversity relationships.
HOUSER, JEFFREY N.,1 PATRICK J. MULHOLLAND1 and KELLY
MALONEY.2 1 Oak Ridge National Laboratory, Oak Ridge, TN, USA;
2
Auburn University, Auburn, AL, USA. Disturbance of upland soil and
vegetation affects the export of nutrients and sediments to streams
during storm events.
The inputs of sediments and nutrients to streams are affected by watershed
characteristics. The role of riparian zones in buffering streams from watershed disturbance has been well studied. Our objective was to determine
the impact of upland soil and vegetation disturbance on nutrient concentrations in streams with largely intact riparian zones. Spatial variability in
the intensity of military training (infantry and tank maneuvers) at the Ft.
Benning military base (Columbus, GA) results in the uplands of some
stream catchments being highly disturbed while others remain relatively
undisturbed. The watershed disturbance intensity was quantified as the pro-

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Abstracts

portion of the watershed area that consisted of either roads or bare ground
on a slope greater than 3 %. We selected 11 headwater streams located in
11 different catchments across this gradient of disturbance intensity: 3 reference sites, 4 sites of low to moderate disturbance, and 4 sites of high
disturbance. We examined nutrient concentrations in these streams during
baseflow conditions and storm events. Suspended sediments showed the
strongest pattern across the disturbance gradient. In streams draining highly
disturbed catchments, suspended sediments (dominated by the inorganic
fraction) were significantly higher during baseflow conditions and showed
larger increases in concentration during storms. Under baseflow conditions
PO43- concentrations were lower in the disturbed streams (perhaps due to
adsorption by the high level of inorganic sediments in these streams), and
NH4+ and NO3- showed no clear pattern across the disturbance gradient.
During storm events, PO43-, NH4+ and NO3- generally showed larger increases in concentration in the disturbed streams. These results indicate
that streams with intact riparian zones can be affected by upland disturbance.
HOVERMAN, JASON T.,* JOSH R. AULD, ADAM C. MARKO and
RICK A. RELYEA. University of Pittsburgh, Pittsburgh, PA. Integrating
predator-induced behavior, morphology, and life history: Why choose
a single trait?
The number of studies documenting predator-induced plasticity has grown
at an amazing rate. This work has shown that prey are capable of deploying
a variety of responses to predator. However, few studies have examined
how prey integrate their responses to predators and how predator-induced
responses are integrated across different resource levels. This experiment
examined the responses of a common pond snail, Helisoma trivolvis, to
two predators, crayfish (Orconectes rusticus) and giant water bugs (Belostoma flumineum), using a factorial combination of caged predator density
(0, 1, or 2) crossed with food ration (2.5%, 5%, or 10% Spirulina). Over
the course of the experiment we recorded habitat use (the proportion of
snails at the water surface), size-independent morphological responses (six
shell traits), and life history traits (size at first reproduction, number of egg
masses laid, the number of eggs per egg mass, and final mass). The data
showed several important results. Snails did not behaviorally respond to
caged Belostoma or to the food rations by altering their use of the water
surface. However, caged crayfish increased the use of surface habitats by
the snails. Caged Belostoma induced the formation of longer, higher, and
thinner shells but no differences in mass whereas caged crayfish induced
the opposite shell traits and greater mass. Food ration significantly increased snail mass but had no effect on morphology. Analysis of the life
history traits showed that higher food levels decreased time to and age at
first reproduction and increased the number of egg masses and eggs per
egg mass. Snails reared in the presence of predators showed increased time
to first reproduction and fewer egg masses and eggs per egg mass. This
study shows how prey are able to integrate their diversity of phenotypic
responses according to the food and predator regime that it encounters.
HOWE, HENRY F.1 and DIANA LANE.2 1 University of Illinois-Chicago,
(hfhowe@uic.edu), Chicago, IL; 2 Stratus Consulting, Boulder, CO. Voledriven dominance in experimental restorations.
We used 18 replicated plantings of 16 tallgrass prairie species in wet-mesic
soils in western Wisconsin to test for the effects of vole (Microtus pennsylvanicus) herbivory over 48 months on perennials that had reached reproductive age after 26 months of protection. Once voles were admitted,
an otherwise common legume (Desmodium canadense) and grass (Elymus
virginicus) were all but eliminated (combined cover 27% where voles were
excluded, 4% where voles were admitted). Vole herbivory accelerated an
existing succession to domination by an unpalatable mint (Pycnanthemum
virginianum) and black-eyed Susan (Rudbeckia subtomentosa) (combined
cover 33% where voles were excluded, 63% where voles were admitted).
Cover of other common species (Echinacea purpurea, Heliopsis helianthoides, Lobelia silphitica, Silphium integrifolium) changed over four years
without being influenced directly or indirectly by vole access. Repeated
measures ANOVA revealed that net decreases (D. canadense, E. virginicus)
or increases (P. virginianum, R. subtomentosa, Solidago graminifolia) in
cover as a function of vole herbivory masked a variety of first order, second
order, and third order effects that imply transient vole-driven dynamics

during divergent successional processes. Access of vegetation protected

from for 26 months to voles resulted in a .50% drop in proportional
diversity during the ensuing 4 years, reflecting vole-driven simplification
of these communities. Unlike previous experiments in this genre, in which
voles gained access to designated plots at planting, vole-driven changes in
species composition were not accompanied by drastic changes in aggregate
characteristics (species richness, standing crop, cover) that might confound
effects of selective herbivory by voles.
HOWE, KATHERINE M.* SUNY-Stony Brook, Stony Brook, NY. Plant
community composition and natural enemies affect invasion success of
a grassland plant.
Biological invasions are a leading threat to biodiversity, yet we still know
little about what makes some species successful invaders. Two factors believed to improve invader success are reduced damage from predators or
pathogens on exotic species relative to native species and superior competitive ability of invaders compared to native species. I examined the role
of both of these factors in affecting the success of a grassland forb introduced from Eurasia, Hieracium aurantiacum, and its co-occurring, native
congener, Hieracium longipilum. I tested the effects of natural enemies on
both species by experimentally excluding herbivores and pathogens from
plots planted with both species. I also examined the effects of neighboring
plant community composition and edaphic properties on the success of both
species. In all years from 1999-2002, both species showed increased growth
when insect herbivores were experimentally excluded, and neither species
showed a significant response to the exclusion of foliar fungal pathogens.
The native species had increased flower number and taller flowering stalks
in plots where deer were excluded, but there was no effect of deer exclusion
on the exotic species. Success of both species was also related to the composition of the surrounding plant community. H. longipilum achieved larger
sizes in areas with greater cover of legumes, graminoids, and leaf litter,
while H. aurantiacum size was negatively correlated to percent cover of
legumes and leaf litter. H. longipilum size appeared to be unrelated to soil
characteristics, but H. aurantiacum individuals were smaller in areas with
higher levels of soil moisture. The results of this study suggest that although H. aurantiacum appears to have an advantage over the native species resulting from lower rates of herbivory, its invasion is likely to be
limited to areas with suitable soil characteristics and plant community composition.
HOWE, ROBERT W.,1,* JAMES R. KARR2 and AMY T. WOLF.1 1 University of Wisconsin-Green Bay, Green Bay, WI; 2 University of Washington, Seattle, WA. Historical and recent approaches to the assessment
of ecological condition.
The search for effective ecological indicators must first identify what we
are trying to indicate, in other words the conceptual endpoints of field
measurements. Diverse approaches to indicator development reflect the
complexity of human needs and values. Whether the emphasis is on human
health or ecosystem health, however, effective ecological indicators must
emphasize biological condition. Analyses of physical stressors alone do not
adequately address ecological integrity, the implicit endpoint of most environmental laws and regulations. In some cases, simple variables such as
species richness or indicator species abundances may be useful for environmental assessment, but these approaches are likely to overlook important characteristics of local or regional ecosystems. More ambitious strategies evaluate ecological integrity or ecological health as products of multiple physical and biological variables. In this paper we review the science
of ecological indicators and identify the strengths and weaknesses of different approaches to indicator development. We conclude by examining the
management and policy consequences of using (or ignoring) scientifically
rigorous ecological indicators.
HOWELL, JONATHAN P. and DEBORAH A. WALLER. Old Dominion
University, Norfolk, VA, USA. Population densities of the fungus-growing ant Trachymyrmex septentrionalis in southeastern Virginia.
This study was conducted over a period of two years in the Blackwater
Ecological Preserve (BEP) of southeastern Virginia. BEP is undergoing a
program of prescribed burns to convert the loblolly pine habitat to longleaf

pine. Four populations of T. septentrionalis separated by 50+ meters were

located and mapped. Vegetation cover was measured within the populated
areas and in adjacent unpopulated areas at two heights by densiometer.
There were significant differences in cover between populated and unpopulated areas. Population densities were calculated for each nesting site,
ranging from 0.072 to 0.145 nests/m2. No association was found between
cover and population density.
HOYE, KATHERINE, ANDREW J. MCELRONE, CHANTAL D. REID
and ROBERT B. JACKSON. Duke University, Durham, NC. How does
elevated CO2 affect a foliar fungal disease of field-grown Acer rubrum?
Little is known about the impact of disease on plants grown at projected
elevated CO2 concentrations. To address these poorly understood interactions, we surveyed Acer rubrum trees for a leaf spot disease caused by the
fungal pathogen, Phyllosticta minima, at the Duke FACE experiment, Durham, NC. Disease incidence and severity were lower under elevated CO2
with 21% fewer trees and 23% fewer leaves per tree exhibiting leaf spot,
and reduced mean lesion area on infected leaves (5.12 and 6.35 mm2 for
elevated and ambient CO2, respectively). Because the fungus enters the leaf
through stomata, we investigated whether a decrease in leaf stomatal density and/or size with elevated CO2 could explain the reduced fungal infection. In the field, we tested this hypothesis by measuring leaf stomatal
density and size of the stomatal aperture on leaf impressions of A.rubrum
collected in the Fall 2002 at Duke FACE. Leaf impressions were examined
with a scanning electron microscope for determination of fungal infection
and stomatal morphology. For leaves grown at elevated CO2, stomatal size
and aperture were increased with elevated CO2 while stomatal density was
reduced. In growth chambers, we grew red maple seedlings at ambient and
elevated CO2 (350 and 560 mmol CO2 mol-1, respectively) and inoculated
young fully expanded leaves with P. minima to follow its mode of entry
into the leaf and rate of infection. Scanning electron micrographs of infected leaves were taken at regular intervals following inoculation. The
direct impact of elevated CO2 on the growth and reproduction of P. minima
in culture was also monitored. Results suggest that changes in stomatal
morphology under elevated CO2 partly explain the reduced fungal infection
on A. rubrum. Changes in leaf surface wax deposition may also contribute
to reduced infection by altering fungal hyphae signaling and growth towards the stomata.
HSU, CYNTHIA L.* and GEORGE HEIMPEL. University of Minnesota,
St. Paul, MN. Evaluating a parasitoids potential to influence the evolution of insecticide resistance in its host.
Agricultural crops that are genetically modified to express the Bacillus
thuringiensis(Bt) toxin are extremely effective at killing targeted pests, resulting in strong selective pressure to increase the frequency of resistant
genotypes. Repeated use of such crops over time and space can increase
the rate of resistance evolution in pest populations. Factors other than selection intensity can influence this rate, but the relative importance of these
factors is not clear. Natural enemies have the potential to increase or decrease the rate of resistance evolution in a pest population by differentially
impacting susceptible and resistant individuals. In the Bt-corn system, the
European corn borer, Ostrinia nubilalis, is attacked by a specialist parasitoid, Macrocentrus grandii. It is hypothesized that limited dispersal or a
positive density dependent response by M. grandii could create a spatial
refuge for resistant ECB larvae that survive in a Bt-field, favoring an increase in the frequency of the resistant genotype, while a high dispersal
ability and a strong inverse density dependent response may increase the
probability that M. grandii locate resistant ECB larvae. We investigated
the parasitoids dispersal range and tested for spatial density dependence
by collecting ECB larvae at 200 sample sites in a grid pattern within 8 ha
non-Bt corn fields. We found little evidence of spatial structure in the distribution of ECB larvae or parasitoids, and a weak inverse density dependent response by the parasitoid. Parasitoids were found throughout the
fields, demonstrating that dispersal is not a limiting factor in the potential
of M. grandii to locate hosts at this scale. Results suggest that M. grandii
has the dispersal potential, but may not have a sufficient behavioral response to locate and parasitize rare resistant ECB larvae that would survive
in Bt corn fields.

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157

HU, SHUIJIN.* North Carolina State University, Raleigh, NC, USA. Nitrogen limitation to soil microorganisms under elevated CO2.
The impact of atmospheric carbon dioxide enrichment on terrestrial ecosystem C balance (in sign or magnitude) is not clear because the resulting
alterations in C input, plant nutrient demand and water use efficiency often
have contrasting impacts on microbial decomposition processes. One major
source of uncertainty stems from the impact of elevated CO2 on N availability to plants and microbes. We examined the effects of atmospheric
CO2 enrichment (ambient + 370 ppm) on soil microbes and microbial N
acquisition both in the field and in model ecosystems of annual grasses.
Elevated CO2 in the atmosphere enhanced the colonization of plant roots
by arbuscular mycorrhizal fungi and plant N uptake. Increased mycorrhizal
infection of roots coincided with the enhancement of plant competition for
mineral soil N, as measured with 15N as a tracer. Elevated CO2 increased
C but reduced N availability for soil microbes, increasing the microbial
biomass C:N ratio and fungal contribution to the soil microbial community.
These results indicate that CO2 enrichment in the atmosphere tightens N
cycling in terrestrial ecosystems and may facilitate C sequestration through
promoting N conversion from soil to vegetation.
HUBBARD, JAKARA,* TANA WOOD and DEBORAH LAWRENCE.
University Of Virginia, Department of Environmental Science, Charlottesville, VA. Litter addition effects on root biomass and nutrient concentrations in a wet tropical forest in Costa Rica: an experimental approach.
Tropical forests typically occur on weathered, nutrient poor soils. In order
to compensate for low nutrient availability, it is likely that fine roots play
an important role in maximizing nutrient acquisition in these systems. Litterfall is the major mechanism for transferring nutrients from the above
ground vegetation to the soil. Hence, we hypothesized that increased litter
fall would lead to an initial increase in root biomass followed by root death
once nutrients are mineralized from the litter. Additionally, we expected
that an increase in nutrients released from the additional litter would lead
to higher nutrient concentration in roots than sites with less litterfall. To
simulate this experimentally in the field, we established six sites varying
in age and soil type at La Selva Biological Station in Costa Rica. Leaf
litter was removed from a 400m2 plot and added it to a 100m2 plot with
two replicates per site. We sampled fine roots to 5cm depth in the addition
and control plots both 7 months and 1 year after litter addition. Originally,
a comparison of live and dead roots showed that live root biomass was
significantly lower in the addition plots (t-test; p50.005) and dead root
biomass was significantly higher in the addition plots (p50.023) than in
the control plots. However, after one year of decomposition, the control
and addition plots did not differ significantly in their live/dead root ratio.
These results support our hypothesis that increased litterfall leads to greater
initial exploration by roots followed by root death as nutrients become more
available. The live roots were digested using a modified Kjeldahl digestion
and the digestate was analyzed colorometrically for phosphorus concentration. Mean root phosphorus was 0.503 mg/g and, despite slightly higher
phosphorus concentrations in the addition plot, did not differ significantly
between addition and control plots as expected. By repeating this experiment in other locations over a longer period of time with a greater level
of litter addition, perhaps we will gain a better understanding of the nutrient
cycling process in wet tropical forest.
HUBBELL, STEPHEN P.,1 JAYNATH R. BANAVAR2 and AMOS MARITAN.3 1 University of Georgia, Athens, Georgia; 2 Pennsylvania State University, University Park, Pennsylvania; 3 International School for Advanced
Studies (S.I.S.S.A.), Trieste, Italy. Are ecological communities symmetric? Frequency dependence and the statistical mechanics of relative
species abundance.
A simple generalization of Hubbells stochastic neutral theory of relative
species abundance (RSA) leads to an analytical solution for the distribution
of RSA that incorporates symmetric density- and frequency-dependence
(rare species advantage). The predicted RSA distribution is a modified Fishers logseries with an interior mode and a shape controlled by the functional
relationship between species abundance and the ratio of the birth rate to
the death rate (b/d). The ratio b/d . 1 for species with abundances below

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a value near the modal abundance, and b/d 1 for species with abundances
above the mode. Our combined theory shows that two independent symmetric mechanisms, frequency dependence and dispersal limitation, are
both sufficient to explain the skewed RSA distributions observed in large
samples from natural ecological communities. Excellent fits of the theory
to data on RSA are obtained. We also demonstrate a remarkable mathematical concilience with physics in that these distributions are closely related to classic problems in statistical mechanics, including the Boltzman,
Fermi-Dirac, and Bose-Einstein equations. In ecology, our combined results
provide a very promising theoretical foundation for reconciling the niche
assembly and dispersal assembly perspectives.
HUBERTY, ANDREA F.* and ROBERT F. DENNO. University of Maryland, College Park, MD. Responses of saltmarsh-inhabiting insects to
salt-stressed host plants.
Plant stress has been traditionally viewed as a force underlying insect herbivore population dynamics, due to stress-induced phytochemical changes
during times of stress. Previously we conducted laboratory experiments to
determine the impact of salt-stressed plants on an assemblage of phytophagous insects. The results of these studies conclude there is no generalized
response governing the performance of insects on stressed host plants; even
congeners exhibit divergent responses to stressed plants. Therefore, we conducted studies to determine if experimentally stressed host plants in the
field elicit similar herbivore responses as the laboratory experiments. Treatments of salt-addition or no salt (control) were randomly assigned to plots
of Spartina alterniflora. Insects were sampled throughout the season to
determine if those insects species that showed positive responses in the
laboratory (Prokelisia marginata and Sanctanus aestuarium) were found
in higher densities in the salt-addition plots. We also predicted those insects
that showed negative responses in the laboratory (Prokelisia dolus and
Draeculacephala portola) to have lower densities in the salt-addition plots.
The results of our field studies correspond closely to the results from our
laboratory experiment. The effect of stressed host plants on herbivore population dynamics varies with insect species, casting doubt on the relevancy
of plant stress as the underlying factor promoting outbreaks for all herbivorous insects.
HUDSON, PETER J.,1 PETER WHITE,2 MIKE BOOTS,2 ROGER
TROUT,2 NAOMI FORRSETER2,3 and ERNEST GOULD.3 1 Penn State
University, State College, PA; 2 Stirling University, Stirling, Scotland, UK;
3
CEH, Oxford, England, UK. The sudden emergence of rabbit hemorrhagic disease.
Rabbit Hemorrhagic Disease emerged suddenly as a virulent and highly
infectious disease in a group of domesticated rabbits being air transported
to China in 1984. Subsequently, the disease caused massive mortality in
domestic and wild rabbits throughout Eurasia and Australia. This paper
examines a series of questions relating to why and how the disease
emerged. First: Will the disease have a major impact on rabbit populations
in all countries? We undertook a large scale sera survey in the United
Kingdom and found that most populations had already been exposed to the
virus. Serial sampling indicated that an epidemic may pass through the
population without massive mortality. Moreover, sera samples collected
before the emergence of the disease also found evidence of previous exposure leading us to surmise that an avirulent form of the disease had been
circulating. However, sequencing of the capsid gene found no evidence of
a clear virulent or avirulent strain. Second: What conditions and evolutionary selective pressures may cause the rapid evolution and emergence of a
highly virulent disease? Using a generic, individual-based SIR model we
show theoretically that large, stable shifts in virulence may occur in pathogen populations due to a bi-stability in evolutionarily stable virulence
caused by the contact/social structure of the host population. We postulate
recombination coupled with changes in the social structure of rabbit populations may have initiated the rapid evolution of a virulent strain and
caused the observed pandemic.
HUEBNER, CYNTHIA D.* NE Research Station, USDA Forest Service,
Morgantown, WV, USA. Predicting early plant invasions in West Virginia public forests.
Western West Virginia public forests have experienced comparatively less
invasion than forests in more urban regions and may serve as an optimal

landscape for modeling early establishment (as opposed to spread) of exotic

invasive plants. Documentation of the importance of disturbance and environmental variables as predictors of invasion in forests is lacking. I sampled the understory (herbs, shrubs, vines and tree seedlings) of 24 undisturbed (80 or more years old) and 24 disturbed (15-year old clear cuts)
sites in the Monongahela National Forest Cheat Ranger District. Sites,
which averaged 3 ha in size, were randomly selected such that three common ecological land types (sugar maple/basswood, sugar maple/red oak,
and red oak dominated forests) were equally represented in both undisturbed and disturbed sites. Plant species were grouped into exclusive categories: exotic invasive, exotic non-invasive, native invasive, native noninvasive weed, and native non-invasive non-weed. Eleven environmental
and 7 disturbance variables were measured. Data were analyzed using logistic regression analysis. In all sites, exotic invasive weeds had low ranking relative importance values, with Rosa multiflora and Berberis thunbergii being the most common among the 11 exotic invasives present. High
herb/shrub/vine richness and diversity values and high levels of native noninvasive weeds and exotic non-invasive weeds best predicted invasion by
exotic invasives in the undisturbed forests; only high herb/shrub/vine richness was a reliable predictor of invasion in disturbed sites. Undisturbed
sites were more likely to be invaded the closer they were to a paved road.
In the disturbed sites, northeast-facing slopes, lower moss/lichen cover,
higher bare ground cover, shallower slopes, lower light levels, and shorter
distances to gravel and paved roads best predicted invasion. A combined
analysis showed that disturbed sites and sites with northeast facing slopes
were most likely to be invaded. These results may be used with caution to
predict invasion into other similar forests and indicate that, while disturbance is an important predictor, sites with certain environmental characteristics are more prone to invasion.
HUENNEKE, LAURA F.* New Mexico State University, Las Cruces, NM.
The Research-Education Link: Looking toward the Future.
Academic ecologists in higher education have historically played an important role in introducing K-16 students to research (through undergraduate research participation programs, especially). However, the context for
educational outreach and involvement for those ecologist faculty members
is changing dramatically. Federal agencies and some private foundations
have modified their definitions of target groups for participation in undergraduate research and internships. Awareness of the critical link between
K-12 experience and college has led to increased pressure for outreach to
K-12 teachers and students, a type of outreach for which many faculty are
unprepared. The balance between research and teaching (including research
mentoring and outreach) is shifting on many campuses, thanks to recent
Carnegie reports and highly publicized moves by funding agencies. Agencies such as NSF have incorporated broader impact considerations into
funding decisions, but have collected little information to evaluate those
impacts. Finally, the financial crisis looming in many institutions today
places constraints on relatively-expensive research and outreach programs.
Speakers in this symposium present aspects of what we have learned in
recent years about the impact and effectiveness of these programs, and
identify key concerns. I outline some essential components that should be
put into place to make involvement with education and research experience
at these different levels a more viable and sustainable enterprise for research-active faculty.
HUFFMAN, JEAN M.,1,* WILLIAM J. PLATT,1 HENRI GRISSINO
MAYER2 and CARLA J. BOYCE.3 1 Louisiana State University, Baton
Rouge, LA; 2 University of Tennessee, Knoxville, TN; 3 Apalachicola National Estuarine Research Reserve, Eastpoint, FL. Fire history of a Florida barrier island slash pine savanna.
Fire regimes of pine savannas on barrier islands along the coast of the Gulf
of Mexico are unknown. We determined the years and seasons of fires and
turpentine operations using dendrochronological techniques to precisely
date scars from 54 slash pines (Pinus elliottii var. elliottii) on Little St.
George Island, an undeveloped barrier island. We also mapped the spatial
distribution of past fires. Five separate periods with different fire frequencies were found. Fires were frequent between 1866 and 1904 (mean firereturn interval of 4 years). No scars were found from 1905-1923, years
during which turpentine operations (1912-1918) protected trees from fires.

Frequent fires were again recorded from 1924-1945, (mean fire return interval of 4 years). During the period from 1945-1962, turpentine operations
(1949-1956) protected trees from fires and no fire scars were found. The
most recent period, 1963 to the present, had a mean fire-return interval of
9 years with active, although not entirely effective, fire suppression. Most
historic fires occurred during the lightning season. Although the trees used
in this study were not old enough to determine pre-settlement fire frequencies, the data does reveal much about fire regimes on this barrier island
during the past 145 years, showing that historic fire regimes of this barrier
island slash pine savanna consisted of predominately growing-season fires
with short fire return intervals.
HUFFORD, KRISTINA M.* and SUSAN MAZER. Department of Ecology, Evolution and Marine Biology, University of California, Santa Barbara, CA. Local adaptation and restoration of California native grasslands.
California prairies have been dramatically altered by the invasion of Old
World grasses, which have displaced many native perennial grass species
throughout the state. Restoration via seed introduction is critical to reestablish native species, but a new concern the adaptation of plants and
seeds to their site of origin has raised the serious question of the suitability of long-distance transfers of seeds produced by native plants. We
conducted reciprocal transplants of three native grass species in order to
test the hypothesis that local genotypes will have higher fitness when grown
at their home site. The presence of a home-site advantage suggested that
at least two of the native grass species are adapted to local environmental
conditions. Soil characters were analyzed at each site in a preliminary effort
to examine the factors contributing to local adaptation of each species.
These results are discussed in light of: 1) current research to examine the
ecotypic differentiation of native grasses in central California and the nearby Channel Islands, and 2) future research to determine the consequences
of mixing ecotypes during restoration of native grasslands.
HUGHES, R. FLINT* and JULIE S. DENSLOW. Institute of Pacific Islands Forestry, USDA Forest Service, Hilo, Hawaii. Invasion by the Nfixing tree, Falcataria moluccana, alters ecosystem processes in wet forests of Hawaii.
Invasive species pose major threats to the integrity and functioning of ecosystems. Previous studies have shown that when invasive species introduce
new biological processes into an ecosystem, they can change the fundamental rules by which other species survive and reproduce. Here we describe the effects of an invasive alien N-fixing tree, Falcataria moluccana,
on ecosystem processes and stand structure of wet lowland forests on recent
volcanic substrates of Hawaii. Substrates ranged in age from 50 to 300 yr
and, as such, supported a gradient of developing stands undergoing primary
succession. On each of 3 substrates, study sites were established in stands
either invaded by, or free of, Falcataria. Inputs of N via litter fall were 4
to 55 times greater, and P inputs 2 to 28 times greater, in forests invaded
by Falcataria relative to native-dominated forests on comparable substrates. Decomposition rates differed substantially in response to Falcataria
as well. Decomposition rate constants (k) of Falcataria leaf litter were 3
times greater than those of leaf litter of the dominant native species, Metrosideros polymorpha, and rate constants of leaf litter were 2 to 7 times
greater in Falcataria-invaded sites compared to native forest sites. Further,
the presence of Falcataria led to significant increases in plant-available
forms of N and P; combined values of NO3-N and NH4-N captured by ion
exchange resins were 12 to 46 times greater,and PO4-P were 2 to 57 times
greater, in Falcataria-dominated sites compared to native-dominated sites.
These Falcataria-mediated changes in ecosystem processes were accompanied by alterations in forest structure and species composition as well.
While Metrosideros dominated the native forests, almost 100% of Metrosideros suffered mortality following Falcataria invasion. In contrast, the
alien tree Psidium cattleianum was absent from native-dominated sites, but
reached densities as high as 14,000 stems ha-1 within Falcataria-invaded
sites. By transforming ecosystem functions, Falcataria has facilitated the
invasion of species that would otherwise be scarce in these early successional forest systems.

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159

HUI, DAFENG,1,* TAO XU,1 YIQI LUO1 and STEVEN WOFSY.2 1 University of Oklahoma, Norman, OK; 2 Harvard University, Cambridge, MA.
Quantitative assessment of climatic effects on seasonal and interannual
variabilities in net ecosystem exchange.
Seasonal and interannual variabilities in net ecosystem exchange (NEE) of
CO2 are observed at many FluxNet sites. These variabilities have been
linked to direct effect of climatic variations and climate-induced functional
changes (e.g., changes in growing season length and the timing of leaf
emergence). However, we still lack the quantitative understanding of climatic effects on NEE variations. In order to study the climatic and functional effects on NEE, we applied a recently developed homogeneity-ofslopes (HOS) model (Hui et al. 2003, Tree Physiology, in press) and wavelet analysis to a long-term dataset with eight years9 measurements of NEE
and climatic variables in Harvard Forest. HOS model showed that photosynthetically active radiation, soil temperature, relative humidity and vertical velocity (ustar) had significant influences on daytime NEE. Functional
change contributed 2.7% of variation in NEE. The direct interannual climatic variations contributed 4.8% and seasonal climatic changes contributed 77.5% of NEE variation. Soil temperature and ustar were two important factors influencing nighttime ecosystem respiration (RE). About 10.0%
variation in RE was contributed by the functional change. The direct interannual climatic variations contributed 4.4% while the seasonal climatic
changes explained 60.2% of variation in RE. When NEE and climatic variables were decomposed to the fourth level using wavelet analysis, 95.4%
of the variation in approximation of NEE was explained by those of climatic variables and 86.4% of the variation in detail of NEE was explained
by those of climatic variables. The fact that less than 30% of the variations
in detail of NEE at lower levels were explained by climatic variables confirmed that functional change and other factors might have significant effects on NEE.
HULL-SANDERS, HELEN M. and MICKY D. EUBANKS. Auburn University, Auburn, AL. Inbred plants in the field: Can plant defense theory accurately predict plant-insect interactions involving inbred
plants?
Inbreeding (self-pollination) frequently occurs in many plant populations.
Inbreeding typically results in broad, detrimental changes in plant morphology and physiology. These phenotypic changes may be analogous to
other environmental stresses. Two plant defense theories may predict plantinsect interactions involving inbred plants. The Plant Stress Hypothesis
predicts that stressed plants will be more susceptible to generalist herbivores. The Plant Vigor Hypothesis predicts that specialist herbivores prefer
and perform better when feeding on the most vigorous plants (low stress
plants). We studied the interactions between insect herbivores and their
inbred and outbred host plants in the field in an attempt to determine if
plant defense theory could accurately predict the effect of inbreeding on
plant-insect interactions. Our work with the morning glory, Ipomoea hederacea var. integriscula (Convolvulaceae), and its generalist and specialist
herbivores suggests that plant defense theory may accurately predict the
outcome of interactions between inbred plants and their specialist herbivores, but may have differing effects with the suite of generalist herbivores
that attack in the field. While inbreeding depression in I. hederacea has
been documented in the greenhouse, the effects of inbreeding are less clear
under field conditions and vary according to season. Specialist herbivores
may be sensitive to small physiological changes in inbred plants in the
field, but the suite of generalist herbivores may attack plants indiscriminately and may not be sensitive to I. hederacea defense characteristics.
HUMPHREY, LEWIS D.1,2,* and EUGENE W. SCHUPP.1 1 Department of
Forest, Range and Wildlife Resources, Utah State University, Logan, UT,
USA; 2 1437 Hillcrest Drive, Buford, GA, USA. A test of a long-lived
and a short-lived perennial grass for use in restoring Bromus tectorum
communities.
The introduced annual grass Bromus tectorum has replaced native species
in much of the western USA. We compared growth and establishment of
a short-lived (Elymus elymoides) and a long-lived (Pseudoroegneria spicata) native perennial grass in the face of B. tectorum competition. In
autumn 1997, field plots were sown with a range of densities of B. tecto-

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rum, E. elymoides and, P. spicata in a factorial design on an arid and a

semiarid site in central Utah. For E. elymoides and P. spicata target plants,
we estimated biomass per plant from morphological measurements for
spring and summer 1998 and 1999, and summer 2000, and monitored survival and flowering. Bromus tectorum neighbors greatly reduced first-year
biomass for both species on both sites (P , 0.001). On the semiarid site,
this effect was greater on P. spicata than on E. elymoides (p , 0.001), but
effects on spring-summer relative growth rate (RGR) did not differ for the
two species. On both sites second- and third-year RGR was unaffected by
B. tectorum, but second-year biomass and survival were similarily lower
for both species, as a result of initial B. tectorum competition. On the arid
site, first-year effects of B. tectorum on both biomass and RGR were greater, and second- and third-year survival lower, for E. elymoudes than for P.
spicata, and survival was still related to first-year B. tectorum densities in
the third year. The proportion of plants flowering was greater for
E.elymoides than for P. spicata at the semiarid site, but the reverse was
true at the arid site. The short-lived E. elymoides may have ruderal growth
traits, allowing it to effectively exploit the higher resource levels of the
semiarid site, even in competition with B. tectorum. P. spicata better tolerated the abiotic stress of the arid site, while E. elymoides was unable to
compete with B. tectorum there. Yet, P. spicata appeared ruderal-like in
that it quickly established and flowered by the second year on both sites.
HUMSTON, ROBERT,1,2,* DAVID A. MORTENSEN1 and OTTAR N.
BJRNSTAD.1 1 Penn State University, University Park, PA, USA; 2 Washington and Lee University, Lexington, VA, USA. Assessing patch dynamics of common sunflower (Heliathus annuus) in response to weed management in row crop agriculture.
Establishment and spread are important processes in pest weed invasion,
yet few studies have quantified these dynamics with respect to weed management approaches in row-crop agriculture. Weed patches with a range of
initial starting densities were established under a range of management
practices and spread measured over a three year period. Common sunflower, Helianthus annuus, patches were established in year 0 in 1.3 m2 quadrats
centrally located in 9.91 m by 15.85 m plots at three densities: 1) low (500
seeds / 1.3 m2); 2) medium (1500 seeds / 1.3 m2); and 3) high (2500 seeds
/ 1.3 m2). Patches were established in corn and in soybean and monitored
for three years in a corn-soybean rotation cropping system. High and low
management intensities were implemented by applying a full or half
strength herbicide application (bentazon). Cultivation, herbicide application
and harvest activities were oriented unidirectionally across the experimental
area. Spatial variation in density of H. annuus was described by quantifying
the number of seedlings in every 76 x 61 cm (0.46 m2) cell of the overlain
sampling grid. Patch structure was evaluated with one- and two-dimensional metrics to examine extent of patch development in the first two years
of growth. Reaction-diffusion models were then parameterized by assessing
patterns of spatial patch density resulting from seed dispersal in year 0 and
seedling emergence in year 1. Results indicate that crop type influences the
intrinsic rate of increase of the number of plants in a patch, as well as the
relative contribution of natural and anthropogenic forces to seed dispersal
and patterns of patch expansion. Initial seed density had the most significant impact on patch dynamics, influencing growth rate, coefficient of diffusion, and lateral velocity of patch expansion. Experimental treatments
had negligible impact on the asymptotic velocity of spatial expansion of
the weed population. Restricting the magnitude of the initial infestation
appears to hold the greatest potential as a management strategy for controlling expansion of pest weed populations in agricultural systems.
HUNGATE, BRUCE A.* Northern Arizona University, Flagstaff, Arizona.
Progressive nitrogen limitation in grasslands in response to elevated
CO2.
Due to widespread nitrogen (N) limitation of plant productivity, changes
in the soil N cycle can strongly shape the responses of terrestrial ecosystems to rising CO2, including responses such as carbon uptake and storage.
Elevated CO2 can alter N transformations that influence N distribution within ecosystems (mineralization, immobilization, nitrification, plant uptake).
While some of these changes can increase N availability and plant growth
in the short term, in the long term and in the absence of major changes in
ecosystem N stocks, changes in the internal N cycle in response to elevated

CO2 are expected to reduce N availability to plants, and therefore to constitute negative feedbacks to enhanced primary and ecosystem productivity.
Such progressive nitrogen limitation does occur in some CO2 enrichment
experiments in grasslands, but it is not apparent in others, possibly because
the productivity responses are too small to elicit strong feedbacks, because
year-to-year variation in climate masks them, or because CO2 increases N
inputs or decreases N losses thereby enhancing ecosystem N capitol and
relieving progressive N limitation. In this talk, I will review grassland CO2
enrichment experiments in light of the concept of progressive N limitation.
HUNTZINGER, MIKAELA1,* and DAVID AUGUSTINE.2 1 University of
California, Davis, CA; 2 Syracuse University, Syracuse, NY. Disparate
herbivore lineages not functionally redundant.
Herbivores can affect the aboveground biomass of plant communitiesbut
are those effects taxon-specific, or are disparate herbivore taxa functionally
redundant? If one herbivore taxon is removed from a system, can another
taxon feeding on the same food resource compensate in biomass consumption? In East African shrub savanna, grasshoppers rely on the same food
resources as ungulates such as impala, cattle, zebras, and buffalo. Grasshoppers were compared in paired, replicated ungulate exclosures in adjacent bush and glade habitats in East Africa to determine whether they could
compensate for the excluded ungulate herbivores. After two years, grasshoppers were sampled in two seasons at all sites and then dried and
weighed. Grasshopper biomass was used as a measure of competition with
ungulates. Grasshoppers increased by 78-256% in the absence of ungulate
herbivores, suggesting that they are suppressed by ungulates. Thus, the
increase in grasshoppers could potentially allow for compensation by the
grasshoppers for the excluded herbivores. However, there is a 117-154%
increase in the peak biomass of Cynodon sp., the monodominant grass,
where ungulates are excluded. Therefore, even when released from competition, these insect herbivores cannot consume as much biomass as large
mammalian herbivores. Thus, different suites of herbivores may not be
functionally redundant even when they compete for shared resources. These
results are important because if habitats are not managed to retain their
original suite of herbivores, changes in the other herbivores and in the plant
community may be inevitable. If plant biomass increases sufficiently when
mammalian herbivores are eliminated, changes in detritivore dynamics or
fire regimes are likely.

mean of 22 percent and 40 percent on Cache and Coosawhatchie, respectively; high concentrations of organic material may have important implications for global carbon cycling.
HURTON, LENKA V.,1,2,* ELIZABETH A. WALLS1,2 and JIM BERKSON.1,2 1 Virginia Polytechnic Institute and State University, Blacksburg,
Virginia; 2 Horseshoe Crab Research Center, Blacksburg, Virginia. Postbleeding mortality of Horseshoe Crabs, Limulus polyphemus, used in
the biomedical industry.
The horseshoe crab is a living fossil, whose existence can be traced back
at least 200 million years. Many aspects of the ecology, life history, and
physiology of the horseshoe crab are unique. The blood of the horseshoe
crab is copper-based, rather than iron-based. Biomedical companies extract
blood from horseshoe crabs for the production of Limulus Amebocyte Lysate (LAL). This purified extract from Limulus hemocytes is used worldwide for detecting the presence of endotoxins in medical and pharmaceutical products. The biomedical companies catch the crabs, bleed them, and
then return them to the ocean. In order to better understand the persistence
of the horseshoe crab, we have worked with BioWhittaker, the largest producer of LAL, to study the effects of blood loss. In the first research component, we measured the amount of blood removed from 98 male horseshoe
crabs. Mean weight of blood extracted was 78.3 g. In the second component, we estimated the post-bleeding mortality rate of male horseshoe crabs
from the biomedical process. Results indicate mortalities to be approximately 7.5%. In the third component, we estimated the total blood volume
of horseshoe crabs over the horseshoe crabs adult size range. An average
sized male horseshoe crab weighing 1 kg was found to have 251 ml of
blood. Results of these studies show that despite losing an average of 31%
of their blood, the post-bleeding mortality rate is quite low. The horseshoe
crab has been in existence in its present form for over 200 million years.
We believe that its ability to survive a significant loss of blood is one of
the unique adaptations that has allowed its continued persistence, while
other species have evolved and/or gone extinct through the millennia. We
will discuss other physiological and ecological adaptations thought to contribute to the horseshoe crabs persistence as well.

HUPP, CLIFF R.* and SAMMY L. KING. U.S. Geological Survey, Reston,
Virginia, USA. Sediment trapping along brown- and blackwater Coastal Plain riparian forested wetlands.

HUTCHINSON, TODD F.,1,* ELAINE K. SUTHERLAND2 and DANIEL

A. YAUSSY.1 1 USDA Forest Service, NERS, Delaware, OH; 2 USDA Forest Service, RMRS, Missoula, MT. Can oak forest structure and sustainability be restored with prescribed fire alone?

Sediment trapping is cited as a major function of riparian-forested wetlands.

Few studies have compared local sedimentation processes on brown- versus
blackwater systems. Sedimentation rates were estimated along the brownwater Cache River, Arkansas and the blackwater Coosawhatchie River,
South Carolina along transects normal to streamflow, using dendrogeomorphic (tree ring) and clay-pad marker techniques. Rates were related to
several physical parameters, woody vegetation, and its debris. Mean deposition ranged from 0.20 to 0.36 cm/yr and from 0.02 to 0.20 cm/yr on
the Cache and Coosawhatchie, respectively. The Cache carries a suspended
load (TSS) of about 100 to 350 mg/l, whereas the Coosawhatchie carries
about 5 to 25 mg/l. Sedimentation patterns varied strongly with elevation
and thus, vegetation patterns. Hydraulic connectivity (degree of flow-path
connections to the main channel) appears to strongly affect sedimentation
rates; greatest deposition occurs along sloughs with a direct flow path to
the river. Whereas, low areas poorly connected to the channel have relatively low deposition rates, presumably due to diminished replenishment
of suspended sediment during the hydroperiod. Sedimentation rates are
highest in and near sloughs and their anabranches and lowest in areas
unaffected by sloughs even though both floodplains are largely inundated
during peak hydroperiod. Woody vegetation, including woody debris, may
also play an integral part in concentrating flow paths. Low velocities facilitate deposition of fines, particularly organic material, however relatively
moderate velocities may ensure a continuous supply of sediment-laden water, particularly mineral fines. The Cache (187.6 kg/ha/yr) and the Coosawhatchie (24.5/ha/kg/yr) trap substantial amounts of sediment, reinforcing
the water-quality functions of these forested wetlands Fines deposited over
the clay pads contained substantial amounts of organic material with a

Witness trees recorded in the original land surveys (ca. 1800) indicate that
oaks dominated presettlement upland forests in southeastern Ohio. Forest
density estimates suggest that open woodlands (, 100 trees/ha) were uncommon but low-density forests (100-200 trees/ha) were the modal frequency class. Through the 1800s, nearly all forests were harvested, and
fire was frequent until suppression began in 1923. After .75 years of
effective fire suppression, current forests are more dense and gradually
succeeding to dominance by shade-tolerant species (e.g., maples). In 1995
we initiated a long-term study at four 75 ha sites to determine the effectiveness of prescribed fire to improve oak regeneration and thus sustainability of the ecosystem. Most stands originated ca. 1870-1910. Prior to
the reintroduction of fire, oaks comprised 67-72% of tree basal area and
median tree density was 368 trees/ha. Shade tolerant species (e.g., maples)
were abundant in the midstory and comprised .85% of saplings. Annual
(1996-1999) and periodic (1996 and 1999) low-intensity fires were conducted on units at each site and an unburned unit served as a reference.
By 1999, sapling densities were reduced by 85% on burned units, whereas
overstory trees were largely unaffected. Hemispheric photographs taken in
1998 indicated that light availability was not significantly affected by fire.
Without significant changes in overstory tree density and light availability
on burned units, there was no significant difference in the relative abundance of large (. 30 cm height) oak seedlings among treatments by 2002.
Large seedlings of shade-tolerant species were more abundant than oaks,
and did not differ among fire treatments. We conclude that even repeated
burns are inadequate to reduce forest density and improve oak regeneration
in long-unburned forests of this region.

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161

HUXEL, GARY R.,1,* DREW M. TALLEY1,2 and FRANCISCO SANCHEZ-PINERO.3 1 University of South Florida, Tampa, FL; 2 University of
California, Davis, Davis, CA; 3 Universidad de Granada, Granada, Andalucia, Spain. Ecological stoichiometric differences on islands in the Gulf
of California.
Our previous research has demonstrated significant differences among tenebrionid beetles on islands in the Gulf of California. The islands vary is
use by seabirds and are classified as non-bird, roosting, and nesting islands.
On roosting and nesting islands, seabird carcasses are the primary carbon
sources, while on non-seabird islands plant detritus is the major resource
of carbon for the beetles. However, significant differences in nitrogen and
phosphorus concentrations between plant and carrion tissues, and within
and across island types exist. We address whether these differences influence patterns of resource flow. We find that on non-bird islands while C
comes mostly from plant tissue, C contributes the vast majority of nitrogen
and phosphorus. On nesting islands, C, N, and P are all mostly derived
from carrion resources. However, on roosting islands because plant tissue
reaches its highest density, and N and P concentrations are the highest in
plants, C, N, and P are derived nearly equally from plant and carrion tissues. These differences in stoichiometry have significant influences on the
diversity of tenebrionid beetles (both alpha and beta diversities) on islands
in the Gulf of California.
HUXMAN, TRAVIS E.* Ecology and Evolutionary Biology, University of
Arizona, Tucson, AZ. Plant invasions and elevated CO2: interactions
with climate variability and plant growth form.
Increasing concentrations of atmospheric CO2 influence energy and material exchange between plants and their environment, altering a number of
important plant functional and morphological traits. In semi-arid and arid
ecosystems, there has been focus on how CO2 influenced changes in two
traits may alter the ability on non-native species to invade novel habitats;
growth rate and stress tolerance. Changes in plant growth rate and stress
tolerance have been used to understand the potential for plants to extend
their ranges into new systems, but have not consistently provided the
framework for general prediction. What has emerged as important factors
are the interactions between the direct effects of elevated CO2 on plants
and variation in other climate variables. Additionally, how shifts in growth
rate interact with patterns and plasticity in plant development may significantly influence invasive ability. Including these interactive effects of climate and secondary effects of changes in growth rate into our conceptual
and experimental frameworks may be the key to understanding the relationship between invasive species performance and global change.
HYATT, LAURA1,2 and JESSICA GUREVITCH.1 1 SUNY Stony Brook,
Stony Brook, NY; 2 Rider University, Lawrenceville, NJ. Exotic and native plant responses to nutrient amendments in the field.
Our observational work on Long Island, NY has revealed that exotic, invasive species appear far less frequently in nutrient-poor pine barrens habitats than in adjacent nutrient-rich, mesic hardwood forests. This work has
also shown that exotic invaders are not only strongly associated with high
plant diversity, but are also found more frequently on nutrient-rich soils.
We experimentally tested the hypothesis that low resource availability prevents exotic species from invading pine barrens forests through a series of
field manipulations. In uninvaded barrens forest plots, we altered light
availability, Ca and N in a crossed design. Into these plots we planted
seedlings of both native and exotic species. Overall, exotic species were
more likely to establish and grow than native species, a result that supports
the notion that successful invaders are more vigorous than natives. However, adding nutrients did not enhance survival or growth for exotics or
natives. Under nutrient addition, exotic species were more likely to shrink
in size while native species were more likely to die. This effect was especially pronounced in N-addition plots. We suspect this pattern is partly
due to the interaction of selective herbivory by insects and mammals and
study species defense mechanisms. Individual species responses to nutrient additions and the interactive effects of light and nutrients varied strongly. Our results suggest that community immunity to invasion is a mechanistically complex attribute that bears further experimental exploration.

162

Abstracts

N
EZ, INES,* JAMES S. CLARK and SHANNON L. LADEAU. Duke
IBA
University, Durham, North Carolina. Climate variability and tree species
recruitment, implications on species coexistence,.
Climate variability will affect natural plant communities, in part, through
its impact on recruitment dynamics. In this study, we estimated the differential influence of climatic patterns on seedling establishment of several
dominant tree species. Using ten years of data on climate, seed rain, and
seedling censuses, we develop a hierarchical Bayes model to quantify establishment of dominant species at five stands of secondary temperate forest in the Southern Appalachians. Establishment (logistic regression) was
estimated as a function of environmental variables (temperature and soil
moisture) and adjusted for seed rain. Coefficients linked with each climatic
variable are reported here. Years having the lowest minimum temperature
during the germination season resulted in poor establishment of the two
Betula species (coefficient mean6SD: -0.18860.08), both of which occur
primarily in the upper elevation stands. On the other hand, low spring
temperatures enhanced recruitment of Acer rubrum (0.4760.02) and Nyssa
sylvatica (0.5860.14), species that grow in warm, low-elevation stands.
Quercus rubra establishment was also most successful in years with low
temperatures (0.3260.14). In addition to temperature, precipitation (measured as soil moisture content during the germination season) affected seedling establishment. Betula spp. (0.0960.008), Liriodendron tulipifera
(0.2560.08), and Q. rubra (0.1360.03) experienced enhanced establishment during years with relatively high precipitation. N. sylvatica, a species
that occurs primarily on dry sites in our region, established more successfully during dry years (-0.1860.03). Results from this study confirm the
influence of climate variability on seedling establishment, and its distinct
effect on particular species.
IGNACE, DANIELLE D.* and TRAVIS E. HUXMAN.* University of
Arizona, Tucson, AZ, USA. The effects of soil surface on community
structure and plant function of Larrea tridentata in the Sonoran Desert.
In the Sonoran Desert, strong plant community and geomorphic soil associations have been illustrated. We evaluated the ecophysiological patterns
that underlie these plant and soil interactions by measuring various aspects
of plant function seasonally for Larrea tridentata on previously characterized Pleistocene and Holocene surfaces at Tumamoc Hill, southern Arizona.
The Holocene soil surface is characterized as sandy with no strong horizonal development, while the Pleistocene surface is a soil that effectively
reduces rooting volume. Larrea on the Holocene have greater canopy volumes compared to Larrea on the Pleistocene soil surface. During the monsoon, Larrea on the Holocene had higher fluorescence, leaf nitrogen content and less negative predawn water potentials than the Pleistocene Larrea,
but there were no differences in leaf photosynthesis and stomatal conductance. Following the monsoon, Larrea on the Pleistocene surface experienced greater water stress than the Holocene Larrea. Despite differences
in water stress on the different soil surfaces, assimilation rates were statistically insignificant and slightly negative. There was greater mortality of
Larrea during the dry 2002 growing season on the Pleistocene soil surface
as compared to the Holocene site. The results may show how soil surface
affects plant function and may shape community structure during episodic
drought.
IMHOFF, MARC L.,1 LAHOUARI BOUNOUA,1,2 TAYLOR H. RICKETTS,3,4 COLBY LOUCKS,3,* ROBERT HARRISS2 and WILLIAM T.
LAWRENCE.5 1 NASA Goddard Space Flight Center, Greenbelt, MD;
2
University of Maryland, College Park, MD; 3 World Wildlife Fund, Washington, DC; 4 Stanford University, Stanford, CA; 5 Bowie State University,
Bowie, MD. Global distribution of human NPP appropriation.
Human population and consumption profoundly affect global ecosystems
in numerous ways. One method of measuring the aggregate impact of human activities is to estimate the amount of total net primary production
(NPP) we appropriate for our own use. While previous studies have estimated this fraction for the globe, here we develop spatially-explicit maps
of both terrestrial NPP and of human NPP appropriation that allow spatial
analysis of human impacts (based on a 0.25-degree grid). To estimate human appropriation, we multiply the population density of each grid cell by
country-specific consumption factors, gleaned from FAO statistics and

back-calculated to annual per capita mass of carbon consumed. We estimate

global NPP to be 56.8 Pg C per year, and human NPP appropriation to be
9-20% of this total, both of which correspond reasonably well to previous
global estimates. More importantly, the geographic distribution of NPP
appropriation is extremely heterogeneous, due to global patterns of both
population density and per capita consumption. In some areas, humans
consume several times the local NPP 9supply,9 while in others we appropriate only a fraction of a percent of local NPP. In general, areas of high
consumption do not correspond to the most productive regions of the world.
Comparisons of NPP appropriation with global distribution of biodiversity
indicate areas of maximum conflict between humans and other species for
the products of photosynthesis.
IMM, DONALD W.1,* and KEN W. MCLEOD.2 1 USDA-FS Savannah
River, New Ellenton, SC; 2 Savannah River Ecology Laboratory, Aiken,
SC. Fire and twenty years of change in a Carolina sandhill.
Sandhill woodlands are fire-prone communities with xeric, infertile sands
in the southeastern coastal plain. Unlike pine savannas, fires in sandhills
are less frequent due to limited litter production and varied fire behavior
due to fuel quantity and presence of a hardwood mid-story. Over the past
20 years, three controlled fires and one escaped fire have occurred in the
study site, dominated by few scattered longleaf pine and large oaks in the
canopy and a near-continuous mid-story of turkey oak, bluejack oak, sand
post oak, and other hardwoods. Based on tree cores, diameter growth of
longleaf pine was reduced during the first year following summer burning
but unaffected by winter burning. Oak diameter growth was reduced for 13 years following burning. Cumulative oak mortality increased with each
successive fire. Growing season fires led to greater top kill and mortality
than dormant season fires. Probability of oak mortality and top kill, both
a function of size, decreased with each successive fire. Eleven dominant
ground cover types were present, each with a different composition, environment, and response to burning. Dwarf huckleberry, oak-sprout, diffuse-herbaceous, and matrix-grass groundcover types increased in area following burning. Dense-herbaceous, deerberry, poison oak-mixed herbaceous, and bracken fern ground cover did not significantly change following fire. Dense litter and sparkleberry ground cover declined following fire.
Immediately following fire, ground cover increased in diversity, but compositional changes were temporary (no significant difference beyond 2
years). However, with each successive burn, composition was less variable
in trajectory and less resilient to return to pre-burn ground cover conditions.
These results suggest that sandhill woodlands are slow to change using
prescribed fire, independent of season.
INGRAM, ELLA L.* and MAXINE A. WATSON. Indiana University,
Bloomington, IN. Seasonal change in the composition of the mycorrhiza
of mayapple, Podophyllum peltatum.
Mutualisms can vary seasonally in the intensity of interactions among participants or in the benefit participants derive from the association. For arbuscular mycorrhizas, the relative presence or absence of mycorrhizal structures such as arbuscules and vesicles can indicate seasonal changes in the
nature of the mutualistic interaction. To understand how mycorrhizal function changes through time, we examined seasonal variation in hyphal colonization and in arbuscule and vesicle presence (also measured as % root
length colonized (%RLC)) within the roots of the clonal perennial mayapple, Podophyllum peltatum. Hyphal colonization and arbuscule presence
peaked in spring and fall, while vesicles were relatively abundant only in
fall. Arbuscule presence (and in fall, vesicle presence too) peaked first,
followed one month later by a peak in hyphal colonization. The levels of
hyphal peaks were roughly equivalent in spring and fall, as were the levels
of arbuscular peaks. During peak times, arbuscules accounted for at least
40% RLC. Vesicles always were present at low frequency, peaking at
,10% RLC. We conclude that spring and fall are particularly active times
of resource exchange in the mayapple mycorrhiza, with these active periods
of exchange followed by rapid hyphal growth. In contrast, resource storage
within the mayapple mycorrhiza occurs more in the fall than during other
times in the year. Our results suggest that examination of hyphal colonization alone, or total root colonization alone, may obscure important aspects of the functional dynamics of mycorrhizal associations.

INNIS, ANNE F.,1,2,* IRWIN N. FORSETH1 and DENNIS F. WHIGHAM.2

1
University of Maryland, College Park, MD; 2 Smithsonian Environmental
Research Center, Edgewater, MD. Reproductive life history traits and
competition as explanations for the success of the invasive perennial
Rubus phoenicolasius.
There is a growing presence of invasive species in global ecosystems. Invasive species displace native species, threaten biodiversity and alter the
function of ecosystems. Determining both the characteristics of invasive
species and those of the invaded habitat will enhance management and
prevention of further invasions. The most effective studies of invasive species involve a comparison of closely related invasive and native species.
This study used two perennial Rubus congeners, an invasive species, wine
raspberry (Rubus phoenicolasius), and a native species, sawtooth blackberry (Rubus argutus). Rubus phoenicolasius is invading fields and successional forests, R. argutus habitat, along the Atlantic Coast of the United
States. In 2002 two studies were conducted at the Smithsonian Environmental Research Center, Edgewater, MD. In the study of reproductive life
history traits, 40 individuals of each Rubus species were examined for selfpollination by excluding pollinators, with pollen bags, and recording fruit
set. R. phoenicolasius was shown to self-pollinate more than R. argutus.
During the growing season, 901 individuals of R. phoenicolasius and 1159
individuals of R. argutus were surveyed for age and reproductive effort.
Of the mature individuals, R. phoenicolasius were more likely than R argutus to be reproductive. R. phoenicolasius produced greater numbers of
fruit with more seeds per fruit than R. argutus. Self-pollination, high fruit
and seed set of R. phoenicolasius may drive rapid population growth. In
the competition study, 72 1m2 plots with existing populations of Duchesnea
indica were used to determine the impacts of R. phoenicolasius and R.
argutus on local species. Planned comparisons between the two Rubus
treatments showed no significant difference in the response of D. indica.
The drought that was in effect in Maryland may have suppressed growth
of the three species, thus masking any competitive interactions. With the
worldwide problem of invasive species, it is necessary to understand what
characteristics define invasive species and quantify the effects of invasive
species on plant communities.
INOUYE, BRIAN D.* Florida State University, Tallahassee, Florida. Invasive ants and gall-wasp mutualisms and commensalisms.
Oak trees host diverse communities of cynipid gall wasps, parasitoids and
inquilines. Ants can mediate several interactions among gall wasps and
their parasitoids found on Quercus lobata and Q. douglassi. I present experimental and observational data on a mutualism between ants and a cynipid wasp whose galls secrete honeydew, and an indirect commensalism
with a different species of cynipid wasp that benefits from the presence of
ants but provides no reward. Using ant exclusions, I show that the presence
of tending ants lowers the rate of attack by a diverse assemblage of parasitoids, and also changes the set of species that can successfully use different galls. Additionally, the species of tending ants (Linepithema humile
vs Liometopum occidentale) affects the overall rate of parasitoid attack and
the proportions of different parasitoid species that emerge from galls. Invasive argentine ants fulfill a similar role to native ants in some mutualisms
(but not all), but have a much weaker interaction than native ants in an
indirect commensalism with a gall wasp.
INOUYE, DAVID W.1,2 1 University of Maryland, College Park, MD, USA;
Rocky Mtn. Biological Laboratory, Crested Butte, CO, USA. Modification of plant-pollinator interactions through consequences of global
and regional climate changes.
2

A 30-year study of the timing and abundance of flowering at 2,900m in

the Colorado Rocky Mountains shows that there is great variation in both
of these aspects of flowering, and that much of it can be explained by
abiotic factors. Timing of flowering for all species studied to date is explained by the timing of snowmelt in the spring, which is largely a function
of how much snow fell the previous winter. For several species the abundance of flowering is strongly correlated with depth of winter snowpack,
either through some mechanism yet to be discovered, or in some cases
mediated by the interaction between time of snowmelt and probability of
late spring frost damage to flower buds. Sometimes most flower buds of

Abstracts

163

some species are killed by frost, with consequences for flower visitors,
herbivores, seed predators, and their parasitoids. Variation in snowpack and
snowmelt in this part of the Rockies is influenced strongly by global factors
such as climate warming and regional influences such as the El NinoSouthern Oscillation and the recently discovered North Pacific Oscillation.
The NPO has about a 50-yr cycle, and much of the study presented here
was encompassed by the last wet phase (starting in 1977). A 1998 phase
change of the NPO appears to have triggered an abrupt decline in winter
precipitation, with effects (primarily negative) on flowering by many species. Winters since 1998 have had declining snowfall, which has resulted
in reduced flowering by many species (through a combination of frost
damage and unknown mechanisms). Climate change is also changing the
synchrony of plants and pollinators, as the plants and animals at this site
are apparently responding to different environmental cues or respond differently to the same cues.
IRVINE, ROBYN L.* University of Calgary, Calgary, Alberta, Canada.
Detecting disturbance in aquatic ecosystems in theory and in the field:
Matching variance and experimental design.
The magnitude and structure of temporal and spatial variance have direct
ecological importance for how clearly community patterns in nature can be
seen, and how large perturbations have to be to move systems beyond
background variation. Here, I examine the spatial and temporal variability
of benthic communities in montane stream ecosystems and consider the
implications of the communities variance to the detection of disturbance.
Benthic invertebrates inherently vary through space and time, so I pose
two questions: 1) what size of disturbance can be reliably detected using
changes in invertebrate community structure? and 2) does the spatial and
temporal structure of the variance in the benthic community make one
experimental design more sensitive to detecting a change than another? I
conducted my study in the headwaters of the McLeod River, Alberta, Canada. Benthic community data were collected from the five streams that
comprise the headwaters in 1985, 1993, 1995, 1999, and 2000 to estimate
the spatial and temporal structure of the variance in the benthic community.
I contrast three experimental designs to assess each designs ability to detect changes in these streams given the variance inherent in the systems
and using a disturbance effect size drawn from the literature. Specifically,
I compare a BACI design (Before After Control Impact), a paired ControlTreatment design, and a multiple treatment (4), one contro1 design. The
results of my work have direct management implications, because these
streams sit on a site slated for an open-pit coal mine. Benthic invertebrates
are a key indicator of water quality employed in Canadas Environmental
Impact Assessment (EIA) procedures.
ITO, AKIHIKO,1,* WILFRED M. POST2 and GEORGE A. ALEXANDROV.3 1 Frontier Research System for Global Change, 3173-25 Showamachi, Kanazawa-ku, Yokohama, Kanagawa, Japan; 2 Environmental Sciences Division, Oak Ridge National Laboratory, Oak Ridge, Tennessee;
3
Center for Global Environmental Research, National Institute for Environmental Studies, Tsukuba, Ibaraki, Japan. Simulation of Carbon Dynamics and Fire Regimes in Boreal Forests of Eurasia and North
America.
Conifer forests increased exploitation and climatic change make modeling
structure and function increasingly important. Sim-CYCLEs five carbon
pools and 16 fluxes simulate production, allocation, and decomposition
ecophysiologically. Simulations of boreal forests in Russia and Alaska now
include burning. Fire frequency, severity, and extent are simulated with a
cellular automaton for 100 km2 (200 rows and 200 columns). East Siberian
(62N, 119E) larch, a deciduous conifer, compares with evergreen conifers,
e.g. spruce, near Fairbanks (65N, 147W). Wildfires increase heterogeneity
of carbon budgets. Siberian fire recurrence interval averages 50 years, restricting carbon storage, despite slow decomposition. The cellular automaton simulates spatial heterogeneity in landscapes. Over 600 years, wildfire
can reduce regional C-storage by 20%. But prescribed fire suppression may
not increase storage, because fire severity increases with fuel accumulation.
Spatial heterogeneity (sub-grid scale for global models) should be included
in continental studies, using parameterization and GIS weighting. Besides
the subroutines concerned with fire and its spatial relations, cells have a
compartment for debris (including standing dead) that is separated from

164

Abstracts

other litter, mostly more decomposable. Charcoal is a (nearly) inert part of

soil carbon, analogous with "passive" humus in Posts Global Terrestrial
Ecosystem Carbon (GTEC) model template. GTEC separates decomposers
(bacteria, fungi; possibly soil animals) from their substrates, and coarse
roots (.2mm?) from fine ones. GTEC grid simulations suggest Ontarios
boreal forest and Great Lakes conifer, mixed and "northern hardwoods"
region as a credible region for part of the "missing sink" of excess greenhouse gases. Local "LoTEC" and SimCYCLE template performance may
be compared for forest types and zones of that region or for larch and
mixed forests in Hokkaido (Sapporo eddy flux covariance tower).
IVERSON, LOUIS R.,1,* ANANTHA M. PRASAD1 and MARK W.
SCHWARTZ.2 1 USDA Forest Service, Delaware, OH, USA; 2 Department
of Environmental Science and Policy, University of California, Davis, CA,
USA. Modeling potential suitable habitat and migration of trees in the
eastern United States using forest inventory data and contrasting climate scenarios.
We automated species range maps compiled by Elbert Little in the early
1970s and summarized current forest inventory data, at a resolution of 20
x 20 km cells, for 135 species in the eastern United States to enable generation of tree biodiversity hot spot maps. The 135 species represent almost
all tree species in the east that have a distribution range of at least 100 20
x 20 km cells, and do not include many endemic species from Florida.
Highest diversities are present in the southern Appalachians and in the
lower Ohio Valley. We next used a statistical model, DISTRIB, based on
recursive partitioning, to estimate changes in potential suitable habitat for
each of the species, assuming equilibrium of 2xCO2, under two climate
change scenarios: the Canadian Climate Center model (CCC) and the Hadley Climate Center model (Had). These were the two scenarios used by the
U.S. National Assessment for Climate Change. Summarizing these outputs
allowed us to compile how the suitable habitat may change for this collection of tree species, and whether the biodiversity hot spots may eventually
shift. As expected, the more severe climate change scenario, CCC, resulted
in greater potential shifts in suitable habitat. Next, we ran a spatially explicit cellular automata model, SHIFT, to estimate colonization probability
via migration, by 1 km cell, over the next 100 years, on five southern
species (persimmon, sweetgum, sourwood, loblolly pine, and southern red
oak). By evaluating the probability of colonization (via SHIFT) within the
potential new suitable habitat (via DISTRIB), we can estimate the proportion of new suitable habitat that may be colonized within 100 years. This
proportion is very low for all five species, suggesting that migration rates
similar to that occurring through unfragmented forests during the Holocene
will lag behind the rate of climate change.
IVEY, CHRISTOPHER T.,1,* DAVID E. CARR1 and MICKY D. EUBANKS.2 1 University of Virginia, Boyce, VA; 2 Auburn University, Auburn, AL. Inbreeding alters tolerance and resistance to herbivory in
natural settings.
Inbreeding is common in natural plant populations and is one of the most
fundamental influences on plant biology. A few recent studies have reported that inbreeding also can alter ecological interactions with other species. We performed two experiments to test the effects of inbreeding on
tolerance and resistance to herbivory in four populations of Mimulus guttatus in Napa County, California. Self- and outcross-fertilized plants were
transplanted into field sites and scored for herbivore damage (resistance)
and fitness at the end of the growing season. In a second experiment, selfand outcross-fertilized plants were transplanted into field sites, and half the
plants were exposed to herbivory from a single spittlebug (Philaenus spumarius) nymph. Inbreeding increased resistance to herbivory in one population but had no effect in the other three. Inbreeding reduced tolerance
to spittlebug herbivory in two populations, increased tolerance in a third,
and had no effect in the fourth population. The presence of herbivores
significantly magnified inbreeding depression for plant biomass in two populations and reduced inbreeding depression for plant developmental rate in
a third population. Variation among populations in inbreeding effects is
common and may reflect differences in inbreeding history. We also observed significant variation among plant families for inbreeding effects on
tolerance, indicating the opportunity for selection by herbivores to improve
population-level tolerance. These results emphasize that the indirect effects

of inbreeding on fitness through altered ecological interactions are important to consider, especially for the management of small and fragmented
populations where inbreeding risk is higher.

JACKSON, ROBERT B.* Duke University, Department of Biology and

Nicholas School of the Environment, Durham, NC, USA. Bioigeosciences
and the elements of interactions.

IYER, MEERA,* RICHARD K. KOBE and MICHAEL B. WALTERS.

Michigan State University, East Lansing, MI. Carbon allocation in roots:
to forage or for storage?

Since Haeckel coined the term ecology more than a century ago, the physical environment has played a central role in ecological theory and ideas.
The sophistication with which we can monitor the environment today, and
the sheer volume of environmental information available, highlights the
need for stronger links between ecologists and physical scientists. The new
Biogeosciences section within ESA is a welcome step in this direction. In
this presentation I outline three kinds of interactions that are important for
the success of the new section. The first acknowledges the importance of
material interactions in the field, including links between the global carbon
and nitrogen cycles and between water and nutrient cycling. The second
centers on interactions with other scientists and societies, such as the American Geophysical Union. The last highlights the importance of education
and public outreach in our interactions with the public. Through all three
interactions, we can more successfully understand the basic ecology of the
earth system and provide the answers to pressing societal problems of the
environment.

To understand the functional significance of intra- and inter-specific variation in carbon allocation to storage versus soil resource uptake, we examined total nonstructural carbohydrates (TNC) and specific root area
(SRA) in the roots of seedlings of eight northern hardwood tree species.
Seedlings were grown in a greenhouse under two levels of light (2% and
22% full sun) and nitrogen availability (0.5 mg/L and 50 mg/L of N in
a modified Hoagland solution). These resource levels span the range of
variation found in northern Michigan forests. Seedlings were harvested 12,
40, 85 and 105 days after germination. SRA (root surface area / total root
mass) showed no change with nitrogen availability in most species although
it increased under low light, suggesting greater foraging for nutrients under
lower light. However, the usual calculation of SRA is confounded by TNC:
we expect stored reserves to be less relevant to nutrient uptake than structural root mass. We therefore introduce the concept of structural specific
root area (SSRA), calculated by normalizing root surface area with only
root structural biomass (hence excluding TNC pools which made up 25 to
80% of root mass across species and treatments). SSRA provided a different picture of plant strategies. Species associated with low-fertility sites
such as Quercus velutina and Q. alba maintained constant allocation to
foraging (i.e. SSRA) across light and nutrient levels, but increased TNC
under low nitrogen availability. Acer rubrum, often associated with intermediate and higher fertility sites, strongly increased SSRA under low nitrogen levels. Light-demanding Betula papyrifera maintained high SSRA
across nitrogen levels and had the lowest stored reserves among the species
examined. Our results demonstrate the importance of distinguishing between root biomass allocation to structural and labile carbon pools in understanding the functional significance of both root foraging and carbon
storage.
JABLONSKI, LEANNE M.,1,* TARA C. POLING,1 MICHELE G. BANKER,1 ROBERT A. KLIPS3 and DONALD R. GEIGER.1,2 1 Marianist Environmental Education Center, Dayton, OH; 2 University of Dayton, Dayton, OH; 3 The Ohio State University, Marion, OH. Soil and vegetation
factors influencing establishment of conservative vegetation on a restored prairie.
Herbaceous and moss vegetation have been assessed periodically since the
reclamation of eastern tallgrass prairie on a sand and gravel borrow pit at
Mount St. John in Greene Co. near Dayton, Ohio. Prairie was initiated in
1986 on the 7-ha. semi-urban site, by a uniform hydro-seeding with eight
native grasses, and by transplanting native forb seedlings. Two permanent
transects of 350 meters total length were established in 1995, encompassing
a gradient in elevation and edaphic factors with 88% of the sampled area
being dry prairie and 12% in wet prairie. Soil nutrients, organic matter, pH
and mycorrhizae and the herbaceous vegetation composition and cover
were all assessed beginning in 1995 in 36 1.0 m2 quadrats. By 2001, transects represented 54% of the total herbaceous species diversity on the dry
prairie on the soil which ranged from fertile agricultural soil with 100%
vegetation cover to a bottomland pit with pH . 8 and 62% cover. Moss
species, began to appear in 1999, with cover being 26.2% in 2001, and
moss diversity was quantified in 2003. The relationship between edaphic
and vegetation factors associated with quality and their change through
time was examined to explore the role of soil, moss and herbaceous species
associations in successful species establishment. Factors associated with a
higher quadrat level species conservatism included soil moisture and organic layer development, and herbaceous species diversity. Biomass increases did not correlate with diversity or conservatism. Our results suggest
that even with sub-sampling of small areas, repeated monitoring of soil
and vegetation can assist understanding of the factors contributing to successful establishment. Given limited personnel and seed source resources,
knowledge of the natural succession of species in a restoration provides
important guidance to land managers regarding the best timing of species
introductions to enhance the quality of the prairie community.

JACKSON, STEPHEN T.* University of Wyoming, Laramie, WY. Environmental change, climate variability, and biotic interactions in the
assembly of late Quaternary plant communities.
Paleoecological studies of the late Quaternary reveal continual forcing of
vegetational changes by climatic change. Plant species have invaded new
landscapes and abandoned others, populations have expanded and contracted, and species associations have come and gone. All of these phenomena
are predictable from the temporal plasticity of multivariate environmental
space and the differing niche structures of species in environmental space.
The role of biotic interactions in these dynamics is often overlooked or
downplayed by paleoecologists, at least in part because consequences of
biotic processes are hard to discern in the fossil record. However, the paleoecological record includes a number of natural experiments that can
unmask biotic influences. A classic example is the species removal experiment comprising the catastrophic mid- Holocene hemlock decline in eastern North America, which was followed by density compensation and habitat expansion in several tree species. A new set of experiments is emerging
from studies that link independent records of climate variability and vegetational history. Recent studies from both eastern and western North
America indicate that late Holocene species range extensions and population expansions often followed abrupt transitions between dry and wet climate extremes. These patterns probably represent drought-related mortality
of incumbent species, creating opportunities for invasion and expansion of
other species during subsequent wet extremes. Community composition,
while ultimately governed or constrained by the prevailing combination of
environmental variables, may be contingent on the recent and not-so-recent
history of climate variability and disturbance in the context of the available
species pool.
JACOBS, ALYSSA B.,1,* JOHN C. VOLIN1 and ERIC L. KRUGER.2
Florida Atlantic University, Davie, FL; 2 University of Wisconsin-Madison, Madison, WI. Responses of vegetation in the Big Cypress Swamp
to increased inputs of phosphorus and other potentially limiting nutrients.

The Big Cypress Swamp and the Everglades of South Florida are inherently
oligotrophic ecosystems that in recent decades have been subjected to increasing agricultural inputs of phosphorus and other nutrients. Inputs of
phosphorus in particular are thought to have resulted in widespread and
deleterious changes in the structure and function of these ecosystems. This
tenet is supported by extensive research in the herb-dominated Everglades,
but there is a lack of comparable data about the impacts of phosphorus
within the adjacent Big Cypress Swamp. In a 12-month field study we
assessed the responsiveness of Big Cypress vegetation to increased inputs
of phosphorus and/or other nutrients, including potassium and nitrogen.
The study focused on two major tree species, Taxodium distichum and
Fraxinus caroliniana, as well as understory herbaceous vegetation, in the
Kissimmee Billie Slough on the Big Cypress Seminole Indian Reservation.
One of six different nutrient amendments (nitrogen, phosphorus, potassium,

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165

2 x potassium, phosphorus + potassium or control) was applied to the soil

around each of 210 randomly selected trees located throughout the 1,600-ha
slough. During the 12 months following nutrient application, we measured
leaf photosynthesis and nutrient status, as well as stem diameter growth,
on trees. Additionally, we monitored changes in composition, growth and
nutrient content of understory herbaceous communities. Nutrient concentrations in groundwater and/or surface water were also measured at monthly
intervals throughout the year. In general, treatment responses indicated that
the constraint on ecosystem productivity imposed by phosphorus availability may not be as important in the Big Cypress Swamp as it is in
herbaceous ecosystems of the Everglades.
JACOBSEN, ANNA L.,1,* STEPHEN D. DAVIS,1 FRANK W. EWERS,2
KRISTINE A. KERN2 and WILLIAM A. PADDOCK.2 1 Pepperdine University, Malibu, California; 2 Michigan State University, East Lansing,
Michigan. Tradeoffs among xylem efficiency, safety, and strength in
twenty-two species of chaparral shrubs.
Dehydration tolerance and minimal seasonal water potential are often associated with high resistance of xylem conduits to cavitation caused by
water stress in woody plants. It remains unclear what xylem traits, other
than pore size in pit membranes, may enhance resistance to cavitation. Does
increased cavitation resistance require increased carbon allocation to xylem? This question was examined by measuring xylem specific hydraulic
conductivity (Ks), mean hydraulic vessel diameter, water potential at 50%
loss in hydraulic conductivity (C50), and minimum seasonal water potential
(Cmin) relative to wood density, stem modulus of rupture (MOR), and estimated vessel implosion resistance [(t/b)h2] in twenty-two species of chaparral shrubs found in the Santa Monica Mountains of southern California.
Wood density, MOR, and (t/b)h2 were all found to be significantly correlated
with C50 (r2 5 0.46, 0.40, and 0.23, respectively; p ,0.05) and Cmin (r2 5
0.57, 0.58, and 0.23, respectively; p ,0.05). In contrast, MOR and (t/b)h2
were not significantly correlated with Ks and mean hydraulic vessel diameter. Wood density was weakly correlated with Ks (r2 5 0.19; p ,0.05).
These correlations suggest a possible mechanistic relationship between the
mechanical properties of xylem and resistance to cavitation. In contrast,
the mechanical properties of chaparral xylem have little impact on water
transport efficiency. Carbon allocation for structural support of xylem may
represent a considerable cost of xylem cavitation resistance among chaparral species of southern California.
JALICS RAUSCHERT, EMILY S.* and MICHAEL J. RUBBO. The Pennsylvania State University, University Park, PA. Models of salamander
population dynamics and response to human-mediated disturbance.
Amphibian populations have suffered widespread declines and extinctions
in recent decades. Some declines are hypothesized to be the result of mortality caused by factors such as chemical pollutants, direct human habitat
disturbance (i.e. logging), and climate change. However, the populationlevel consequences of mortality have not been examined explicitly, and
may be complicated by density dependence in intervening life-history stages. We examine the population dynamics of 4 salamander species: Ambystoma maculatum, Plethodon cinereus, Desmognathus fuscus and Notophthalmus viridescens under various types of human disturbance. Although
they are not currently declining, these species are ideal model organisms
because they are well-studied and likely to be increasingly impacted by
human urbanization at different life-history stages. We develop demographic models to examine how various forms of human disturbance affect the
relationship among stage-specific survival rates and long-term population
dynamics of the salamanders. Using stage-structured matrix models with
parameters estimated from the literature, we analytically solve deterministic
models. Simulations then allow us to incorporate the influence of environmental variability, including human disturbances and stochastic pond drying. Our results highlight the need to a priori identify critical life stages if
we are to assess accurately how stressors are likely to affect amphibian
populations of conservation concern.
JAMES, JEREMY J.* and JAMES H. RICHARDS. University of California Davis, Davis, California. Seasonal nitrogen capture among coexisting desert plants is differentially affected by water and phosphorous
availability.
In deserts water and nitrogen limit productivity and are mainly available
to plants in brief pulses following rainfall. Consequently, the ability to

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Abstracts

rapidly increase N capture through morphological and physiological adjustments determines, in part, survival and competitive ability. While such
responses are well-described, little is known about how the seasonal timing
and magnitude of N pulses affect plant N acquisition or the extent limitations of other resources may alter plant response to an N pulse. We applied
pulses of 15N labeled K15NO3 at two rates (low, 28 mg N m-2 or high, 5 g
N m-2) three times during the growing season (Early, Mid, or Late Spring)
to the three dominant species at our cold desert study site (two C3 shrubs,
Sarcobatus vermiculatus and Chrysothamnus nauseosus and a C4 grass Distichlis spicata). We also manipulated soil water and phosphorous availability concurrently with these N pulses to determine if plant N capture
during a pulse is affected by the availability of these resources. We quantified plant 15N capture, and soil and microbial N pools for each of the
three pulse periods. All species increased N capture at higher rates of N
addition. Relative to controls, the shrubs did not increase N capture following water addition during any pulse period but N capture in Distichlis
increased 1.6 and 7-fold during the mid and late spring pulses. Chrysothamnus N capture increased 3-fold following P addition in the late spring
pulse but did not increase N capture in the other species. These results
suggest that N capture is not only significantly influenced by both the
magnitude and timing of the N pulse but depends on the availability of
other soil resources. Moreover, these results demonstrate that the ability
these coexisting species to capture N may be differentially affected by soil
water and P availability.
JAROSZ, ANDREW M.* Michigan State University, East Lansing, MI,
USA. Changes in host life history and population dynamics due to
pathogen epidemics.
Pathogen infections are well known for their effects on fitness, which can
ultimately alter life history of the host. These pathogen-induced changes
can have cascading effects by influencing population structure and growth.
My lab has been investigating the population level consequences of epidemics in two systems. Our work on Jack-in-the-pulpit indicates that infection by the systemic rust, Uromyces ari-triphylli, reduces survivorship
and growth. Infected plants also change their pattern of reproduction by
displaying an increased propensity to produce flowers, while reducing the
number of asexually produced cormlets. At the population level this results
in altered size structure within populations and reduced population growth
rates. This decrease in growth rates seems to be exacerbating a trend in
population decline that may be due to habitat fragmentation. We are also
investigating how changes in virulence in the chestnut pathogen Cryphonectria parasitica influence populations of American chestnut. Virulence
is altered by the presence of a double-stranded (ds) RNA hyperparasite that
infects the pathogen intra-cellularly. Reduced virulence in C. parasitica
does not always change population dynamics. Size structure and growth
rates of the chestnut population are determined ultimately by subtle changes
in growth and survivorship of sub-adult plants.
JASTROW, JULIE D.* Argonne National Laboratory, Argonne, IL. Mean
residence time of soil carbon pools: controlling factors and implications
for soil carbon cycling.
Soil organic matter (SOM) is a heterogeneous mixture of plant, animal,
and microbial materials in all stages of decay combined with a variety of
decomposition products of differing ages and complexity. Because the turnover of these components varies continuously, any estimate of the mean
residence time (MRT) for SOM as a whole simply represents an overall
average. The use of whole soil MRT for predicting responses to management practices or environmental perturbations can be misleading because
soils with similar average MRTs can have very different distributions of C
among pools with varying turnover rates. Recent advances in approaches
to fractionating SOM have improved our ability to isolate functionally
meaningful pools tied to the mechanisms that control soil C cycling, and
isotopic techniques enable the MRTs of soil fractions to be estimated and
compared. Even so, predictions of responses to perturbations are problematic because (1) estimates of MRT are made under the assumption of steady
state conditions and (2) the MRT of C in physically or chemically protected
pools is not controlled by its inherent decomposability. Thus, under aggrading conditions relatively new C can accumulate in pools with long
MRTs, and disturbance can accelerate the loss of old C pools by exposing

protected but otherwise labile C to decomposers. Similarly, responses to

environmental changes that influence decomposer activity, such as temperature or moisture, cannot be directly predicted from estimates of pool MRTs
without accounting for effects on the transfer between pools and knowledge
of the saturation levels for protective mechanisms. Hence, simulation models that account for variations in turnover rates for different soil C pools
and the transfers between pools are necessary to generate realistic predictions of soil C dynamics.

lower vertebrates, such as frogs may also be relevant for predicting potential health effects in humans and other aquatic organisms. We are extending
this work to include mammalian models. Studies using rat C6 neural cells
showed that pre-exposure to PCBs causes neuroimmune suppression. Gene
expression profiles were analyzed using Affymetrix microarrays, followed
by quantitative analysis of specific genes with Taqman PCR. Exposure to
PCBs suppressed expression of genes that regulate the neural-immune response, for example, GFAP, SOCS-3, IL-6 and PAI-1.

JAYACHANDRAN, KRISH,* SEEMA SAH, JAY SAH and MIKE ROSS.

Florida International University, Miami, FL, USA. Phosphorus biogeochemistry of the Everglades National Park tree island soils.

JENKINS, DAVID G.,* KATHRYN KIEHL and SUSAN RYAN-SCHNEIDER. University of Illinois at Springfield, Springfield, IL. Local- and regional-scale models of habitat suitability for the Illinois chorus frog
(Pseudacris streckerii illinoensis) in isolated wetlands.

Soils serve as a media for plant growth, exchange and storage of nutrients.
Also, their structural profiles provide a record of soil formation and site
history. Tree Island soils biogeochemistry and nutrient cycling, primarily
phosphorus (P) in the Everglades National Park, FL is of particular interest
to us. The objectives of the soils component of the project as follows: How
are the recent changes in hydrology affecting the vegetation mosaic and
soils? Are these soils functioning similarly to the adjacent marsh soils? Are
these soils nutrient sinks or sources? Do they differ in their elemental
composition? And how does this elemental composition affect vegetation?
With these questions, we describe physical and chemical properties of soils
from three tree islands - Black, Gumbo Limbo, and Satinleaf Hammocks.
Composite surface soil samples at 0 to 10 cm depth were collected from
three tree islands in November 2001. By using a 10 cm diameter soil auger,
several 0-10 cm deep cores were taken at each sampling spot, composited
in a sampling bag, and transported to the laboratory for physicochemical
analysis. Soils were homogenized in the bag and refrigerated until further
analysis. Sub-samples from each bag were transferred to a plastic sample
cup, weighed wet (g), and measured for sample volume (cm3). Later, these
samples were analyzed for dry weight (80 C), sample bulk density (g dry
cm-3), fractional water content, pH, EC, total C, carbonate content, inorganic and organic C, organic matter content, total N, and total P. We have
established baseline physicochemical properties of soils on three tree islands, Black Hammock, Gumbo Limbo, and Satin Leaf. Gumbo Limbo
Hammock seems to be different from Black and Satinleaf Hammocks in
several soil properties. In general, soils in the hardwood hammock communities, particularly the raised part of the community, have alkaline pH,
low organic matter, abundant carbonate minerals, and low N. The P content
seems to be very high in all three tree islands, especially in the hammock
portions. Based on the high levels of total P in these soils, we postulate
several theories on P retention, transport, cycling, and biogeochemistry that
are being tested in our research, with field and laboratory data.

The Illinois chorus frog (Pseudacris streckerii illinoensis) exists in Illinois

only in isolated wetlands among sandy soils on the east edge of the Illinois
River valley. In order to better identify critical habitat for conservation of
this state-listed (threatened) amphibian, we collected a suite of abiotic and
biotic data in wetlands to develop a local-scale model of habitat suitability.
We also collected GIS data across three counties for regional-scale models
of habitat suitability, and sampled subsets of wetlands within the three
counties to test the effect of scale on models. We then developed logistic
regression models to predict Illinois chorus frog presence/absence, based
on the local (in-wetland) data and regional (GIS) data. Among all the abiotic and biotic variables collected in wetlands, the presence of large red
water mites was most strongly and negatively associated with Illinois chorus frog presence (coefficient 5 -12.812, R2 5 0.84, model accuracy 5
94%). Wetlands that support large red water mites appear to not support
Illinois chorus frog populations. Regional-scale logistic models improved
from poor (R2 5 0.2) to good (R2 5 0.8) as spatial scale was reduced from
large (. 250 sites across 7000 m) to small (,20 sites across 10s of meters)
scales. Landscape heterogeneity interfered with predictive models at large
spatial scales and based on GIS data alone. The best predictive models for
Illinois chorus frog habitat are based on neighborhood-scale sets of habitats,
and will be greatly improved by the inclusion of in-wetland data, such as
the presence/absence of red water mites.

JELASO, ANNA M.* and CHARLES F. IDE. Western Michigan University, Kalamazoo, MI. Gene Expression Patterns as Indicators of Health
Effects Induced by Exposure to Environmental Contaminants.
Environmental contaminants disrupt ecosystem homeostasis and produce
numerous health effects in wildlife and humans. Decisions regarding the
clean-up of contaminated sites are complicated by economic, political and
scientific considerations and are often met with much dissent, as it is difficult to establish direct cause and effect relationships between exposure to
specific contaminants and specific health effects for ecosystems and for
humans at a particular site. New genomics based technologies, such as realtime PCR and DNA microarrays that measure changes in gene expression
provide sensitive, molecular tools for improving eco-risk assessment. These
technologies can rapidly and accurately determine not only whether an
exposure to a specific contaminant has occurred, but also the qualitative
and quantitative biological effects related to that exposure. We have developed genomics based technologies to define molecular bioindicators of
exposure to environmental contaminants, such as polychlorinated biphenyls
(PCBs). Studies using laboratory surrogates of aquatic species demonstrate
that gene expression signatures are useful bioindicators of exposure and
are predictors of impending or overt health effects. In studies with developing Xenopus laevis frogs, expression levels of physiologically relevant
genes were measured using real-time PCR (Taqman). Specific gene expression signatures correlated with exposure and/or the appearance of overt
health effects and decreased survivorship. Due to the highly conserved
nature of many genes, genes that serve as health effects bioindicators in

JENKINS, MICHAEL A.,1,* SHIBU JOSE2 and PETER S. WHITE.3 1 National Park Service, Gatlinburg, TN; 2 University of Florida, Milton, FL;
3
University of North Carolina at Chapel Hill, Chapel Hill, NC. Influence
of soil, topographic, and stand characteristics on Cornus florida distribution before and after anthracnose.
In Great Smoky Mountains National Park, heavy losses of Cornus florida
(flowering dogwood) have been observed as a result of infection with Discula destructiva, the fungus that causes dogwood anthracnose. We used
vegetation, soil, and topographic data from 83 permanent forest plots first
established in 1977-79 and resampled in 1995-2000 to examine the influence of soil, topography, and stand structure on the distribution of C. florida
before and after anthracnose infection. According to multiple linear regressions, in 1977-79 (prior to anthracnose infection), C. florida basal area
increased with increasing soil pH (P , 0.001), percent slope (P 5 0.017),
and soil potassium saturation (P 5 0.025). In 1995-2000 (after anthracnose
infection), C. florida basal area increased with increasing soil potassium
saturation (P,0.001) and magnesium saturation (P,0.001), but decreased
with increasing total stand basal area (P 5 0.008). Prior to anthracnose
infection (1977-79), soil, topographic, and stand structure variables explained relatively little of the variability in C. florida basal area (adjusted
R2 5 0.28). However, after anthracnose infection (1995-2000), these variables accounted for a greater amount of the variability in C. florida basal
area (adjusted R2 5 0.44), suggesting that the distribution of C. florida
trees was more dependent upon site conditions following heavy morality
from anthracnose. Although C. florida has been shown to concentrate large
amounts calcium in its foliage, post-anthracnose basal area was not related
to soil calcium saturation (P 5 0.396).
JEONG, KWANG-SEUK,* KYONG HA, MYOUNG-CHUL KIM and
GEA-JAE JOO. Pusan National Univ., Busan, S. Korea. Influence of
drought on the phytoplankton bloom formations in a river-reservoir
system.
This study presents the influence of drought on the phytoplankton proliferations in a river-reservoir system. In the case of freshwater ecology, re-

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167

duction of discharge caused by droughts can cause diverse changes of

ecosystem functions and structures at the point of time. When considering
flow regulation, the river systems may show a different pattern responding
to droughts. The lower Nakdong River in South Korea is a good example
of a eutrophic river-reservoir system which contains multi-purpose dams
and an estuarine barrage at the river mouth (chlorophyll a concentrations,
43.6676.7 L-1, n5385). Due to the monsoon and typhoons, the river had
concentrated rainfall during summer (. 60% of total rainfall in June to
August; average annual rainfall, 1,250 mm). The flow regulation can shuffle
the lentic and lotic characteristics in this river. Ten years (1993-2002) of
weekly study exhibited that Microcystis aeruginosa and Stephanodiscus
hantzschii blooms occurred in summer and winter respectively. Seasonally,
winters are dry (5% of total annual rainfall), and S. hantzschii proliferated
in every year. Summer blooms of M. aeruginosa showed inter-annual variability and dominated about 90% of algal species during dry years (19941996, , 800 mm of annual rainfall). A non-linear clustering by Self-Organizing Map on the algal proliferation in this river revealed that the increased heat energy, accumulated nutrients and decreased flow were responsible for both phytoplankton blooms. Furthermore, in the case of S.
hantzschii, larger magnitude of the cell density could be observed when
summer drought occurred (r250.61). It might be due to the regulated discharge from upstream dams. This study could suggest that drought can
give favorable conditions on the algal bloom formation, and the inter-annual climate changes can have time-delayed influence on the algal species
succession in river-reservoir systems.
JETZ, WALTER,1 CHRIS CARBONE2 and JAMES H. BROWN.1 1 Biology Department, University of New Mexico, Albuquerque, New Mexico,
USA; 2 Institute of Zoology, Zoological Society London, London, London,
UK. General rules for the scaling of home range size and density in
mammals.
The energetic equivalence rule states that densities N should scale to the
inverse of an animals metabolic needs Bi resulting a population energy flux
BT that is independent of body size M: BT 5 Bi x N 5 b0 M3/4 x a0M-3/4
(Damuth 1981). There has been considerable evidence that this relationship
holds true for a wide variety of mammals from different regions, and recently, Carbone & Gittleman (2002) showed that the fit of this scaling
relationship in carnivores can be enhanced by including information on
prey productivity. Secondary consumers appear to show a somewhat steeper scaling than primary consumers. A separate line of research has examined the scaling of home range size in mammals, and found a scaling of
M-1 (Kelt & Van Vuren 2001). Carnivores appear to scale more steeply
than herbivores, and recently fractal geometry of resource distribution has
been invoked to explain this difference (Haskell et al. 2002). Here we
present a model and empirical data that attempts to unify these disparate
scaling relationships. Analyzing several hundred studies of mammalian
home ranges and densities we find that density-derived per individual area
estimates consistently show a scaling with an exponent that is by 1/4 smaller than for home range size. We show that this difference is explained by
a positive 1/4 scaling of home range overlap. Higher degrees of overlap in
larger mammals come from both increased sociality and increasingly soft
home range boundaries, and can potentially be explained by the principles
underlying the scaling of day range. Inclusion of information on prey productivity significantly enhances the scaling fit for both home range and
density. Parsimony suggests that fractal resource distribution is not necessary to explain the steeper and trophically differing scaling of home range
size.
JHEE, EDWARD M.,1,* ROBERT S. BOYD1 and MICKY D. EUBANKS.2
Auburn University, Department of Biological Sciences, 101 Life Sciences,
Auburn University, AL, USA; 2 Auburn University, Department of Entomology and Plant Pathology, 301 Funchess Hall, Auburn University, AL,
USA. Hyperaccumulated Ni defends Streptanthus polygaloides (Brassicaceae) against Plutella xylostella (Leptidoptera).
1

Nickel hyperaccumulating plants contain more than 1000 mg Ni kg-1 dry

mass when grown on Ni-amended soils. We hypothesized that Ni hyperaccumulated by Streptanthus polygaloides serves as an elemental defense
against the Diamondback Moth, Plutella xylostella by deterring oviposition.
An oviposition experiment used 2 X 5 arrays of S. polygaloides grown on

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Abstracts

Ni-amended (high-Ni) soil interspersed with plants grown on unamended

soil (low-Ni). Diamondback Moths exhibited a significant oviposition preference for low-Ni plants. Moths allowed to oviposit on aluminum foil
sheets dipped in extracts of hi-Ni or low-Ni plants significantly preferred
low-Ni extracts. To determine if Ni was the factor influencing oviposition
preference, we allowed moths to oviposit on foil sheets coated with lowNi plant extract to which varying amounts of Ni were added. No difference
in oviposition was observed, suggesting that factors other than Ni influence
oviposition preference. Previous research has shown that hyperaccumulating S. polygaloides plants have a lesser concentration of glucosinolates.
This trade-off might influence oviposition by an oligophagous herbivore
since glucosinolates stimulate oviposition and feeding by Lepidopteran specialists on members of the Brassicaceae. To contrast fitness of high- and
low-Ni plants, eggs were allowed to hatch and larvae feed freely among
plants in the arrays. High-Ni plants had significantly greater biomass as
measured by change in number of leaves and plant height. Reproductive
effort (number of flowers) was also significantly greater for high-Ni plants.
Ni hyperaccumulation may be an effective plant elemental defense against
an oligophagous herbivore. By hyperaccumulating Ni, fitness of S. polygaloides is increased by toxicity to P. xylostella caterpillars and by decreasing oviposition by adults.
JIANG, LIN* and PETER J. MORIN. Rutgers University, ljiang@rci.
rutgers.edu, New Brunswick, New Jersey. Resource or predator control
of community structure: The importance of generalist and specialist
predators.
Ecologists have long recognized the importance of resources and predators
in structuring ecological communities. Despite several decades of research,
however, the question of what controls the relative strength of resource and
predator limitation of community structure remains elusive. Using aquatic
microcosms in which nutrient resources and predators were independently
manipulated, we show that predator characteristics can largely determine
the relative strength of resource and predator control of both species abundance and diversity of the middle trophic level in a food web with three
trophic levels. Predation dominated in communities with generalist predators, whereas resource impacts were more important in communities with
specialist predators. These results suggest that natural communities dominated by generalist predators and those dominated by specialist predators
may exhibit differential responses to nutrient enrichment imposed by human activities.
JIMENEZ-LOBATO, VANIA and TERESA VALVERDE. Lab. Especializado de Ecolog
a, Facultad de Ciencias, Universidad Nacional Autonoma
de Mexico (UNAM)., Mexico, D.F., Mexico. Population dynamics of the
shrub Acacia bilimekii in a semi-desert region in Central Mexico.
Shrubs are important elements of semi-desert vegetation. They provide microhabitats for the establishment of many species under their canopies thus
maintaining community structure and diversity. Yet, little is known regarding their population processes and the factors that affect them. Recent soil
erosion resulting from habitat degradation related to human activities may
potentially affect the regeneration of shrub populations, thus altering the
dynamics of complete semi-desert communities. We studied the population
dynamics of the shrub Acacia bilimekii in the Tehuacan Valley through the
analysis of population projection matrices (2001-2002). Additionally, we
carried out germination and seedling establishment experiments in two
sites: a well preserved and a relatively eroded one; in each site, experiments
were performed both under the shade of shrubs and in the open. The results
of these experiments were used to calculate the fecundity entries for transition matrices and project population dynamics in different theoretical scenarios (i.e. different disturbance degrees, and presence/absence of clonal
propagation). Germination was higher in the eroded than in the well preserved site, and germination percentages were similar under the shade and
in the open. However, seedling establishment followed the opposite trend,
with highest values in the well preserved site and under the shade of shrubs.
Population growth rate was significantly higher than unity in all cases.
Stasis was the demographic process that contributed mostly to l (c.a. 60%),
followed by growth and fecundity. According to the germination and establishment result, the population would grow slightly faster (l51.23) in
the well preserved site compared to the eroded site (l51.20). The scenario

that incorporated clonal propagation projected a population with a slightly

higher growth rate (1.29 and 1.26 respectively). These results suggest that
A. bilimekii populations have the potential to persist, withstanding relatively
high levels of disturbance conferring semi-desert communities a certain
degree of ecological resistance.
JIN, VIRGINIA L.,1,2,* CHRISTOPHER S. ROMANEK1,3 and REBECCA
R. SHARITZ.1,2 1 Savannah River Ecology Laboratory, Aiken, SC; 2 Department of Plant Biology, University of Georgia, Athens, GA; 3 Department of Geology, University of Georgia, Athens, GA. Evaluation of the
ammonia diffusion method for determining soil gross N mineralization
rates by 15N dilution.
The ammonia diffusion method is commonly used for the stable isotope
dilution technique to determine soil gross N mineralization rate. We evaluated the ammonia diffusion method developed by Brooks et al. 1989 using
laboratory incubations of stable isotope-labeled field soils extracted with
2M KCl. Temperate hardwood (H) soils and sandhills (S) soils were sampled in the field, brought back to the laboratory, and labeled with
(15NH4)2SO4 (d15N 537) to increase soil N by approximately 2 mg N kg1
dry soil. Control incubations were also conducted using 2M KCl spiked
with 300 m g NH4+-N. Initial samples were extracted and diffused immediately after addition of label, and labeled samples were incubated for 48
hours at room temperature then extracted and diffused. Triplicate samples
of extracts and diffused N from controls and each soil type for initial and
incubated soils were taken daily over a five-day period to characterize
changes in extract NH4+-N concentrations and isotope values of diffused
N. After the diffusion period, percent recovery of both total N and 15N
were highest in controls (88-103% and 97-115%, respectively), then H soils
(64-70% and 54-61%, respectively), then S soils (31-53% and 26-57%,
respectively). Isotopic values for diffused N in the controls were similar
for initial and incubated soils; however, d15N values of diffused N were
higher for initial soils than incubated soils in both H and S diffusions.
Diffused NH4+ d15N values from S soils were higher than H soils for both
initial and incubated soils on each day of the diffusion period. Overall
recovery of N label using the diffusion method appears positively correlated to the initial NH4+-N concentration of the diffusing solution. Our
findings suggest that more accurate calculations of gross N mineralization
rates can be made if the initial N of labeled soil extracts for diffusion
approximate 300 mg NH4+-N.
JOERN, ANTHONY. University of Nebraska-Lincoln, Lincoln, NE, USA.
Grasshopper abundance and species richness resulting from fire, topography and ungulate grazing in tallgrass prairie.
How does the abundance and species richness of common insect herbivores
respond to fire, topography and large ungulate grazing, major drivers in
determining the structure and function of tallgrass prairie? I evaluated
grasshopper species (Acrididae) from Konza Prairie to test the habitat heterogeneity hypothesis that predicts species diversity and overall abundance
will increase with increased structural heterogeneity and number of plant
species. Grasshopper species richness and abundance at Konza Prairie increased in response to grazing by bison, but fire interval and topographic
effects had little impact in this study. Habitat structure and heterogeneity
were positively related to both grasshopper species richness and overall
abundance. However, densities of common individual grasshopper species
differed significantly with regard to effects of fire, grazing and topography.
Abundances of some species are very responsive to bison grazing, while
other species are not. Densities in these latter species reflect responses to
fire interval, but in different ways. Clearly, it is important to evaluate responses at both species and community levels to understand the significance of major system drivers in tallgrass prairie to important insect consumers. Results will be discussed with respect to managing tallgrass prairie
for promoting insect biodiversity and conservation.
JOHANSEN, MATHEW P.,1,2,* DAVID D. BRESHEARS,3 JEFFREY J.
WHICKER3 and JOHN E. PINDER.2 1 Department of Energy, Los Alamos
Area Organization, Los Alamos, NM; 2 Colorado State University, Fort
Collins, CO; 3 Los Alamos National Laboratory, Los Alamos, NM. Comparative measurements of wind and water erosion in semiarid grassland, shrubland, and forest ecosystems.
Wind erosion is viewed as an important process in dryland ecosystems, yet
measurements are lacking, especially for dryland grasslands and forests, as

are consistently obtained estimates for cross-ecosystem comparisons. Further, erosion can result from both wind and water, yet measurement-based
estimates of both processes within the same ecosystem are lacking. Here
we compare wind and water erosion rates from three different semiarid
ecosystem types: a shrubland near Carlsbad, New Mexico, a grassland near
Denver, Colorado, and a forest near Los Alamos, New Mexico. In addition,
we propose a framework for comparing the horizontal transport due to
wind- and water-erosion. We used field measurements of wind erosion,
rainfall simulations, and extrapolation methods to compare the two types
of erosion within and among ecosystems. We found that wind erosion exceeded water erosion at the shrubland and forest sites, but not the grassland
site, where the high clay content of the soils contributed to greater amounts
of water erosion. Horizontal transport by wind was greater than that by
water for all three systems, overwhelmingly so in the shrubland. Horizontal
transport by wind exceed vertical transport by wind by more than an order
of magnitude in all three systems. Our results, which include some of the
only wind erosion measurements to date for semiarid grassland and forests,
provide a basis for hypothesizing trends in wind and water erosion among
ecosystems, highlight the importance of wind erosion in semiarid ecosystems, and have implications for land surface geomorphology, contaminant
transport, and ecosystem biogeochemistry.
JOHNSON, DALE W.,* JAY A. ARNONE, PAUL S. VERBURG, GITANE ROYCE, LINDA WALLACE, YIQI LUO and NANCY ZERBACH.
NSF-IRCEB interannual climate variability and ecosystem processes
in tallgrass praire: Non-destructive measures of soil nutrients.
A key element of the overall hypothesis in the ICREB study is to measure
soil N availability. This is a challenging proposition at best, as no truly
reliable and meaningful measure of soil N availability has yet been found;
in the case of the ICREB study, we have the additional challenge of very
limited plot size, making routine, destructive sampling of soils infeasible.
In order to address this problem, we have tested two non-destructive methods for assessing soil N availability: Plant Root Simulator Probes (PRS),
which consist of anion or cation exchange membranes imbedded in plastic
stakes (Western Ag Innovations, Inc., Saskatoon, Canada) and the WECSA
Soil Access System (Warrington Ecological Analysis, Ft. Collins, CO)
which employs a mixed-bed cation/anion exchange resin capsule fitted to
the end of a permanently-installed pvc tube. Soil N availability indices by
each of these methods are compared to each other, to standard destructive
soil analyses (C, N, mineral N, incubations), and to aboveground biomass
production. The advantages and disadvantages of each method are discussed.
JOHNSON, DANIEL M.,1,* MATTHEW J. GERMINO2,* and WILLIAM
K. SMITH.1,* 1 Wake Forest University, Winston-Salem, NC, USA; 2 Idaho
State University, Pocatello, ID, USA. Carbon gain in young seedlings of
Abies lasiocarpa and Picea engelmannii at alpine treeline.
Most research concerning the occurrence and stability of alpine timberlines
has focused on broad correlations with annual temperature regimes, without
addressing specific mechanisms. Timberline migration to higher altitude is
dependent upon new seedling establishment away from the forest edge into
the timberline ecotone. Reductions in carbon gain in these establishing
seedlings have been shown to decrease survival. The purpose of this study
was to compare photosynthetic carbon gain and mortality in seedlings of
two codominant tree species of the subalpine (Abies lasiocarpa and Picea
engelmannii), across a representative altitudinal gradient from the subalpine
forest to the upper treeline (2965m-3198m). Differences in environmental
factors (temperature, light, and water) that might be responsible for differences in carbon gain were also examined. Mean daily photosynthesis in
seedlings of both Abies lasiocarpa and Picea engelmannii was reduced (19
and 29 % respectively) at the higher-elevation site. Minimum nighttime
temperatures below 28C were more frequent at higher elevation and were
correlated with reduced maximum photosynthesis the following day. Reduced water status was correlated with reduced carbon gain equally at both
sites early in the season before snowmelt, as well as late in the season
when soils began to dry. Reductions in photosynthesis at the high-elevation
site do not appear to be related to water status because similar values of
xylem pressure potentials were observed between sites. Sunlight regimes
were highly variable at both sites due to structural shading, although this

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variation was substantially reduced on cloudy days (44% of all days measured). Moreover, cloudy days resulted in greater integrated daily carbon
gain, especially at lower elevation, based on a photosynthesis model that
compared potential (simulated) versus actual (measured) daily carbon gain.
Photosynthetic responses to temperature, sunlight and water indicated that
sunlight and nighttime temperature were the major limiting factors.
JOHNSON, DEREK M.1,2,* 1 University of Miami, Coral Gables, FL, USA;
2
Penn State University, State College, PA, USA. Temporal variability in
source-sink dynamics.
Source-sink models are used to address spatial questions about organisms
living in heterogeneous environments. In traditional source-sink models,
vital rates and movement probabilities are assumed to be temporally homogeneous. Numerous studies, however, have demonstrated that temporal
heterogeneity in many forms is the rule rather than the exception in natural
systems. In this study, I present a temporally-variable source-sink dynamic
of a neotropical beetle, Cephaloleia fenestrata, driven by flooding disturbance. A flood completely inundated a subset of C. fenestrata host plants,
Pleiostachya pruinosa during the study. Survival and movement probabilities of C. fenestrata were estimated from data on approximately 1,200
individually-marked beetles. I demonstrate that a stochastic flood event
caused a 6-fold increase in mortality in the flood zone compared to the
upland habitat. In non-flood periods survival probability did not differ between habitats. Thus, the flood zone acts as a sink only during flood periods. Movement among the habitats was low (1-5%) during non-flood
periods, but during a flood event increased over 10-fold (69%). Despite
this, the net movement of beetles was from the upland to the flood zone.
Given the estimated flood frequency of once per year, the overall population
growth rate in the flood zone was determined to be , 1. Given estimates
of flood probability, these results are consistent with the flood zone acting
as a sink habitat for C. fenestrata. Ultimately, whether or not the flood
zone is a sink or pseudo-sink habitat, and how strong a sink it is, depends
on the flooding frequency and density-dependent effects.
JOHNSON, JERALD B.* Getting from pattern to process in evolutionary ecology: testing alternative models of life-history evolution.
Studies of natural selection in the wild almost always begin by examining
patterns of association between phenotypic adaptations and environmental
factors thought to shape evolutionary change. Unfortunately, many studies
pay little attention to the effects of model selection on the evolutionary
inferences drawn from such correlative data. In this study, I employed a
candidate model analysis to examine four potential causes of life-history
evolution in the livebearing fish Brachyrhaphis rhabdophora. Combining
factor analysis with path analysis, I constructed a nested set of 17 models
that represent the hypothetical effects of four putative selective agents
(mortality, density, resource availability, and habitat stability) on life-history evolution in this species. These models represent both direct and indirect of effects of selection on the life history. Using the Akaike Information Criterion to distinguish among models, I found that simple models
that contained only single selective agents most parsimoniously explained
life history divergence among 27 B. rhabdophora populations. Surprisingly, the four putative selective agents could not be distinguished, suggesting
that the selective environment could be composed of a single selective
agent confounded with other environmental factors, or could be composed
of a suite of environmental factors that act in concert to shape the life
history. In addition, comparisons among more complex models indicated
that direct effects of selective agents appear to have primacy over combinations of indirect selective interactions in explaining intraspecific variation
in B. rhabdophora life histories.
JOHNSON, KELLY S.* Ohio University, Athens, OH. Plant phenolics
as beneficial antioxidants for insect herbivores.
High foliar phenolics have frequently been associated with decreased performance of insect herbivores, but their physiological mode of action is
not well understood, and in many instances this relationship is not apparent.
One hypothesized mode of action is that phenolics damage tissues and
reduce food quality via pro-oxidant effects (by generating reactive oxygen
species when oxidized), particularly in herbivores with alkaline, oxidizing

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gut environments. We investigated the pro- versus anti-oxidant behavior of

five common plant phenolics (quercetin, caffeic acid, catechin, chlorogenic
acid, and tannic acid) to test the alternative hypothesis that dietary phenolics may offer antioxidant benefits to some herbivores. Using an in vitro
radical cation decolorization assay (ABTS), we found that both gut fluid
and hemolymph of Manduca sexta exhibit strong radical quenching properties. All of the phenolics except caffeic acid behaved as antioxidants
when added to these body fluids at 2-10 uM concentrations. Some phenolics (chlorogenic acid) enhanced antioxidant capacity of body fluids in
an additive manner. These results suggest that many plant phenolics may
provide beneficial antioxidant benefits to insect herbivores, and thus play
more diverse ecological roles in plant-herbivore interactions than chemical
defense.
JOHNSON, MARC T.J.* and ANURAG A. AGRAWAL. University of
Toronto, johnson@botany.utoronto.ca, Toronto, Ontario, Canada. Community genetics: The importance of genetic and environmental variation in affecting arthropod community composition on Oenothera biennis.
Genetic and environmental influences on plant resistance are well known
to affect insect populations, but their effects on arthropod community composition are not well understood. We tested how plant genotype interacts
with the environment at two spatial scales to affect herbivory, arthropod
diversity (Simpsons D), species richness, and species abundance. We grew
14 clonal families of Oenothera biennis in each of five common gardens
in sourthern Ontario. Each garden represented a unique habitat of O. biennis, and spatial blocks within gardens were created to examine environmental variation within habitats. All arthropod species on plants were identified and counted every 2-4 weeks during the summer. The amount of
plant damage by early season herbivores varied nearly 2-fold between clonal families, but the effects of plant family were dependent on environmental
variation within habitats. Plant family also affected arthropod diversity,
species richness and abundance on individual plants, but in all cases, the
effects of family were dependent on the habitat in which plants grew. Species richness and abundance varied by as much as 140% and 280%, respectively, between clonal families within a garden, but genetic variation
affected herbivores more strongly than it did predators. Mean species richness was positively correlated (r 5 0.68) with the proportion of plants
within a clonal family that flowered. Plant genetic variation was frequently
more important than within-habitat environmental variation in affecting arthropod community composition. At larger scales, however, variation between habitats was the most important factor. These results demonstrate
that intraspecific genetic variation in plants has large ecological consequences for arthropod communities, but its influence decreases with increasing geographic scale.
JOHNSON, MICHELE A.* and JONATHAN M. CHASE. Washington
University, St. Louis, MO. Community age and species composition:
Invasibility and turnover in natural zooplankton assemblages.
The patterns of species composition among communities can give important insights into the process of community assembly. Previous hypotheses
regarding community assembly include the achievement of (1) a single
equilibrial state (composition determined by a combination of environment
and species interactions), (2) multiple equilibrial states (composition determined by environment, species interactions, and invasion history), (3)
cyclical equilibrial states (same as 1, except composition changes cyclically
through time), and (4) the absence of equilibria where composition is continually in flux (owing to neutral processes such as ecological drift). To
distinguish among these four hypotheses, we studied the assembly of zooplankton communities among constructed ponds that varied with age and
thus the time for communities to establish. We established natural zooplankton communities from the ponds of different ages in experimental
mesocosms and then introduced potentially invading zooplankton species
from within the regional species pool. We found that older pond communities were less likely than younger communities to experience species
turnover when invaded by species from the region. This pattern, in which
older communities appear more stable in species composition when perturbed with invasions from the regional species pool, conforms to concepts
that assume that species composition achieves some sort of equilibria

through time. In addition, we found that when invaders were rare with
respect to the natural community, the community experienced less species
turnover then when invaders were common with respect to the natural
community. This suggests that species interactions are an important factor
in establishing the species composition of communities, and that they appear to have multiple stable equilibria.
JOHNSON, PIETER T., * KEVIN B. LUNDE, DEREK A. ZELMER and
KIRWIN WERNER.4 1 Center for Limnology, University of Wisconsin,
680 North Park St., Madison, WI, USA; 2 The Roberts Environmental Center, Claremont McKenna College, 925 North Mills Avenue, Claremont, CA,
USA; 3 Department of Biological Sciences, Emporia State University, 1200
Commercial St., Emporia, KS, USA; 4 Department of Environmental Sciences, Salish Kootenai College, Pablo, MT, USA. Amphibian malformations and parasite infection: An emerging disease?
1,

Widespread reports of malformed amphibians have generated concern over

the possibility of an emerging threat to amphibians and other wildlife.
Recent field and laboratory evidence has implicated infection by a trematode parasite (Ribeiroia ondatrae) as an important cause of such deformities. Prior to 1999, however, there were no records of Ribeiroia from wildcaught amphibians. Whether these parasite-induced malformations represent a new or expanding phenomenon in wetland habitats remains conjectural. I investigated the causal role of Ribeiroia infection in nine North
American wetlands that historically (1946-1988) yielded high frequencies
of severe malformations in amphibians. Amphibian voucher specimens corresponding to these sites were redescribed using current malformation terminology and dissected to isolate trematode metacercariae. Between 1999
and 2002, historical malformation sites from California, Colorado, Idaho,
Mississippi, Montana, Ohio, and Texas were resurveyed for deformed amphibians and the presence of Ribeiroia. Malformation patterns were compared with data presented in the original historical accounts and with recent
reports from the western USA. Direct identification and classification by
discriminant function analysis indicated that historical malformations at six
of eight sites were associated with infection by Ribeiroia, dating back as
far as 1946. Malformations recorded historically at these sites were consistent with the documented effects of Ribeiroia infection, including extra
limbs, cutaneous fusion, and bony triangles. Of the six sites that still supported amphibians upon resurvey, three continued to support severe limb
malformations at frequencies of 7 to 50% in one or more species. Although
no pesticides were detected, amphibians from each of these sites were
infected with Ribeiroia metacercariae. The disappearance of amphibian
populations at four of the five sites associated with Ribeiroia raise concerns
about possible long-term impacts of infection and malformations. Taken
together, these results suggest that Ribeiroia infection has historically been
an important cause of mass malformations in amphibians.
JOHNSON, SCOTT N.* and PETER J. GREGORY. University of Reading,
Reading, United Kingdom, United Kingdom. Root weevil attraction to
clover roots 2 the roles of general and specific signals.
The clover root weevil (Sitona lepidus) exclusively attacks white clover
(Trifolium repens) throughout Europe and North America, where it is valued both as a forage legume and for the nitrogen enrichment it provides
to grassland. The adult weevil feeds on leaves above-ground where it lays
eggs that fall to the soil surface. Soil-dwelling larvae emerge from eggs
and feed on roots, initially feeding on root nodules containing the N2 fixing
rhizobia, before feeding on progressively larger roots. Using X-ray microtomography we have shown that newly-hatched larvae move to the root
nodules directly. We tested the hypothesis that chemical signals from the
roots or root rhizobial nodules could be exploited by S. lepidus to locate
white clover roots. In arena experiments, .85% of responsive S. lepidus
selected host plant roots (T. repens) over non-host plant roots (e.g. ryegrass,
red clover, vetch). When given a choice between T. repens roots with and
without root nodules, 60% of responding insects selected roots with nodules and 40% chose those without. In choice tests between T. repens roots
and excised root nodules, .80% of responsive S. lepidus selected nonnodulated roots rather than the excised root nodules. Respiratory emissions
of CO2 from roots are known to attract some root feeding insects. S. lepidus
showed a dose-dependent attraction to CO2 in arena experiments. CO2 concentrations in the rhizosphere of T. repens with root nodules were .20%

higher than the rhizosphere of T. repens without root nodules, and .30%
higher than the rhizosphere of non-host plants. It is proposed that specific
chemical signal(s) are implicated in the attraction of S. lepidus to T. repens
roots, but general signals, such as CO2, might also play a role in root
location.
JOHNSTON, CAROL A.,1 BARBARA BEDFORD,2 MICHAEL BOURDAGHS,3 TERRY BROWN,3 CHRISTIN FRIESWYK,4 LYNN VACCARO2 and JOY ZEDLER.4 1 South Dakota State University, Brookings, SD;
2
Cornell University, Ithaca, NY; 3 Natural Resources Research Institute,
Duluth, MN; 4 University of Wisconsin, Madison, WI. Plants as environmental indicators in Great Lakes coastal wetlands.
Coastal wetlands are the focal point of much human activity, both direct
and indirect, that threatens their condition and existence. Wetland plants
and their assemblages can provide early warning of environmental stress
in coastal ecosystems. Plants have the advantage over many faunal indicators that they remain in place, simplifying sampling and increasing the
likelihood that the biotic indicator is spatially coincident with in situ stressors. Wetlands and their vegetation have also been extensively mapped,
providing a basis for analyzing landscape-scale wetland attributes and their
relationship to stressors. Field studies were conducted in 70 coastal wetlands spanning the Great Lakes coast from Duluth, Minnesota to Cape
Vincent, New York. Wetlands were sampled from three types based on
their dominant water source and connectivity to the lake: lacustrine, riverine, and barrier-protected. Emerging indicators focus on selected species
(invasives), assemblages (e.g., biodiversity measures), and landscape characteristics (e.g., interspersion of wetland types).
JOHNSTON, JILL A.,1,2,* MICHAEL REAGON1 and ALLISON A.
SNOW.1 1 Department of Evolution, Ecology and Organismal Biology,
Ohio State Universtiy, Columbus, OH, USA; 2 Department of Plant Biology, University of Minnesota, St. Paul, MN, USA. Potential for gene flow
from cultivated sorghum to shattercane (both Sorghum bicolor ssp. bicolor) in the midwestern U.S.
Close taxonomic relationships between some crops and agricultural weeds
create the potential for beneficial crop genes to introgress into weed populations. Cultivated sorghum has two close relatives that are pests throughout many areas of the U.S. Johnsongrass (S. halepense) is the best-known
Sorghum weed, but shattercane shares subspecific status with the crop and
therefore has greater potential for rapid crop to weed gene flow. To evaluate
the potential for crop-to-weed gene flow, we surveyed six microsatellite
loci in eight populations of shattercane and three types of cultivated sorghum from Kansas and Ohio. Four shattercane populations were collected
from sorghum fields (contact) and four were collected from other crop
fields (non-contact). Based on the shared proportion of 60 alleles, crops
are most genetically similar to each other together and to one shattercane
population growing in contact with crops. Only one of the shattercane
populations that is relatively isolated from cultivated sorghum clusters
away from the crops and other shattercane populations. The remaining
shattercane populations are very similar genetically. Geographic distance
and contact with crop plants does little to explain patterns of genetic similarity among populations. Migration estimates based on Fst statistics are
fairly low (Nm 5 0.5), and overall there seems to be a large amount of
genetic diversity partitioned among populations (Fst 0.3-0.4). While this
could suggest a fair degree of isolation among populations, we believe it
is more likely the result of recent founding events in these weed populations. Our data show that the most closely related populations are separated
by hundreds of miles. This seems most likely due to similar founders or
recurring input of similar alleles from nearby crops.
JOHNSTON, TAMMY* and KEITH CLAY. Indiana University, Bloomington, Indiana. Host plant influences parasite transmission mode in
a grass-fungal endophyte association.
Theories of the evolution of virulence generally rely on a positive correlation between parasite transmission and virulence. It is particularly important to test the correlation in a system where a range of both vertical
and horizontal transmission modes co-occur to critically evaluate how
transmission mode affects the evolution of virulence and to explore the

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171

potential for interspecific associations to evolve towards reduced virulence

and possibly mutualism. Importantly, the degree to which the host versus
the parasite controls transmission mode is not well understood. Such an
understanding would enlighten models of the evolution of virulence, which
often assume that the transmission mode is directly linked to traits of the
parasite. The plant/fungus interaction, Elymus hystrix infected with the endophyte Epichloe elymi, exhibits both vertical and horizontal transmission
and thus provides an ideal system to explore the relationship between virulence and transmission mode. In a common garden, we detected significant variation in horizontal transmission potential across eight maternal
half-sib lines of infected plants and this variation was consistent across
years. Vertically infected offspring resembled maternal plants in horizontal
transmission. No significant variation was observed in vertical transmission
to seeds or in the number of inflorescences produced. While all seeds were
infected initially, longevity of the infection in dormant seeds varied across
the maternal lines. Thus significant genetic variation in transmission mode
exists upon which selection can act. A second experiment using artificial
host/fungus combinations revealed significant host, fungus and interaction
effects on the number of plant tillers, which plays a significant role in
horizontal transmission. These results demonstrate that both the host and
fungus shape the expression of parasite transmission. Thus, evidence from
a host-parasite interaction that spans a continuum from vertical to horizontal transmission highlights the importance of both plant and parasite in the
evolution of transmission and virulence.
JONES, ANDY.* Department of Plant Biology, University of Georgia, Athens, GA, 30602. Genetic reconstruction of seed dispersal patterns in a
neotropical moist forest.
Seed dispersal is a central process in a number of the major hypotheses
proposed to explain the maintenance of high tree species diversity in tropical moist forests. These include the escape and enemies hypothesis (Janzen
1970, Connell 1971), the dispersal limitation hypothesis (Tilman 1994,
Hurtt and Pacala 1995), and a neutral theory of biological diversity (Hubbell 2001). However despite the ecological and evolutionary importance of
seed dispersal, there is a considerable void of empirical data on the frequency, patterns, and mechanisms of dispersal. This is particularly evident
in long distance dispersal events within and among natural plant populations. I employ a simple genetic method to identify the exact maternal
parent of seeds of Jacaranda copaia dispersed to seed traps within the 50ha Forest Dynamics Plot (FDP) on Barro Colorado Island (BCI), Panama
to determine the patterns of seed dispersal for this wind dispersed tree
species. Using microsatellite genetic markers developed for Jacaranda, I
genotyped remnant maternal tissue from dispersed seeds and matched this
genotype to the source adult maternal genotype in the population. This
method allows for determination of exact seed dispersal distances. I address
a number of questions including: (1) How frequent is long distance seed
dispersal (here defined as dispersal at distances greater than 100m) in Jacaranda? (2) How well do current inversely modeled dispersal curves estimate and predict the dispersal patterns measured above? (3) What biotic
and abiotic factors determine dispersal distances? And (4) Are Jacaranda
seeds disproportionately dispersed to tree fall gaps? I will also present
preliminary results for other wind-dispersed tree species in Panama.
JONES, CRYSTAL A.* and GARY S. KLEPPEL. University at Albany,
State University of New York, Albany, NY. Biodiversity education in
New Yorks secondary schools: an assessment of current curricula.
That biodiversity is critically linked to ecosystem function and stability,
often in complex ways, is increasingly appreciated at the level of professional ecology. The public in general appears far from any such understanding or appreciation. The secondary education system is viewed by
some as an appropriate vehicle for transmitting that crucial appreciation to
the public sector. To accomplish this, it is critical to ascertain the current
state of biodiversity education in the nations schools and to identify the
constraints, as well as the opportunities, for teaching biodiversity in the
classroom. We are partnering with the Biodiversity Research Institute at
the New York State Department of Education in an effort to develop a
report entitled "The State of Biodiversity Education in New York." A questionnaire focusing on teaching methods, facilities and infrastructure, teacher
preparation, and evaluation of existing curricula has been distributed to 300

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Abstracts

randomly selected high schools in the state. Of particular importance is the

identification of difficulties and shortcomings in teacher preparation and
curriculum delivery. Responses are being compiled, formatted and analyzed
to identify statistically significant concensi. Suggestions for curricular modification are being developed to overcome current limitations.
JONES, DAVID T.,* MICHAEL S. ROSS, BERNICE HWANG, JOSHUA
WALTERS, STEVEN OBERBAUER and KRISH JAYACHANDRAN.
Florida International University, jonesd@fiu.edu, Miami, Florida. Effects
of Simulated Hydrologic Regimes on Twelve Tree Species of Everglades Tree Islands.
The responses of tree species to environmental gradients within Everglades
tree islands have rarely been tested under controlled conditions. We describe the response of 12 tree species to three realistic hydrologic regimes
that were applied while holding other variables (e.g., light, nutrients) constant. Seven species (Annona glabra, Chrysobalanus icaco, Ilex cassine,
Magnolia virginiana, Myrica cerifera, Persea borbonia, Salix caroliniana)
were characteristic of Shark Slough (Everglades National Park, Florida)
bayhead and bayhead swamp forests, while five (Bursera simaruba, Coccoloba diversifolia, Eugenia axillaris, Mastichodendron foetidissimum,
Simarouba glauca) were common in hardwood hammocks (uplands) in the
region. Recently emerged seedlings of all species were collected from Everglades tree islands in May-June 2001 and raised in a shadehouse. The
experimental design was randomized complete block, with 36 species x
treatment combinations represented twice in each of four pools (blocks).
Water levels in the pools were managed to mimic variation among weekly
means of water depths at a known marsh location. Treatments representing
hardwood hammock, bayhead, and bayhead swamp hydrologic regimes
were maintained in each pool at the appropriate relative heights. Commencing in April 2002, plant structural and physiological responses were
monitored at one to six week intervals for 24 weeks. Upland species
showed signs of both morphological and physiological stress from inundation between weeks 6 and 12. Some species typically found in bayhead
and bayhead swamp sites appeared to be better adapted to rising water
levels than others. Our data suggested that Annona, Magnolia, Myrica, and
Salix responded most positively to flooding, while Chrysobalanus, Ilex, and
Persea were less flood-tolerant. The arrangement of species according to
their response to the flooding treatment roughly paralleled their spatial distribution in the marsh landscape. Such experimental work may allow us to
isolate the mechanism and timing of morphological and physiological responses in tree species, possibly permitting early warning of flooding stress
in tree islands and modification of water distribution that may benefit natural resources management in the Everglades.
JONES, ROBERT H.,* JULIA M. SHOWALTER, GLEN N. STEVENS
and MATTHEW A. NEATOUR. Department of Biology, Virginia Tech,
Blacksburg, VA. Natural re-invasion and artificial restoration of upland hardwoods in the coastal plain of South Carolina.
In the upper coastal plain of the southeastern USA, past agricultural activities have reduced the abundance and distribution of some valuable upland
hardwood species. To identify ecological factors limiting natural re-invasion, and to develop methods for restoration, we planted seeds of five
hardwood species in spring 1997, and bare-root seedlings of four species
in spring 1998. Soils were mesic to xeric, and stand structures included
uncut versus recently clearcut 40+ year old pine plantations. Some understory seedlings were located in trenched plots, and others were released by
removing the pine overstory three years after planting. Five-year survival
and growth were influenced by our treatments (i.e., presence of an overstory and timing of overstory release), but also by factors that we did not
plan or control, including competition with herbs, soil moisture during the
dormant season, and seed predation. At age five, very few planted seeds
had emerged and survived (0-42% depending on species and site), and all
survivors were short (, 20 cm). High seed predation of Carya spp. occurred in one of four test sites. Compared with planted seeds, planted seedlings had superior performance. Quercus alba seedlings performed best
when planted in clearcuts (mean five-year survival .60% and height .110
cm). Cornus florida seedlings performed best when planted in the understory and then released after three years (mean five-year survival . 70%
and height .210 cm). Carya spp. seedlings did very poorly regardless of

treatment (five-year survival ,20% and total height , 20 cm). In trenched

understory plots, five-year height and base diameter of Q. alba and C.
florida seedlings were nearly four times greater than in untrenched plots.
Our experiments suggest that dispersal is a limiting factor to natural reinvasion. Furthermore, seedling planting with attention paid to minimizing
competition may be a practical option for restoration.
JONSEN, IAN D.* Dalhousie University, Halifax, NS, Canada. Metaanalysis of aquatic and terrestrial animal movement using state-space
models.
The study of animal movement and behavior is being revolutionized by
technology, such as satellite tags and harmonic radar, that allows us to track
the movements of individual animals. However, our ability to analyze and
model such data has lagged behind the sophisticated collection methods.
We review problems with current methods and suggest a more powerful
and flexible approach, state-space modelling, and we illustrate how these
models can be posed in a meta-analytic framework so that information from
individual trajectories may be combined optimally. State-space models enable us to deal with the complexity of modelling animals interacting with
their environment but, unlike other methods, they allow simultaneous estimation of measurement error and process noise that are inherent in animal
trajectory data. A Bayesian framework allows us to incorporate important
prior information when available and also allows meta-analytic techniques
to be incorporated in a straightforward fashion. Meta-analysis enables both
individual and broader-level inference from observations of multiple individual pathways. Our approach is powerful because it allows researchers
to test hypotheses regarding animal movement, to connect theoretical models to data, and to use modern likelihood-based estimation techniques, all
under a single statistical framework. We illustrate the approach with examples using ARGOS satellite data collected from marine vertebrates such
as Leatherback turtles (Dermochelys coriacea) and radio transmitter data
collected from terrestrial vertebrates such as fishers (em Martes pennanti).
JUDD, KRISTIN E.,1,* BYRON C. CRUMP,2 DON R. ZAK,1 ROBERT
L. SINSABAUGH3 and GEORGE W. KLING.1 1 University of Michigan,
Ann Arbor, MI; 2 Ecosystem Center, Marine Biological Lab, Woods Hole,
MA; 3 University of New Mexico, Albuquerque, NM. Dissolved organic
carbon in soils, streams, and lakes of an arctic catchment: linking bioavailability, chemistry, and microbial community comoposition.
The transport rate and processing of dissolved organic carbon (DOC)
through catchments depends, in part, on its bioavailability to the microbes
it encounters as it travels down-slope. DOC chemical characteristics and
microbial community physiological capabilities determine DOC bioavailability; however, little is known about the relative importance of these
factors at various locations in a catchment or over the course of the growing
season. To better understand the major controls on spatial and temporal
patterns of DOC bioavailability and microbial activity, we examined microbial community composition and DOC chemistry along a terrestrialaquatic gradient in an arctic catchment and conducted experiments to determine the relative importance of these two factors in controlling carbon
processing. Both DOC chemistry and microbial community structure (phospholipid fatty acids) differed between tussock, birch-willow, and wet sedge
soils. In mesocosm experiments, we simulated down-slope transport of
DOC by adding DOC from up-slope soil and surface water sources to
stream and lake microbial communities. Microbial production was monitored over the course of four weeks, and DOC chemistry (phenolics, simple
carbohydrates, and amino acids) and microbial community composition
(denaturing gradient gel electrophoresis (DGGE)) was measured at the beginning and end of the experiment. Addition of up-slope DOC altered microbial production such that it was similar to microbial production of DOC
source communities. Addition of both wet sedge and tussock soil-water
DOC to stream and lake microbial communities greatly enhanced microbial
production relative to controls. Addition of stream water to a lake inoculum, however, depressed microbial production in lake water communities.
Over the course of the experiment, microbial community structure in treatment mesocosms shifted in the direction of the DOC source community,
and the addition of soil-water DOC had the strongest effect. These results
suggest that DOC chemistry strongly drives the structure and function of
microbial communities in arctic tundra systems.

JUICE, STEPHANIE M.1 and MELANY C. FISK.2,* 1 Cornell University,

Ithaca, NY, USA; 2 Appalachian State University, Boone, NC, USA. Decomposer fungal community response to calcium additions in a northern hardwood forest, NH.
Soil pH is likely to be a master variable influencing the structure and
function of soil decomposer communities. We tested the influence of calcium additions (as the mineral wollastonite) and an associated rise in pH
on the species composition of basidiomycete fungi in the northern hardwood forest at the Hubbard Brook Experimental Forest, NH. Fungal communities were compared using the terminal restriction fragment length
polymorphism approach (t-RFLP), for Oe, Oa, and mineral soil horizons
of paired control and calcium amended plots. Basidiomycete species composition differed significantly among soil horizons and species composition
appears to have shifted in the Oe horizon in response to Ca additions. Our
results suggest that in surface organic horizons, decomposer fungi are influenced by base status of the soil, and this is likely to impact litter decomposition processes.
JULIANO, STEVEN A.,1,* MARIETA A.H. BRAKS,2 NILDIMAR A.
HONORIO,3 L. PHILIP LOUNIBOS2 and RICARDO LAURENCO-DEOLIVEIRA.3 1 Illinois State University, Normal, IL, USA; 2 Florida Medical Entomology Laboratory, University of Florida, Vero Beach, FL, USA;
3
Instituto de Oswaldo Cruz, Rio de Janeiro, Rio de Janeiro, Brazil. Competitive advantage and plasticity in growth and development in competing container mosquitoes.
A clear competitive advantage for one species over another species leads
directly to questions about the mechanisms producing that advantage. For
resource competition, greater feeding rate and greater efficiency are commonly postulated as mechanisms, but another way of realizing competitive
advantage may be greater phenotypic plasticity, expressed as greater flexibility of size and development rate of one competitor, which may buffer
a species against mortality effects of competition. Because of flexibility in
one life history trait (e.g., size at maturity) impacts of competition on other
life history traits more closely related to fitness (e.g., survivorship) may be
reduced, giving a species an advantage in competition with other species
that suffer mortality in competition. This hypothesis predicts that competitively superior species should: 1) suffer lower mortality when in competition; and 2) show a greater range of size or development time in response
to a range of conditions than do competitively inferior species. We tested
these predictions for competing larvae of two mosquitoes, the recently
introduced invasive Aedes albopictus, and the long-established Aedes aegypti. In field experiments on two continents, A. albopictus has a clear
advantage over A. aegypti. In the lab, although this advantage was dependent on conditions, when there was a strong advantage it usually fell to A.
albopictus. This advantage is associated with greater survival to adulthood
under the most severe conditions of inter- and intraspecific density. As
predicted, in several independent studies, A. albopictus has significantly
greater plasticity of body size and development time than does A. aegypti.
This is true for both males and females, but is most evidence for female
body mass. Thus, in this system, it appears that greater plasticity of size
and development time are possible as mechanisms for the competitive advantage of A. albopictus over A. aegypti.
JUNG, DAWOON* and STANLEY I. DODSON. University of WisconsinMadison, Madison, Wisconsin. Identification of contaminantresponsive
protein in Daphnia magna.
Cytochrome P2450 (CYP) enzymes are involved in the metabolism of a
wide variety of external toxicants. They can be useful as tools for the study
of toxic substance metabolism and as indicators of contamination. We have
identified CYP enzymes in Daphnia magna, an important constituent of
the freshwater food web. Experiments using immunoblotting technique indicated the existence of the protein that interacts with antibodies for mammalian CYP 3 family of cytochrome P2450 enzymes. This protein was
roughly 80 kD in size, similar to the P450 enzymes of other species, such
as mosquitoes. The protein showed differential expression in response to
the different amounts of toxaphene when chronically exposed to the chemical. In acute exposure to the same dose of chemicals, the difference in
protein expression was not as pronounced. We were also able to see dif-

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ferences in protein level among different populations within Daphnia pulicaria. Our results indicate that Daphnia acclimate to sublethal doses of
chronical exposure to toxic chemicals. Furthermore, the results imply that
if exposed over a long term, different ability by different populations to
adjust to the contamination may have profound effect on the population
composition in different habitats. In addition, the identified CYP protein
may be a useful tool as an early warning system in evaluating contamination in freshwater system.
JURENA, P. N.* and O. W. VAN AUKEN. University of Texas at San
Antonio, Department of Earth and Environmental Sciences, 6900 N. Loop
1604 W., San Antonio, Texas, USA. Gas exchange rates and water relation responses of Juniperus ashei seedlings to drought.
The central Edwards Plateau, Texas, is dominated by Juniperus ashei
woodlands which comprise . 40% of the canopy cover at ca. 1500 trees/
ha. Survival is reported to be highest beneath the woodland canopy and
decreased at the canopy edge but growth rates are highest at the canopy
edge. Little is known of the drought response of J. ashei seedlings and its
affects on spatial differences in growth and survival. Yearly mean photon
flux density (PFD) measured beneath the canopy was 346.11696.81 mmol
m22 s21. Soil moisture beneath the canopy was 34.8263.72%. The light
compensation point for J. ashei seedlings was between 25 and 50 mmol
m22 s21 PPFD and Amax (90%) was 4.4860.03 mmol m22 s21 at 750 mmol
m22 s21 PPFD. Soil moisture dropped from 43.3[plusmn:]1.5% to
7.661.0% through the greenhouse experiment. Seedling water potential
started at 20.860.1 MPa and did not decline until after the soil moisture
dropped below 20.563.6%(after three weeks). Gas exchange rates did not
decline until three weeks after drought conditions were applied. Juniperus
ashei seedling water potential and gas exchange rates appear to be decoupled from soil moisture levels.
JURGENSEN-ARMSTRONG, MELISSA J.,1,* ALAN R. BERKOWITZ,2
JASON TAYLOR3 and ELVAN GUR-EDEMAN.4 1 Diversity Initiatives in
Science, Flagstaff, AZ; 2 Institute of Ecosystem Studies, Millbrook, NY;
3
Ecological Society of America, Washington, DC; 4 United Negro College
Fund, Fairfax, VA. Yes, no, maybe, I don9t know: The decisions SEEDS
students make about an ecology pathway.
Upon the completion of the first phase of SEEDS (Strategies for Ecology
Education, Development and Sustainability), an initiative designed to promote ecology to undergraduate minority students, we conducted interviews
of 39 students and alumni who had participated in the program. We were
interested in discovering the initial influences that piqued student interest
in ecology and the natural environment, the role of student undergraduate
experience in helping them develop their ecology interest, and how the
combination of initial interest and undergraduate influences interacted to
affect student decisions about pursuing an ecology pathway. Initial findings
indicate that nearly all interview participants had an early exposure to the
natural environment through family outdoor activities, youth groups, their
pre-college education, or a family member in the environmental field. Despite an early exposure to the natural environment, participants usually did
not decide to explore an ecology pathway until their undergraduate career
when many of them experienced multiple, positive influences that encouraged their ecology interest. Participants most often cited SEEDS, undergraduate professors, and ecology courses as influences that prompted them
to take interest in ecology. A few participants, however, appeared to have
a strong motivation toward a particular path regardless of what influences
they experienced. Insights into the larger question of what students decided
about an ecology pathway given this myriad of influences were gained from
comparing students that are pursuing ecology (persisters), chose another
path (leavers), are still unsure of their path (undecided), or have incorporated ecology into another career (modifiers).
KALKHAN, MOHAMMED A.,1,* THOMAS J. STOHLGREN,2 DEBBIE
A. GUENTHER1 and PAUL H. EVANGELISTA.1 1 Natural Resource Ecology Laboratory, Colorado State Unversity, Fort Collins, CO, USA; 2 The
National Institute of Invasive Species Science, USGS, Fort Collins Center,
Fort Collins, CO, USA. Understanding the landscape ecology of cryptobiotic crust cover of Grand Staircase-Escalante National Monument,
Utah: A map and spatial models.
Grand Staircase-Escalante National Monument (GSENM), Utah, USA, represents a complex landscape of plant diversity and covers an area of about

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2 million acres. Key biological parameters can be estimated using multiscale sampling with multi-phase design to provide unbiased estimates of
vegetation and soil characteristics. We evaluated the vulnerability of various habitats to invasion by exotic plants over the entire Monument. This
paper will provide examples only on cryptobiotic crust cover. A total of
367 Modified-Whittaker nested plots (0.1 ha) were established, and 19 vegetation cover types were found. For modeling large-scale and small-scale
variability to predict distribution, presence, and pattern of cryptobiotic and
soil characteristics, we integrated remotely sensed data, GIS, field data, and
spatial statistics. These models are based on trend surface analysis and
stepwise regression. We present results of trend surface models that describe the large-scale spatial variability. Models with small variance were
selected. In addition, the residuals from the trend surface model were then
modeled using regression classification trees (RTC). The final surfaces were
obtained by combining the trend surface model with RTC. Our research
program is using these new tools for forecasting the landscape-scale levels,
especially the ability to predict and map the cryptobiotic crusts with an R2
value of 67%. These models and spatial maps can be used for better resource management for such a large area like the GSENM.
KAMMERDIENER, SUSAN A.* and JAMES R. EHLERINGER.* University of Utah, Salt Lake City, UT, USA. The use of stable isotopes to
study urban nitrogen dynamics.
The urban-rural gradient provides an opportunity to compare plant nutrient
dynamics between systems with different sources and levels of available
nutrients. The similar location, vegetation, and topography of the Salt Lake
and Skull Valleys of northern Utah provide an excellent opportunity for a
study on the effects of urban pollution on the vegetation and soils of Salt
Lake City. NOx, is one of the main components of air pollution in the Salt
Lake Valley; therefore, it is important to monitor its deposition on plants
and soil. Measuring the stable isotope ratio of 15N /14N in plant and soil
materials is one way to do this. Results from the first season support the
hypothesis that the d15N in Bromus tectorum leaves is more enriched in
15
N in areas with high levels of atmospheric NOx deposition than in areas
with low levels of atmospheric NOx. The d15N in the vegetation decreased
significantly with distance from areas of high urban nitrogen deposition to
rural areas with lower concentrations of predicted deposition. Research
from the second season more thoroughly assesses the use of d15N as an
indicator of anthropogenic nitrogen deposition on soils and plants. In addition to leaf and soil samples from every square mile of the urban and
rural valleys, a set of genetically identical potted cheatgrass plants are
grown in the same soil at one urban and one rural site. The plant and soil
samples from the valleys and the potted samples are expected to follow
the pattern found during the first season. In addition, it is hypothesized that
the d15N in the soils supporting the Bromus tectorum will follow the same
pattern as the leaf samples, but will be more enriched.
KANG, SINKYU,1,* JOON KIM,2 FAITH ANN HEINSCH,1 MAOSHENG
ZHAO,1 JOHN S. KIMBALL1 and STEVEN W. RUNNING.1 1 Numerical
Terradynamic Simulation Group, School of Forestry, University of Montana, Missoula, MT, U.S.A.; 2 Department of Atmospheric Sciences, Yonsei
University, Seoul, Rep. Korea. Monitoring local and global vegetation
productivity and surface resistance using Moderate Imaging Spectrometer (MODIS).
Moderate Imaging Spectrometer (MODIS) provides improved remote sensing technology for monitoring global vegetation phenology and productivity, and surface resistance at 1km nadir resolution every eight days. As
well, these biophysical products can be used as inputs for global biosphere
models to improve our understanding of global carbon and water processes.
For this, MODIS biophysical products should be validated using ground
measurements, which represent at least 1km2 area. Currently, fluxtower
gives unique opportunity to provide ground truth data comparable with
MODIS 1km biophysical products. For this purpose, we selected representative fluxtower sites each from evergreen needleleaf forest (ENF), evergreen broadleaf forest (EBF), deciduous broadleaf forest (DBF), savannah,
and crop biomes, respectively. This study (1) demonstrates ability of MODIS to capture seasonal and spatial variability of GPP and surface resistance;
(2) validates MODIS GPP and surface resistance at fluxtower sites across
divers biomes; and (3) discusses practical considerations of the MODIS

products for monitoring terrestrial carbon and water processes at local and
global scales. We also evaluated the spatial patterns of the MODIS products
using fine-scale optical remote sensing data (IKONOS and Landsat ETM+),
incorporated with fluxtower measurements. At sites for which tower eddyflux measurements are available, MODIS GPP follows the general trend,
capturing onset of leaf growth, and in many cases, leaf senescence, while
tending to over-estimate total tower GPP. The MODIS GPP algorithm effectively captures the effects of stress events, such as late-summer drydown, on canopies.
KAPLAN, IAN* and MICKY D. EUBANKS. Auburn University,
kaplaia@auburn.edu, Auburn, AL. Mechanisms underlying variation in
the impact of a keystone predator: Effects of an ant-homopteran mutualism on the numerical response and per-capita effects of ants.
Dominant species, such as keystone predators, can exert tremendous influence on the distribution and abundance of other organisms. However, the
behavior and ensuing impact of these keystone species can be strongly
affected by the presence of key interactors, such as mutualists. Our previous
research documented that red imported fire ants, a keystone predator in
many ecosystems of the Southern U.S., have a greater impact on arthropod
food webs in the presence of honeydew-producing homopterans. These ants
are attracted to honeydew, a sugary solution which homopterans excrete,
and thus shift microhabitats from primarily soil-dwelling insects to foraging
on plant foliage. It is still unclear, however, why ants have stronger community-wide effects when co-occurring with homopterans. The results of
this study indicated that fire ants exhibit a strong numerical response to
aphids; cotton plants with aphids attracted 11X as many ants as plants
without aphids. Behavioral observations suggested that ants also behave
more aggressively in the presence of aphids. Fire ants were more likely to
attack and kill caterpillars while tending aphids than on aphid-free plants.
These experiments imply that both enhanced ant recruitment (i.e., numerical response to aphids) and increased ant aggression (i.e., variation in the
per-capita effect of individual ants) contribute to the keystone impact of
fire ants on insect communities associated with cotton plants.
KAPPEL, CARRIE V.,* FIORENZA MICHELI, KIMBERLY W. HEIMAN, GIACOMO C. OSIO, RAFAEL SAGARIN, SURESH SETHI and
ANDREW O. SHELTON. Stanford University, Palo Alto, CA. Shifts in
species dominance on rocky shores under human and natural disturbance regimes.
Through intensive biodiversity surveys of rocky intertidal habitats of Monterey Bay, CA, we assessed how human and natural disturbances interact
to affect these coastal communities. Specifically we examined whether differences in human disturbance and wave exposure lead to differences in
dominance patterns within these communities and whether disturbances
differ in their effects on common and rare species. The eight survey Areas
span a gradient in human disturbance, taking advantage of existing marine
reserves and other areas representing different levels of access and legal
restriction of human activities. Areas fell into 4 Categories: (1) open access,
no restrictions; (2) open access, reserve, no enforcement; (3) closed access,
no restrictions (de facto reserve); (4) closed access, reserve, restrictions
enforced. Each Area was divided into a wave-exposed and a protected Site
and Transects were stratified by tidal elevation within each. Relative rank
abundance patterns were compared among Areas, Sites, and Categories of
human disturbance. Results suggest that human disturbance through trampling, collecting, and other impacts leads to shifts in species dominance.
Sites with no restriction on access or harvest exhibited decreased dominance of common species and increased abundance of rare species relative
to reserves with restricted access and effective enforcement. This was most
apparent among sessile invertebrates and algae, i.e. those species that compete for primary space, suggesting that patchily distributed human disturbances open up primary space and decrease competitive dominance on
rocky shores. Differences between exposed and protected sites were less
striking than those for human disturbance, but sessile species did exhibit
decreased dominance and increased equitability in wave-exposed sites. Relative abundances of common species were most sensitive to differences in
disturbance. Rank abundance patterns among rare species were generally
similar among Areas and Sites, though identities of these species varied.
Mobile invertebrates exhibited similar patterns across human and physical
disturbance gradients.

KARBAN, RICHARD.* University of California - Davis, Davis, CA. Deciduous leaf drop reduces herbivory.
Trees in temperate latitudes tend to be deciduous while those in the mesic
tropics and at higher latitudes tend to retain green leaves throughout the
year. The deciduous habit is thought to be an adaptation to reduce snow
and ice loading in winter and reduce drought stress for broad-leaved species. In contrast, the evergreen habit is thought to be an adaptation to
increase the time available for photosynthesis, reduce rates of nutrient loss,
and increase nutrient recovery from older leaves. None of these current
explanations consider herbivory to be an important force in selecting for
seasonal deciduous leaf drop. Here I report that oak saplings (Quercus
lobata) that were experimentally forced to drop all of their leaves over the
winter supported much smaller populations (less than 50%) of herbivores
during the following spring compared to control saplings that naturally
retained some of their leaves. One important consequence of a deciduous
habit may be reducing rates of herbivory.
KAREL, IRENE S.,1,* VAL H. SMITH,1 BRYAN L. FOSTER,1 AMANDA
BROTHERS,2 MATTHEW RAMSPOTT1 and KEVIN PRICE.1 1 University of Kansas, Lawrence, KS, USA; 2 Kent State University, Kent, OH,
USA. Effects of experimental nutrient manipulations on a restored
grassland community.
Tallgrass prairie was once widespread in North America, but human activity
has reduced it to a small fraction of its former distribution. In recent years,
the threatened status of tallgrass prairies has inspired an interest in the
restoration of this habitat type. Here, we examine the effects of manipulation of nitrogen and phosphorus levels on the diversity and composition
of restored prairie plant communities at the Nelson Environmental Studies
Area (University of Kansas). Fifty-four separate 16m2 plots were sown with
prairie species in 1996, and we applied two levels of nitrogen depletion,
six levels of nitrogen enrichment, and a phosphorus enrichment treatment
in a factorial design. The plots are burned each spring to discourage the
invasion of woody species. In 2002, we found that species diversity increased significantly in response to nitrogen enrichment. Andropogon gerardii, a native C4 perennial grass, responded negatively to nitrogen addition, while Sorghastrum nutans was unaffected. Setaria faberii, an abundant non-native annual grass, increased significantly in response to nitrogen
addition, while leguminous forbs decreased significantly. Seasonally integrated NDVI (Normalized Difference Vegetation Index) values derived
from high-resolution spectral images of the site reveal a gradient of green
biomass and growth activity between the different treatments. In summary,
nitrogen addition affected overall species diversity, and has caused shifts
in species composition away from native prairie species and towards weedy
species.
KARIM, REZAUL,1 MASAHIKO SEKINE,2,* TAKAYA HIGUCHI,2
TSUYOSHI IMAI2 and MASAO UKITA.2 1 Bangladesh Institute of Technology, Khulna, Khulna, Bangladesh; 2 Yamaguchi University, Ube, Yamaguchi, Japan. Simulation of fish behavior and mortality in hypoxic
water in an enclosed bay.
A model of fish preference for environmental conditions (dissolved oxygen
(DO) and water temperature) and mortality was developed and coded within a combined hydrothermal and eutrophication model. The model was
applied to an enclosed bay located in the western part of Japan, where
eutrophication and associated onset of hypoxia at the bottom waters would
occur in every summer. A field survey of fish behavior under hypoxic and
non-hypoxic waters was conducted by releasing marbled sale in the bay
and tracked its movement. At the same time, vertical profiles of DO, salinity and water temperature were measured at every 0.5m vertical interval
at several locations along the tracking path of fish. Moreover, a series of
preference tests for DO, salinity and temperature in the laboratory were
conducted in order to decide preference parameters of fish. The model
could simulate reasonably the observed behavior and movement of the fish
under both hypoxic and non-hypoxic waters in the bay; thereby, validated
its applicability. Using the model, an assessment of the impact of the ongoing land reclamation project of about 400 ha in the bay on the fisheries
resources was also conducted. The results showed that a substantial mortality of the fisheries resources would take place when hypoxic condition

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175

occurs in every summer and the construction of the artificial land will lower
the mortality rate of fish during the summer period in the bay. Strong
offshore winds cause the upwelling of bottom hypoxic waters in summer
and the initial mortality of the fisheries resources increase under northerly
offshore wind of 5 m/s or more.
KARPANTY, SARAH M.* SUNY Stony Brook, Stony Brook, NY. Behavioral and ecological interactions of raptors and lemurs in Madagascar: A multiple-predator approach.
Predation has been a major topic of debate for ecologists interested in
understanding the origins and maintenance of social behavior in primates.
The most direct method to understand how predation structures primate
groups, populations, and communities and favors particular patterns of social behavior is to conduct studies of the predators themselves. Continuous
research since 1999 of the nesting and feeding ecology of the Madagascar
Buzzard (11 nests, 1,289 hours observation), Madagascar Harrier-Hawk (7
nests, 975 hrs.), and Hensts Goshawk (7 nests, 1,683 hrs.) in Ranomafana
National Park, Madagascar have identified the Harrier-Hawk and Goshawk
as significant lemur predators while Buzzards have never been observed to
feed on lemurs. Both Harrier-Hawks and Goshawks have been observed
feeding on Hapalemur griseus and Microcebus rufus while Hensts Goshawks have also fed frequently on Eulemur sp., Cheirogaleus major, and
Avahi laniger. A combination of raptor prey profiles, feeding rates, and
density with lemur demographics has allowed a quantification of the annual
predation rate by diurnal raptors on the lemur community in southeastern
Madagascar. These data demonstrating that the Hensts Goshawk is the major raptorial predator of the lemur guild corresponds with playback experiments of predator vocalizations that show lemurs exhibit the most intense
anti-predator behaviors to calls of the Goshawk. These predation rates will
be contrasted with other known sources of lemur mortality to emphasize
the potential significance of predation in lemur life history evolution, behavior, and community structure.
KAY, ADAM,1,* SCHADE JOHN,2 AMY NOVOTNY,2 DANIEL HERNANDEZ,1 MEGAN OGDAHL,1 ROBERT W. STERNER1 and SARAH
E. HOBBIE.1 1 Department of Ecology, Evolution and Behavior, University
of Minnesota, St. Paul, MN, USA; 2 Department of Zoology, Arizona State
University, Tempe, AZ, USA. Body size and N:P stoichiometry of insect
herbivores vary with host plant chemistry.
Recent evidence suggests that fast-growing animals contain an abundance
of P-rich ribosomal RNA and characteristically low ratios of N:P. If biochemical allocation depends on the relative availability of substrates, then
the elemental contents and growth rates of consumers may vary with the
C:N:P stoichiometry of their food. In this study, we investigated the relationship between the stoichiometries of two herbivores (the lace bug, Corythucha arcuata, and an unidentified aphid species) and the leaf N and P
contents of their host plant (Quercus macrocarpa). To obtain leaves of
variable quality, we sampled plots across a fire frequency gradient at different points in the growing season (June and August). We found correlated
shifts in oak leaf and lace bug chemistry over the summer, as both leaf P:
C and the P content of adult lace bugs were significantly higher in June
than they were in August. In August samples, the size of adult lace bugs
(measured as dry mass) was positively correlated with leaf C:N and C:P
ratios, and negatively correlated with leaf N:P. These results suggest that
lace bugs reached larger adult size by growing faster on leaves that are
relatively rich in P. However, because lace bug P content decreased with
body size, there was no relationship between lace bug %P and leaf C:P or
C:N ratios. Unlike lace bugs, same-sized aphids contained significantly
more P in August than they did in June, despite the lower C:P ratio in
August oak leaves. In addition, aphid %P in August was negatively correlated with leaf N:P, and was significantly lower in control plots than it
was in periodically burned plots. Our study provides the first demonstration
in a forest ecosystem of herbivore C:N:P stoichiometry adjusting to host
plant quality.
KAYE, THOMAS N.* and DAVID PYKE. Institute for Applied Ecology,
Corvallis, OR. Does correlation among vital rates matter? The effect
of correlation structure on estimates of population viability.
Transition matrix models are one of the most widely used tools for assessing population viability. Demographic parameters may be correlated across

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years and environments and inclusion of this correlation structure in stochastic models may be necessary to avoid overly optimistic estimates of
population viability. However, negative correlations among vital rates are
also possible, and these tend to counteract the effects of positive correlations. Therefore, the effect of correlation structure on population viability
estimates may depend on the nature of the correlations among vital rates,
which, in turn, may differ among species and environments. We used empirically derived data from 27 populations of five perennial plant species
collected over a span of five to ten years to examine the effect of correlation
among transition elements on population viability estimates. We also compared different methods of including stochasticity and checked for interactions between stochastic method and correlation structure. For each method, we estimated stochastic population growth rate (a measure of viability)
as our response variable. Temporal correlation among vital rates in our
stochastic matrix models altered estimates of population viability, but this
effect differed among species and was generally weak. The magnitude of
change in estimated stochastic growth rate for each species examined here
was largely explained by the ratio of positive to negative cross-correlations
of transition matrix elements; as the relative number of negative correlations decreased, the impact of correlation structure increased. When deciding whether or not to include such correlation structure in viability models,
conservationists may want to examine the correlations in their species as
a means of assessing their anticipated effect, and we provide a preliminary
linear regression model for doing so. These results are applicable to a range
of perennial plants and possibly other life histories.
KEARSLEY, MICHAEL J.C.,1,* HELEN YARD,2 NEIL COBB,1,3 DAVID
LIGHTFOOT,4 SANDRA BRANTLEY,4 JENNIFER FREY5 and GEOFF
CARPENTER.4 1 Department of Biological Sciences, Northern Arizona
University, Flagstaff, AZ; 2 Helen Yard Consulting, Flagstaff, AZ; 3 Merriam - Powell Center for Ecological Research, Northern Arizona University,
Flagstaff, AZ; 4 Biology Department, University of New Mexico, Albuquerque, NM; 5 Department of Fishery and Wildlife Sciences, New Mexico
State University, Las Cruces, NM. Integrated monitoring of terrestrial
riparian resources along the Colorado River in Grand Canyon.
Monitoring of diverse resource types in large and complex systems is often
done in piecemeal fashion, with tasks divided among unconnected groups
along taxonomic or methodological lines. Here we present the framework
for monitoring the effects of discharges from Glen Canyon Dam in Grand
Canyon National Park which recognizes the connectedness of many terrestrial elements in the Colorado River corridor. Joint sampling trips simultaneously collect data on habitat productivity (vegetation structure) and the
abundance and composition of breeding birds, waterfowl, small mammals,
herpetofauna and terrestrial invertebrates at a series of sites over the entire
year. Sites are revisited at one to three year intervals in an augmented
serially rotating panel design in order to balance change detection power
against observer impacts and canyon-wide representation. Sites are divided
into three zones based on hydrology: a pre-dam xeroriparian zone, a shoreline zone below the current range of non-emergency flows, and a post-dam
riparian zone between the two. Data collected in the first two years shows
several interesting patterns. First, vegetation density is more stable than we
would expect based on large differences in precipitation in 2001 and 2002.
Second, breeding bird densities within sites differ between years, but shift
among zones depending on the where invertebrates and other food sources
are greatest. Third, vegetation density sets an upper bound on breeding bird
density rather than predicting it outright. Fourth, mammal diversity and
density is highest in the pre-dam zone where productivity is lower. Fifth,
herpetofaunal densities and diversity are highest in the interfaces between
hydrologic zones, especially between the lower two. These patterns have
important implications for efficiencies and insight gained by monitoring
many resources in an integrated fashion.
KEELIN, CHERYL, SEAN OBRIEN and DANIEL R. TAUB.* Southwestern University, Georgetown, Texas. Competition between NAD-ME
and NADP-ME C4 grasses across a nitrogen gradient.
There are three distinct biochemical variants of C4 photosynthesis, each
performed by different C4 plant species. These three variants are called the
NAD-ME, NADP-ME and PCK variants, after the bundle sheath decarboxylation enzyme used in each pathway. Some previous research has sug-

gested that there may be differences amongst species using the different
pathways in responses to nitrogen, but very little work has been done to
explore this possibility. In this study we examined the competitive interactions of NAD-ME and NADP-ME species across a nitrogen gradient
under greenhouse conditions. We grew three NAD-ME grass species and
three NADP-ME grass species in every pairwise combination of an NADME and an NADP-ME species (nine combinations in all), as well as each
species individually. Plants were grown in pots at five different levels of
IBDU nitrogen fertilizer for six weeks, following which the aboveground
biomass was collected, dried and weighed. In monoculture, all species
showed a positive growth response to nitrogen. There were significant differences amongst the species in the slope of the regression of aboveground
biomass on nitrogen supply, but there was no consistent pattern of differences between the C4 variants. In six of the nine pairwise competition
treatments, the NAD-ME species increased as a proportion of aboveground
biomass as nitrogen supply increased. This provides qualified support to
the hypothesis that NAD-ME species are superior competitors to NADPME species under high nitrogen fertility conditions.
KEELING, MATT J.* Dept of Biological Sciences, University of Warwick,
Gibbet Hill Road, Coventry, West Midlands, West Midlands. Modeling
stochasticity: the use of moment closure techniques.
Ecologists and epidemiologists are increasingly focusing on stochasticity
and spatial heterogeneity as important biological factors. With high-powered computers simulation of such systems is a common modelling technique, however we lack a detailed understanding of the processes involved.
Moment closure approximations provide a simple method which can be
used to capture the main features of a wide variety of stochastic models
and to gain a more intuitive understanding. This presentation focuses on
the insights gained from such techniques, and in particular focuses on the
behaviour and persistence of finite metapopulations for two common ecological systems: a single species with density dependent growth, and an
enemy-victim interaction with epidemiological dynamics.
KEESING, FELICIA,1,3,* JACOB R. GOHEEN,2 RICHARD S. OSTFELD,3 BRIAN ALLAN4 and DARCY MISURELLI.5 1 Bard College, Annandale, NY, USA; 2 University of New Mexico, Albuquerque, NM; 3 Institute of Ecosystem Studies, Millbrook, NY; 4 Rutgers University, New
Brunswick, NJ; 5 Mpala Research Centre, Nanyuki, Kenya. Net effects of
large mammals on Acacia seedling survival in an African savanna.
Trees critically affect the functioning of savanna ecosystems through their
effects on nutrient cycling, water availability, and patterns of space use by
wildlife. Therefore, whatever factors influence successful recruitment of
tree seedlings are important filters controlling savanna ecosystem function.
In African savannas, large mammals, especially elephants, have been considered the most important agents of mortality for adult trees, but their
impacts on tree seedlings are not well-known. We investigated the effects
of large mammals on Acacia seedling survival by excluding large mammals
from replicated, 4-ha plots. In areas where large mammals were excluded,
rodents and invertebrates were more abundant and consumed approximately twice as many seedlings as on plots with large mammals. However,
seedlings in areas with large mammals were more likely to die of desiccation. Overall, net seedling survival was approximately twice as high in
the presence of large mammals. Our results indicate that large mammals
may indirectly increase seedling survival for a dominant savanna tree,
which in turn may influence tree recruitment and ecosystem function.
KELLER, EMILY A.,1,* THIMMAPPA S. ANEKONDA,2 BRUCE N.
SMITH,1 LEE D. HANSEN,1 J. BRAD ST. CLAIR3 and RICHARD S.
CRIDDLE.1 1 Brigham Young University, Provo, Utah; 2 Oregon State University, Corvallis, Oregon; 3 USDA Forest Service, Corvallis, Oregon.
Stress and respiration traits differ among four geographically distinct
Pinus ponderosa seed sources.
This study shows that plants adapt their metabolism to specific environmental temperatures (their native climate) and that plants respond to both
their environment and their genetics. This was done using calorimetry and
stable isotopic fractionation. Two-year old seedlings from each of four geographically distinct ponderosa pine (Pinus ponderosa var. ponderosa) seed

sources were studied. Two sources are from Oregon (Willamette and Deschutes) and two sources are from California (Mendocino and Eldorado).
Dark metabolic heat and CO2 production rates were measured on tissue
from elongating shoot tips at five temperatures from 15 to 35 degrees C.
Heat and CO2 production rates were not significantly different among the
four seed sources, but temperature coefficients of both heat and CO2 production rates were higher in the Deschutes seed source than in the other
three sources. The Deschutes seed source is the coldest, driest and most
variable climate of the four sources. Climates of the other three seed sources are very similar. The d 13C values were then obtained on the same
samples to determine the degree of stress associated with growth in different locations. Small differences in fractionation indicate variation in
plant stress. The fractionation ratios were lower in the Mendocino seed
source than in the other three sources, which were all very similar to each
other.
KELLER, TROY A.* and MARIA MARTINEZ. St Johns River Water
Management District, Palatka, FL, USA. Estimating lake-wide sediment
accumulation from historic bathymetry and radiometrically dated
cores.
Heterogeneity caused by water circulation patterns, lake morphometry, and
nutrient loading results in spatial variation in sedimentation. Sedimentation
rates are can be calculated from individual sediment cores by measured
mass accumulation and dating using radiometric analysis. Often the accuracy of lake-wide estimates of sedimentation is limited by the density and
spatial distribution of radiometrically-dated cores. To improve the spatial
accuracy of sedimentation rate estimates, we calculated sedimentation from
bathymetric surveys (1939 and 1996) of Lake Jesup, a 32 km2 lake, hydrologically linked to the St. Johns River. Results from 8 210Pb dated cores
show that 21.1 - 56.2 cm of sediment accrued over the 57 year period.
Mass sedimentation rates for these cores ranged from 6.13 to 92.3 dry
mg*cm-2*yr-1. Bathymetric comparisons indicated a mean sediment accumulation of 25.6 cm of muck (22 SD). The accumulation was highly variable and greatest in the deepest portions of the lake. A preliminary lakewide estimate of 55 mg*cm-2*yr-1 is nearly equivalent to the 58 mg*cm2
*yr-1 mean calculated from the dated cores. This spatial analysis technique
provides a powerful approach for estimating lake-wide sedimentation and
generates a detailed picture of the within basin heterogeneity of sediment
accumulation.
KELLEY, ALEXIA M.,1 HOWARD E. EPSTEIN1,* and DONALD A.
WALKER.2 1 University of Virginia, Charlottesville, VA; 2 Univeristy of
Alask Fairbanks, Fairbanks, AK. Plant community and ecosystem properties in Arctic frost-boil systems.
Frost boils are a type of patterned ground formation that is common in
arctic ecosystems. These landscape features, typically 0.5 to 3 m in diameter, are initially formed through differential freezing and thawing of
soils (cryoturbation) and may persist on the landscape for long periods of
time. The disruption associated with soil heaving causes frost boils to be
distinctly different in terms of plant community structure and soil biogeochemistry from the surrounding inter-boil areas. The properties of frostboil ecosystems are strongly influenced by regional climate gradients. In
this study we are investigating the interactions among nutrient cycling,
plant communities, and cryoturbation. Soil biogeochemistry and plant community traits of frost boils and inter-boil areas were examined along a
latitudinal climate gradient in arctic Alaska. Preliminary data from three
sites on the Arctic Slope of Alaska show that Normalized Difference Vegetation Index (NDVI) and Leaf Area Index (LAI) of frost boils (average
NDVI across sites 5 0.41, average LAI across sites 5 0.19) are less than
those of inter-boil areas (average NDVI 5 0.54, average LAI 5 0.88).
Both NDVI and LAI decrease with increasing latitude, however the NDVI
values on frost boils decrease more rapidly than those outside of frost boils.
Thaw depth is between 19 to 29 cm deeper in frost boils than inter-boil
areas and is on average 23 cm deeper at the more northern sites, as compared to the more southern study sites, due to the insulating effect of vegetation on soils. Soil moisture is lower in frost boils (mean of 42.3% by
volume) than inter-boil areas (mean of 70.2%), largely because of greater
mineral soil content and less organic matter in frost boil surface soils. This
work is part of a larger study that seeks to understand the relationships
among frost boils, climate, vegetation, and nutrient cycling.

Abstracts

177

KELLOGG, CHEV H.,1,2,* SCOTT D. BRIDGHAM3 and CHRIS B.

CRAFT.4 1 U.S. Geological Survey, Corvallis, OR; 2 Oregon State University, Corvallis, OR; 3 University of Oregon, Eugene, OR; 4 Indiana University, Bloomington, IN. Phalaris arundinacea invasion potential increases with soil successional changes.
Invasive plants increasingly threaten to disrupt plant communities and ecosystem function worldwide. Of particular concern in North American wetlands is the invasive grass Phalaris arundinacea (reed canary grass). In
wetlands this grass can invade and become the dominant member of the
plant community, both in restored and undrained wetlands. Although latesuccessional systems have traditionally been thought to be less invasible,
the invasion of undrained wetlands casts doubt on this assumption. A carbon accumulation rate, based on a 47-y chronosequence of restored wetlands, was used to develop a simulated successional gradient to test the
effects of soil successional changes on invasibility. We determined the effects of three levels of organic soil on germination of P. arundinacea seeds
in three water levels (-4, -1, +2 cm from soil surface). Additionally, the
effects of the same organic additions on both intraspecific and interspecific
(five species) competition with water levels 1 cm below the soil surface
were examined. Germination of P. arundinacea was increased by high
levels of organic soil in the driest hydrology and inhibited by intermediate
and high levels of organic matter in flooded hydrology. Aboveground biomass, belowground biomass, and number of new shoots of P. arundinacea
increased with increasing organic content of the soil. Leersia oryzoides
consistently had the greatest competitive effects on P. arundinacea. These
results indicate that successional changes in soils is sufficient to increase
invasibility of Midwestern marshes by P. arundinacea. From a management standpoint, planting or seeding of L. oryzoides may slow the invasion
of P. arundinacea. Additionally, control of hydrology based on organic
content of the soil can be effective in reducing germination rates of P.
arundinacea.
KELLY, CHARLENE N.,* JOSEPH B. MORTON and JONATHAN R.
CUMMING. West Virginia University, Morgantown, WV. Intra- and inter-specific variation of infectivity of isolates of several species of mycorrhizal fungi in response to aluminum exposure.
ABSTRACT: Mycorrhizal fungi (AMF) mediate interactions between
plants and soils, especially where nutrient or metal concentrations limit
plant growth. Variation in fungal response to specific soil conditions may
influence stress tolerance conferred to the host plants. Broomsedge (Andropogon virginicus) colonizes extreme sites, including acidic soils where
aluminum (Al) is phytotoxic. Aluminum resistance in Broomsedge is associated with colonization by acid-adapted AM fungus Glomus clarum.
These studies were extended by assessing inter- and intraspecific variation
of several species of mycorrhizal fungi in their ability to confer Al resistance to Broomsedge. Broomsedge seeds were planted in sand culture inoculated with 5 isolates of 3 species of fungi (Glomus clarum, Scutellospora heterogama, and Acaulospora morrowiae). Plants were exposed to 0
or 400 mM Al nutrient solution and grown for 4 or 9 weeks. Mycorrhizal
infection percentage (MIP) and biomass were measured. Al effects on infection depended on fungal species. Isolate-dependent variation of colonization within species was detected in 4-week-old plants. Of the Glomus
isolates tested, all exhibited increased colonization response to Al, yet only
one Scutellospora isolate showed increased infection. All isolates of Acaulospora were negatively impacted by Al. However, this intra-specific variation was not seen in plants grown 9 weeks.Biomass response to Al in
young plants was isolate-specific, whereas, plants grown for 9 weeks did
not exhibit isolate-dependent responses in biomass. This variation in Al
response in young plants may reflect spore germination inhibition by Al.
Some Al-sensitive isolates exhibited increased tolerance in age, reflecting
secondary colonization of roots by more tolerant hyphae. These patterns
may be attributed to selection of Al-resistant hyphal lines under Al exposure.
KELLY, STEPHANIE J.* University of Rhode Island, Kingston, RI. Estimating age and stage-specific demographic parameters for scup, Stenotomus chrysops, using model selection.
Identifying environmental and demographic processes that account for fluctuations in the abundance of organisms is a central goal of population

178

Abstracts

ecology. For commercially valuable fish species, such as scup (Stenotomus

chrysops) in Long Island Sound, understanding population dynamics is
important in developing regulations and management practices for the harvest of these species. Historically, changes in the abundance or harvest of
commercially important species have been difficult to predict because many
of the factors regulating their dynamics are poorly understood. In many
cases, it is the demographic rates of specific ages or stages that have the
largest influence on the growth rate and resulting population size. The goal
of this study was to determine the demographic structure of the scup population by identifying the number of ages/stages and vital rates justified by
census data. The strength of evidence in the empirical data from the Long
Island Sound Trawl Survey (LISTS) for age and stage-structured models
of scup was examined using maximum likelihood estimation and model
selection. Initial results indicate support for models that distinguish between two age/stage classes. Although scup can be aged quite accurately,
it appears that the identification of individual age classes may not be necessary to the development of accurate models. Additional population effects, including density dependence and environmental variance, will be
incorporated into the age/stage structured models to further evaluate scup
population dynamics. Results of this study will provide a more thorough
understanding of the demographic processes regulating the scup population
in Long Island Sound and serve as a basis for the development of more
effective management strategies.
KEMPES, CHRISTOPHER P.,1,* DAVID D. BRESHEARS,2 JULIANNA
E. FESSENDEN2 and JEFFREY M. HEIKOOP.2 1 McCurdy High School,
Espanola, NM; 2 Los Alamos National Laboratory, Los Alamos, NM. Variation in carbon isotope ratios in Pinus edulis trees following droughtinduced mortality.
Droughts of a high severity often cause a period of massive tree mortality
that may affect one or several species. In northern New Mexico both a
severe drought and a high mortality of Pinus edulis, a co-dominant of
pinyon-juniper woodlands, has recently been observed. The stress an individual tree is under in any given year may be reflected in the Carbon 13/
Carbon 12 ratios in tissue. When a tree is under stress due to either low
precipitation or high temperatures (i.e., drought) then that specific tree is
forced to close its stoma during the day and increase respiration rates overnight and in the process abandon the normal selection of Carbon 12 over
Carbon 13. The drought of the late 1990s /early 2000s is very similar to
that of the 1950s, in that there was massive mortality and pest infestations.
Obviously any trees that are over the age of 50 years survived the more
dramatic portion of the 1950s drought yet those same individual trees may
not have survived this past drought. We evaluated Carbon 13 relationships
from cores of recently deceased trees for different 4-year intervals, including tree age during a given period as a factor. We used a Walter index,
calculated from monthly temperature and precipitation data, to evaluate
relationships between Carbon 13 and periods of water stress. Our results
indicate that Carbon 13 ratios are related to the integrated periods of water
stress reflected in the Walter index. The results on how trees respond to
water stress have implications for reconstructing past climate conditions.
KENDALL, BRUCE E.,1,* GORDON A. FOX,2 GLEN WOOLFENDEN3
and JOHN W. FITZPATRICK.4 1 University of California, Santa Barbara,
CA; 2 University of South Florida, Tampa, FL; 3 Archbold Biological Station, Lake Placid, FL; 4 Cornell University, Ithaca, NY. Individual variation in demographic traits of Florida Scrub Jays.
Our prior theoretical work has demonstrated that variation in demographic
traits among individuals in a population can affect the uncertainty in the
populations demographic fate that is associated with demographic stochasticity. In particular, when an individuals trait is drawn at random and
is uncorrelated with traits of other individuals, then variation in mean fecundity may increase the uncertainty associated with demographic stochasticity in reproduction, but individual variation in survival probability
has no such effect. In contrast, variation associated with cryptic population
structure (so that individual traits are not entirely independent of one another) can reduce the uncertainty due to demographic stochasticity in survival, but may have no effect on fecundity. Here we quantify the magnitude
and effect of individual trait variability in a well-studied population of
Florida Scrub Jays from central Florida. These birds exhibit substantial

individual variation in demographic traits associated with factors such as

genotype, breeding history, territory quality, and the presence of helpers at
the nest. Were we to ignore this population structure (which we might have
to do in a small, poorly-studied population), we would introduce biases
into our estimates of the magnitudes of demographic stochasticity. This in
turn would bias our estimate of the extinction risk of the population.
KENNAMER, ROBERT A.* Savannah River Ecology Laboratory, Aiken,
SC, USA. Wood ducks of the Savannah River Site: Indicators of
changing environments.
Knowledge of the dynamics of biological populations and of their specific
responses to varying environmental conditions provides a foundation for
assessing the potential for projected climate change to reshape ecosystems
and their member components. Critical in the search to understand how
species might adapt to unfolding climate change will be the data from longterm population studies, as in this example using wood ducks (Aix sponsa)
breeding on the upper Atlantic Coastal Plain in west-central South Carolina.
In one investigation, surface waters in local wetlands that are the primary
habitats of wood ducks are shown to be under the influence of global-scale
climatological systems such as the El Nino-Southern Oscillation in the
tropical Pacific. Annual productivity of the breeding population then is
related to an annual index of wetland conditions across the 78,000-ha Savannah River Site, exemplified as higher duckling production in wetter
years. High levels of population productivity in wet years is realized
through increased production by individual females, attempted female
breeding at an earlier age, and a lessening of certain nest predation pressures. Wood ducks, like many organisms, rely on environmental cues to
time reproduction with resource availability. In a second investigation, the
timing of wood duck breeding is examined, showing that since the early
1970s, the annual onset of breeding has shifted almost a month earlier. This
shift in timing of reproduction appears to coincide with a general warming
of the late-fall and winter months over the period of study.
KENNARD, DEBORAH K.,1,* KENNETH OUTCALT,2 GRAEME
LOCKABY3 and ROBIN GOVERNO.3 1 USDA Forest Service, Southern
Research Station, Auburn, AL; 2 USDA Forest Service, Southern Research
Station, Athens, GA; 3 Auburn University, Auburn, AL. Spatial patterns
of fuel loads, fire intensity, and microbial biomass in a longleaf pine
forest.
Characterizing spatial patterns of fire intensity, as well as patterns of soil
resources following burning, is a potentially important and rarely considered means of understanding mechanisms of regeneration following fire.
Fire intensity can vary at fine spatial scales depending on local differences
in fuel loads, moisture, and wind. This spatial variation in fire intensity,
i.e., patchiness, is a characteristic feature of most fires, and provides a
critical mechanism to alter spatial patterns of regeneration. We characterized spatial patterns of pre-burn fuel loads, fire intensity, post-burn soil
microbial biomass, and post-burn fuel accumulation in a longleaf pine forest in the coastal plain of Alabama. These variables were measured at 100
spatially referenced points within a 10 ha plot. Pre- and post-burn fuel
loads were estimated using percent cover and height of live and dead fuels,
and point intercept counts of down-woody fuel. Maximum temperature
during the prescribed burn, which served as an indicator of fire intensity,
was measured using heat-indicating paints. Soil microbial biomass was
measured on soil sampled 6 weeks following the burn and analyzed using
the chloroform fumigation-extraction method. Geostatistical analysis
showed that variability of fuel loads, maximum fire temperature, and soil
microbial biomass indicators exhibited strong spatially explicit structures;
77-95% of their sample variance was spatially dependent. However, the
scale of this spatial dependence differed from 2 m (microbial biomass C),
to 13 m (fire intensity), to 33 m (fuel loads). These results suggest that fire
behavior can change significantly over relatively small scales and that patterns of fire intensity can only partially predicted by pre-burn fuel loads.
KENNEDY, THEODORE A.,1,* SARAH E. HOBBIE2 and JACQUES C.
FINLAY.2,* 1 Grand Canyon Monitoring and Research Center, USGS, Flagstaff, AZ, USA; 2 University of Minnesota, Saint Paul, MN, USA. Longitudinal and seasonal patterns of metabolism along a desert spring
stream.
Longitudinal patterns in physical, chemical, and biological variables are a
prominent feature of lotic ecosystems; however mechanistic studies of these

patterns have rarely been conducted in desert systems or across small spatial scales. We estimated whole stream Gross Primary Production (GPP)
and Respiration (R) along a 3 km reach of Jackrabbit Spring, a spring-fed
stream in the Mojave Desert of southern Nevada, using the single-station
dissolved oxygen change technique. A downstream decrease in GPP along
Jackrabbit Spring is present throughout the year that is associated with a
downstream decline in water temperature. Downstream increases in the
severity of nutrient limitation and rates of detritus deposition, that may
actually smother benthic algae, may also contribute to the longitudinal patterns in GPP. Overall, GPP was higher in spring and summer than fall or
winter. GPP was low (0-2 g O2.m-2.d-1) because of a combination of light
and phosphorus limitation of algal growth. A dense stand of invasive saltcedar (Tamarix ramosissima) that dominates a 1 km portion of the study
reach heavily shades Jackrabbit Spring, further reducing algal growth rates.
Downstream decreases, and seasonal changes, in R were also driven by
changes in water temperature. Longitudinal and seasonal patterns in R appear to drive across sites differences in organic matter accumulation. Additionally, a large manmade pool that is present mid-reach disrupts these
longitudinal patterns, increasing R and reducing GPP at sites further downstream. These results are consistent with predictions from the "serial discontinuity hypothesis", which suggests that longitudinal patterns and linkages in lotic ecosystems are disrupted by impoundments, whether they are
from a dam on a large regulated river or a pool in small stream.
KERBY, JACOB.1,2,* 1 University of California, Davis, CA; 2 California
State University, Northridge, CA. Factors limiting spread of the crayfish,
Procambarus clarkii, in Santa Monica Mountain streams.
Invasive species are a major threat to stream ecosystems; however, research
has seldom identified successful ways of prohibiting their spread. We surveyed 32 stream sections in the Santa Monica Mountains of southern California for two consecutive years to determine the distribution of the invasive crayfish, Procambarus clarkii. Surveys indicated that streams with
large obstructions (waterfalls, culverts, artificial structures) often did not
have crayfish present upstream of these obstructions. A mark recapture
study indicated that P. clarkii displayed significantly more downstream
than upstream movement when observed in three successive stream pools.
Obstructions significantly reduced movement of crayfish between pools
when compared to stream sections without any obstructions. The distribution of crayfish in streams and mark-recapture data thus suggest that
crayfish mainly spread downstream from a point of colonization and are
restricted in their movement to adjacent upstream sections by the presence
of both natural and artificial obstructions.
KERBY, JAY,* DAVID M. ENGLE and SAMUEL D. FUHLENDORF.
Department of Plant and Soil Sciences, Rangeland Ecology and Management, Oklahoma State University, Stillwater, OK, USA. Does foraging
behavior contribute to sexual segregation in bison across all hierarchical scales?
Many species of large ungulates are known to exhibit sexual segregation.
This segregation is expressed at multiple hierarchical levels (i.e. landscapes,
patches, and plants). North American Bison (Bos bison) exhibit sexual segregation in foraging behavior in their diet and within landscapes. Mature
male bison select a diet with higher proportions of warm-season grasses
than do mature female bison. Mature male bison also form independent
groups that preferentially utilize undisturbed habitats, but mature female
bison in mixed groups preferentially utilize recently disturbed habitats. Little is known about the foraging behavior of bison within patches and what
role that behavior plays in sexual segregation. Therefore, the objective of
our study was to investigate the foraging behavior of bison in heterogeneous patches created by a fire-grazing interaction. We describe attributes
of foraging paths of bison in multiple vegetation patches with contrasting
structure and composition through 3 seasons. Mature female bison adjust
foraging paths in response to differences in vegetation structure and composition among patch types and seasons. Mature male bison are generally
unresponsive to changes in vegetation structure and composition among
patches and seasons. These results show that bison exhibit patch-level segregation. This segregation may result from contrasting nutritional requirements of mature female and male bison. Differences in social dynamics of
male groups and mixed groups that include mature female bison may also
explain segregation of foraging behavior.

Abstracts

179

KERKHOFF, ANDREW J.,1,* BRIAN J. ENQUIST,1 JAMES J. ELSER2

and WILLIAM F. FAGAN.3 1 University of Arizona, Tucson, AZ; 2 Arizona
State University, Tempe, AZ; 3 University of Maryland, College Park, MD.
Latitudinal patterns of net primary production and the compensatory
effects of leaf stoichiometry.
Vegetation canopies, via primary production, are the energetic and material
drivers of all terrestrial ecosystems. A growing body of research suggests
that many biological and ecological process rates, including primary production, may be largely controlled by a small number of factors, including
body size, temperature, and C:N:P stoichiometry. However, interrelationships among site temperature, standing biomass, and organismal stoichiometry (as well as many other factors) complicate our ability to make
predictions about global patterns of productivity. For example, ecologists
generally consider the tropics to be substantially more productive than temperate, boreal, and arctic zones. However, adaptive changes in plant stoichiometry in response to colder environments and shorter growing seasons
may obscure, or even eliminate the expected latitudinal gradient in productivity. Here, we develop a simple model integrating allometric, kinetic,
and stoichiometric principles to generate testable predictions concerning
the biogeography of net primary production and plant stoichiometry. Global patterns of productivity and leaf stoichiometry generally support the
predictions made by the model. Specifically, trends in productivity expected
on the basis of temperature appear to be moderated by compensatory
changes in the nutrient content (especially P) of higher latitude plants. The
principles underlying metabolic scaling theory and ecological stoichiometry provide a mechanistic basis for scaling productivity estimates "from
leaf to globe" and for developing a predictive theoretical ecosystem ecology.
KERN, RUTH ANN* and RYAN P. LOPEZ. California State University,
Fresno, Fresno, CA. Fire intensity and shrub cover effects on microclimate and tree regeneration in Sierra Nevada Forests.
A field experiment was conducted to test the effects of fire intensity and
the presence of the montane shrubs Ceanothus cordulatus and Arctostaphylos patula on the growth and survival of Abies concolor and Pinus lambertiana seedlings. Nine experimental plots were located at 2200 m elevation within the Teakettle Experimental Forest, Sierra National Forest,
California, USA. Each plot was roughly circular, approximately 225 m2,
and had extant patches of C. cordulatus and A. patula as well as open
ground before treatment. The plots were subject to one of three burn treatments: hot, light, or no burn (control). In the first spring following the fall
burn treatments, one-year old bareroot A. concolor and P. lambertiana
seedlings were planted in the (former or current) patches of C. cordulatus,
A. patula and in the open. Plots were instrumented and monitored for air
and soil temperature as well as surface and seedling rooting depth soil
moisture. In this way, we intended to study the possible facilitating effects
of nitrogen enhancement in the proximity of C. cordulatus, root grafting
with A. patula, or simple shade vs. the possible competing effects of shrub
presence. Extreme drought led to a small seedling survival rate (2.5%),
compromising meaningful statistical analysis of seedling growth and survival data. There was a positive correlation between seedling growth and
soil moisture, which was highest in the unburned plots. A high proportion
of the surviving seedlings were P. lambertiana, suggesting greater adaptation to hot, dry conditions than A. concolor seedlings. Survivorship and
microclimate data suggest that current conditions present severe difficulty
for tree regeneration. With low moisture and the exposed ridgetop location,
the effect of fire may be to convert patches of forest to shrub community,
thereby resetting the pathway of succession.
KESAVARAJU, BANUGOPAN,* KATIE S. COSTANZO and STEVEN
A. JULIANO. Illinois State University, Normal, IL, USA. Interspecific
differerences in mosquito behavioral responses to water-borne cues to
predation : Costs and benefits.
Aquatic prey often detect the presence of a predator via chemical cues, and
react by showing reduced activity level. The mosquito Toxorhynchites rutilus preys upon other container dwelling insects, including larvae of Aedes
albopictus, which is native to Asia but was introduced into USA. Previous
work has established that the native container mosquito Ochlerotatus tris-

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eriatus adopts low risk behaviors, reducing movement and feeding, in the
presence of predation risk from T. rutilus. It is unknown whether introduced
A. albopictus show similar responses to this predator. Behavior of 4th instar
A. albopictus or O. triseriatus was recorded in water that had held either
A. albopictus or O. triseriatus larvae, respectively, alone (control) and in
water that had held T. rutilus larvae feeding on either A. albopictus or O.
triseriatus, respectively, (predation). Activity and position of larvae were
recorded in 30 minute instantaneous scan censuses. In response to waterborne cues to predation, O. triseriatus adopted low-risk behaviors (more
resting, less feeding and movement), but A. albopictus showed no significant change in behavior. We reared both species in predation and control
waters to assess the costs of low risk behaviors for O. triseriatus. We
predict that development rate and mass at pupation for O. triseriatus, but
not for A. albopictus, will be lower in predation water. We also compared
impact of predation on both species in the two treatment waters to assess
the benefits of low risk behaviors to O. triseriatus. We predict that survivorship of O. triseriatus will be greater than that of A. albopictus in predation water.
KESSLER, ANDRE* and IAN T. BALDWIN.* Max-Planck-Institute for
Chemical Ecology, Jena, Thuringia, Germany. The emission of herbivore
induced phytochemicals and their effect on the insect community of
Nicotiana attenuata in natural populations.
Herbivore attack causes plants to release volatile phytochemicals that can
function as attractants for predators or parasitoids of herbivores and potentially influence plant fitness. However, while a majority of the previous
work has focused on genetically uniform crop plants and genetic variation
in the volatile organic compound (VOC) emission is well described, the
ecological relevance of induced emissions for natural plant populations
remains unclear. We present a repeatable, quantitative method for the collection of VOCs from the headspace of Nicotiana attenuata plants growing
in natural populations. Different natural herbivores were used to elicit VOC
emission to evaluate the signaling function of that emission for natural
enemies of the herbivores. On wild growing N. attenuata we tested the
attractiveness of singly applied VOCs for natural occurring predators and
found that the 5 to 10 fold higher emission of certain compounds (cis-3hexene-1-ol, cis-3-hexenyl butyrate, linalool, bergamotene) and the application of methyl jasmonate (MeJA) can increase predation of Manduca
sexta eggs and 1st instar larvae by Geocoris spec.(Heteroptera). Moreover
we tested the effect of "doping" plants with single VOCs for their effect
on ovipositing adult Manduca quinquemaculata. The moths avoided ovipositing on plants with increased linalool emission and on plants treated
with MeJA or attacked previously by caterpillars. The plant not only mediates the described tritrophic interactions but also competitive interactions
between the leaf bug Tupiocoris notatus and M. quinquemaculata. Our
results demonstrate how induced VOCs are used by both the herbivore and
its generalist predator and hence are capable of influencing the composition
of N. attenuatas herbivore community.
KICKLIGHTER, CYNTHIA E.* and MARK E. HAY. Georgia Institute of
Technology, School of Biology, 310 Ferst Dr., Atlanta, GA, 30332-0230.
Opening up a can of worms: Prey traits that influence the use of predator deterrence strategies.
Antipredation strategies that deter consumers, such as the use of chemical
and structural deterrents, commonly allow species to persist in predatorrich environments. However, little is known about the characteristics of
prey species that rely on these strategies. To investigate this question, marine worms (e.g., annelids, hemichordates) collected from soft-sediment
and hard-substrate habitats from various locations in the Caribbean and the
east coast of the US were examined for palatability to sympatric fishes and
a crab. Chemical and structural traits, refuge use and prey mobility were
examined for the role they play in determining palatability and susceptibility to consumers. Results indicate that species with body parts exposed
to predation for long periods of time (such as during feeding and/or respiration) are unpalatable and chemically or structurally-defended against
predators while species that live completely buried beneath the sediment,
or inhabit a strong tube, are more palatable. Moreover, for some refuging
species, body parts that are in physical refuges are palatable and lack deterrents while more exposed body parts are unpalatable and possess chem-

ical or structural deterrents. Mobility appears to be an important factor as

well. Species that are more mobile and, thus, may be able to avoid predation through escape, are more palatable and lack deterrents compared to
species with restricted movement. In addition, the use of deterrence strategies within a taxonomic group is not necessarily uniform. Not all members
may utilize a chemical defense or there may be variation in which body
parts are defended. Our findings indicate that ecological exposure to predation is a prevalent characteristic of prey that utilize predator deterrence
strategies and susceptibility to predation may override taxonomic trends in
the use of predator deterrence.
KILHAM, SUSAN S.,1,* CATHERINE M. PRINGLE,2 KAREN R. LIPS,3
DANA DRAKE2 and ANTHONY W. RANVESTEL.3 1 Department Bioscience & Biotechnology, Drexel University, Philadelphia, PA 19104,
USA; 2 Institute of Ecology, University of Georgia, Athens, GA 30602;
3
Department of Zoology, Southern illinois University, Carbondale, IL
62901. A pre-extirpation study of the role of stream-dwelling frogs in
neotropical foodweb dynamics.
Stable isotope analyses were used to examine trophic transfer in a neotropical montane stream characterized by abundant larval anurans. The goal
was to assess the role of stream-dwelling frogs in food web dynamics
before they become extirpated, as has already occurred in many regions
throughout the neotropics. We selected a study site in the headwaters of
the Rio Guabal in the Parque Nacional Omar Torrijos H., El Cope, Cocle,
Panama. The site is ca. 700 masl on the Atlantic slope of central Panama,
and is about 300 km east of the closest documented site of frog extirpations.
Samples were collected from May to August, 2000. The d15N in the first
and second trophic levels were a function of rainfall, decreasing as rainfall
increased. Food web fractionation of d15N was much less on average
(1.9per trophic step) in these tropical stream food webs than in well
documented lake food webs (3.4). The d13C of autotrophs appeared to
be affected by stream flow. Our results are important baseline data that will
be used to make predictions regarding how trophic structure will change
post-extirpation, which is apparently just beginning in this area.
KILLILEA, MARY E.,* STEPHEN DEGLORIA, SUSAN RIHA and WILLIAM PHILPOT. Cornell University, Ithaca, NY, USA. Carbon sequestration in New York State forestland.
Quantifying the amount of carbon currently being sequestered in New York
State forests and evaluating how forest site variability and management
practices affect carbon sequestration are important (Cannell 1999). This is
especially true if there are potential economic and environmental benefits
to landowners. Globally, forest ecosystems are estimated to store 330 GtC
in the vegetation and 660 GtC in the soil, and are an important part of the
global carbon budget (IPCC WGI 1996). The percentage of land in New
York State covered by forests has been steadily increasing over the last
100 years, which has led to increased carbon accumulation in areas of
reforestation (Caspersen et al. 2000). Several models (e.g. CENTURY,
TEM, and Biome-BGC) have been created to increase our understanding
of carbon dynamics (Parton et al. 1993, McGuire et al. 1995, Running and
Coughlan 1988, Cramer et al. 1999). Biome-BGC will be used for this
study because leaf area index (LAI) is the primary vegetation input, which
will allow the model to be spatially scaled-up in the future (Running and
Gower 1991 and Running and Coughlan 1988). Currently carbon sequestration is being estimated for 52 sites in the Hudson River Valley of New
York State. During the summers of 2001 and 2002, LAI measurements
were made at each of the 52 sites using the CI-110 Digital Plant Canopy
Imager. In addition to LAI, site, soil, climate, and vegetation data are necessary to run the model. The forest type, latitude, longitude, slope, and
aspect were recorded at each site. Soil data (i.e. effective soil depth, and
percentage of sand, silt and clay) will be extracted from the STATSGO
database. Temperature and precipitation data for the region is available
from the Northeast Regional Climate Center, and additional climate variables can be calculated. Biome-BGC will be initially run using inputs for
current environmental conditions, providing a baseline value of annual carbon stored in the vegetation. Then the input parameters can be manipulated
to examine the effects of various management decisions and potential
changes in environmental conditions (e.g. increased temperature or decreased precipitation).

KILLINGBECK, KEITH T. University of Rhode Island, Kingston, RI;

keith@uri.edu. Models of the determinants of realized and potential
nutrient resorption.
Nutrient resorption, the withdrawal of nutrients from senescing plant tissues, is a complex process whose physiology and ecology have been scrutinized widely. Nonetheless, resorption remains somewhat of an enigma.
Even rudimentary questions such as whether site fertility has significant,
predictable impacts on resorption remain unresolved. However, recent theoretical and empirical advances have led to valuable insights into mechanisms that may account for interspecies and intersite variability in resorption. What has not emerged in the years following the classic paper by
Chapin and Kedrowski (1983) that sparked renewed interest in resorption,
are models that portray the relationships between resorption and the factors
by which it is influenced. Two such qualitative models are offered here to
serve as a framework for an analysis of the mechanisms that control resorption. These predictive models are comprised primarily of testable hypotheses that consider the most important factors known to, or suspected
to, influence both potential and realized resorption. Potential resorption, the
quantity of a nutrient that can physiologically be resorbed from senescing
tissue, is determined by biochemical constraints imposed on nutrient retrieval, and by genetic controls shaped by phylogeny and a suite of plant
and site characteristics. Realized resorption, the quantity of a nutrient that
is actually resorbed, is influenced by potential resorption, and by a wide
array of determinants that are primarily environmental, not genetic. Although realized resorption may reach potential resorption, it is likely that
realized resorption is frequently reduced below potential resorption by conditions that terminate, or impede the retrieval of nutrients from senescing
leaves. Realized resorption will be less than, or equal to potential resorption, but never higher. Because there appears to be no obvious advantage
to inefficient or unproficient realized resorption, it seems likely that selection continually favors adaptations that allow realized resorption to reach,
or closely approach, potential resorption.
KILPATRICK, A. MARM.* University of Wisconsin-Madison, Madison,
WI, USA. The effects of predation and disease on the evolution of
resistance to malaria in Hawaiian honeycreepers (Drepanidinae).
The response of a population to selective pressure is strongly influenced
by other sources of mortality which may act orthogonally to the selective
agent. In the case of predation and disease, the evolution of resistance may
be stymied by non-selective predation, or even halted by a predation-disease interaction. I combined field data with a model to examine the evolution of resistance to avian malaria in Hawaiian honeycreepers. The results
offer insight into which species might be expected to evolve resistance to
malaria and what can be done to facilitate the evolution of resistance.
Predation can have a significant role in altering the temporal course, and
in some circumstances the outcome, of resistance evolution. The results I
will present have broad applicability to other disease-host interactions, especially recently introduced or emerging infectious diseases.
KIM, DONGGIL,1,* SHOU GUO MU,1 DOWON LEE,1 NAMYI CHAI2
and JOON KIM.2 1 Department of Environmental Planning, Graduate
School of Environmental Studies, Seoul National University, Seoul, Seoul,
Korea; 2 Department of Atmospheric Science, Yonsei University, Seoul,
Seoul, Korea. Effects of a pulsing water addition to soil on CO2 efflux.
The objectives of this study was to examine how an abrupt increase of soil
water by rainfall affects soil respiration. We measured soil respiration and
various environmental factors in deciduous, coniferous and mixed forest
sites over two years. We also conducted mesocosm experiments to clarify
how a pulse of water addition to soil affects CO2 evolution. Based on the
field measurements, we derived a regression model to estimate soil respiration with soil temperature. However, observed values of soil respiration
were higher just after summer rainfall events (about 15;30%) than model
predictions in the deciduous and mixed forest sites, and an opposite tendency was observed in the coniferous forest sites. A mesocosm experiment
showed that increase of soil respiration after rapid increase of soil water
content had significant positive correlation with soil temperature in the
range of 10 and 20 degrees C. On the other hand, soil respiration decreased
with increasing water content at 4 degrees C. Also soil respiration was

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181

positively correlated with organic matter content in the range of 10 and 20

degrees C. Our results indicate that the effects of an abrupt increase of soil
water on CO2 efflux is dependent on temperature and soil organic matter
in fields.
KIM, HYUN-SEOK,1,2 RAM OREN1 and THOMAS M. HINCKLEY.2
Duke University, Durham, NC, USA; 2 University of Washington, Seattle,
WA, USA. Comparison between potential and actual net assimilation
of hybrid poplar in eastern Washington.
1

Potential and actual net carbon assimilation of 3-year-old, 8.5 m tall Populus trichocarpa x P. deltoides hybrids was estimated at an irrigated plantation in Eastern Washington. A Conductance-Constrained Carbon Assimilation (4C-A) model was employed based on sapflux-scaled conductance.
Although trees were irrigated 24 hours a day, there was clear evidence of
a significant soil water potential decrease below optimum levels as indicated from predawn leaf water potential. By assuming a constant nonlimiting soil water potential (-0.2 MPa) throughout the season, we estimated potential stomatal conductance based on relationships with photosynthetically active radiation, vapor pressure deficit and boundary layer
conductance. Actual net assimilation (June 14th - October 16th, 125 days)
was estimated at 1688 g C m-2, 49% of which was used for above ground
biomass production (woody biomass: 768 g C m-2; foliage biomass: 55 g
C m-2). Potential net assimilation was estimated at ;16% more than actual
(1953 g C m-2). Assuming that partitioning to wood production remains the
same without soil moisture limitation, this would add 252 g m-2 ground.
Considering the allometrics of this poplar hybrid, its average height at age
4 would have increased from an actual of 10.8 m to a potential of 11 m.
KIM, IN SUNG,1,* YONG-JOO CHO,1 EUN JU LEE,1 HONG KEUN
CHOI2 and CHANG KYUN KIM.2 1 iskim07@snu.ac.kr, Seoul National
University, Seoul, Seoul, S. Korea; 2 Ajou University, Suwon, Kyunggi, S.
Korea. Effects of waste landfill leachate on the growth of Phragmites
australis.
This study intended to investigate the effects of waste landfill leachate on
the growth of Phragmites australis and to analysis the genetic diversity
within and among populations of P. australis in Korea. The study was
carried out in a pilot-scale constructed wetland and selected reed populations from 14 sites in Korea. The experiments were conducted at two different conditions: one was treated with leachate and the other with diluted
by groundwater as a control. Several morphological and physiological characteristics were examined. It was proved that flooding with leachate significantly influenced the growth of reed. Shoot growth, biomass and chlorophyll content grown on pilot treated with leachate were lower than those
of control treatment. However, the populations from landfill sites showed
better growth rate than other populations. Also, genetic diversity within
and between populations were investigated using RAPD analysis. The pherogram was generated by unweighted pair group mathematical average
(UPGMA) cluster analysis of RAPD data. As a result, populations were
divided into four clusters, although clusters did not represent the geopraphical relationship among 14 populations. The genetic diversity within and
between popultions were compared using AMOVA. The genetic variance
within the popultions showed higher genetic diversity than between populations. Generally, a higher genetic diversity was detected in the populations from landfill sites, suggesting that growth conditions of P. australis
could be correlated with genetic diversity.
KIM, RAE-HYUN,1,* YOWHAN SON1 and JONG-HWAN LIM.2 1 Korea
University, Seoul, Korea; 2 Korea Forest Research Institute, Seoul, Korea.
Mass and nutrient content of coarse woody debris in a deciduous forest
of central Korea.
Mass and nutrient contents of coarse woody debris (CWD) were investigated in a natural mixed deciduous forest of central Korea. Also all CWD
was assigned to a decay class (class 1 (least decomposed) to 5 (most highly
decomposed)). Total CWD mass was 20.7 Mg/ha in March, 2002. Carpinus
laxiflora showed the highest mass (9.6 Mg/ha) followed by Quercus serrata
(8.8 Mg/ha) and C. cordata (0.7 Mg/ha). The store of CWD decreased to
19.2 Mg/ha after the 6 month study period. The initial (March) and final
(September) N, P, K, Na, Ca, and Mg contents (kg/ha) for decomposing

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Abstracts

CWD were 15.2 and 16.8, 1.0 and 1.9, 3.9 and 7.7, 0.1 and 0.6, 30.9 and
30.8, and 1.4 and 1.5, respectively. CWD characteristics in a decay class
such as presence of bark, branches, and consistency of wood varied by
species. In general, bark showed the highest decomposition rate, however,
the bark of C. cordata remained to the decay class 5. CWD density decreased with the decay class. Nitrogen and P concentrations in CWD increased with decay class while K, Na, Ca and Mg concentrations showed
no significant changes by decay class.
KIMBRELL, TRISTAN S.* and ROBERT D. HOLT. University of Florida,
Gainesville, FL, USA. Evolution, predator switching, and population
stability: An individual-based approach.
Predator switching has traditionally been viewed as a stabilizing mechanism in multispecies communities. However, recent models have suggested
that predator switching may destabilize population dynamics and even lead
to species extinction. Most of this work has been based on the elaboration
of standard predator-prey models represented as systems of differential
equations. We examined the dynamical and evolutionary implications of
predator switching using a spatially-explicit, individual-based model, in
which switching involved movement between two habitat patches. We compared three different predator types: predators that randomly moved between patches, predators that optimally switched between patches, and
predators with incomplete information that sampled the patches and used
a memory rule to determine when to switch patches. We first explored how
each switching strategy impacted population dynamics. We then allowed
alternative switching strategies to compete, and examined the potential for
sustained coexistence of alternative switching strategies. Evolution was incorporated by adding a mutation rule for the switch decision variable. An
adaptive dynamics approach suggests that switching strategies that are allowed to evolve often result in population dynamics that are more stable
than non-evolving strategies.
KING, AARON A.1,* and SHANDELLE M. HENSON.2 1 Ecology & Evolutionary Biology, University of Tennessee, Knoxville, TN; 2 Department
of Mathematics, Andrews University, Berrien Springs, MI. Which deterministic skeleton? Multiple skeletons and lattice effects in stochastic
population models.
Gone are the days when theoretical ecologists could simply ignore stochasticity. Deterministic models, however, remain very important. Specifically, the dynamics of stochastic models, though often strikingly different
from their deterministic counterparts, can nevertheless frequently be understood with reference to transients or unstable invariant sets in deterministic models. I will illustrate this point with a few examples. In general,
the process of adding noise to a deterministic model is a well-defined recipe
for creating a stochastic model, but the inverse procedure, of "subtracting"
the noise from a stochastic model, is not well-defined. The upshot is that
there can be multiple "generalized deterministic skeletons" corresponding
to a given stochastic model. Although in cases of interest the differences
among these may be small, but the small differences can have surprising
consequences.
KING, ELIZABETH G.* Section of Evolution and Ecology, University of
California, Davis, CA, U.S.A. Aloe secundiflora shrubs facilitate a native
perennial grass in degraded Kenyan rangelands: Implications for restoration.
Over 75% of Kenya is arid and semi-arid land. Human and livestock pressure has led to severe vegetation degradation and soil erosion across vast
areas of Kenyas drylands. I hypothesize the presence of Aloe secundiflora
shrubs positively affects the success of perennial grasses and other vegetation in degraded rangelands. If so, propagating and planting aloes may
be an effective tool in restoration efforts. An observational vegetation study
in the Kerio Valley, Kenya, indicated that vegetation cover, litter cover, soil
retention, and soil seed bank were all greater in the immediate vicinity of
naturally occurring A. secundiflora shrubs. The positive effect of aloes was
likely due to physical protection, shade, and perennial ground cover offered
by the aloes. To test whether mature transplanted A. secundiflora shrubs
can provide similar ecological benefits, I performed a manipulative experiment in northern Laikipia District, Kenya, in which seeds of the native

perennial grass Cenchrus ciliaris were planted next to three different facilitator treatments: around transplanted aloe shrubs, around small piles of
thorny Acacia tortilis branches, and in bare areas with no facilitator. The
success of grasses at each life history stage was compared among the three
facilitator treatments. Aloe presence significantly enhanced C. ciliaris seedling emergence rate, seedling survival, and adult plant survival, while thorn
protection enhanced some of these fitness aspects to a lesser degree than
aloes. Findings indicate that propagating and planting aloes may indeed be
a useful restoration tactic, especially when combined with perennial grass
seed planting. If utilized, this restoration tactic can offer rural communities
additional benefits, because valuable sap can be legally and sustainably
harvested from propagated aloes as the basis of a rural economic enterprise.
KING, RACHEL T.* University of Miami, Coral Gables, Florida. Effects
of succession and microelevation on seedling establishment in a Peruvian river meander forest.
I investigated the effect of river meander microhabitats differing in successional stage and microelevation on seedling establishment of Calophyllum brasiliense, a common canopy tree of seasonally flooded forest along
the Manu River meander zone in Peru. To compare seedling establishment
between microhabitat types, I planted C. brasiliense seeds in six microhabitat conditions combining the effects of 3 successional stages (early,
mid, and mature) and 2 microelevations (ridges and swales). Seedling establishment success in this study was affected by both successional stage
and by microelevation, but microelevation was mostly important in midsuccessional habitats. In general, seedlings in early succession experienced
better conditions than in mature forest; light was higher, herbivory lower
and seedling growth and survival higher. In mid-successional forest, microelevation determined habitat quality; swales had higher light, lower herbivory and higher seedling success than ridges. Mid-successional swales
were similar in quality to early successional forest for seedling establishment, while ridges in that successional stage were the poorest microhabitats
for establishment. Although mid-successional swales are similar to early
succession for seedling establishment, in the long term, seedlings that establish in the mid-succession may have less chance of reaching reproductive size before their habitat ages to mature forest than members of their
cohort that established in early succession. I hypothesize that successful
recruitment of C. brasiliense in the Manu river meander system requires
dispersal to early successional habitat.
KINKEAD, KAREN E.,1,* DAVID L. OTIS2 and ALBERT G. ABBOTT.1
1
Clemson University, Clemson, SC, USA; 2 Iowa State University, Ames,
IA, USA. Genetic variation among Ambystoma populations associated
with restored wetlands.
Several degraded Carolina bay wetlands on the Savannah River Site have
been subjected to experimental management in an effort to functionally
restore these valuable habitats. This project evaluates responses of salamander populations to this restoration effort. Carolina bays are important
breeding habitat for two species of pond breeding salamanders: Ambystoma
talpoideum (mole salamanders) and A. opacum (marbled salamanders). Sixteen bays have been monitored: three reference bays (functionally intact),
three control bays (with active drainage ditches), six treatment bays, and
four intact bays near two of the treatment bays (in effect creating two metapopulations). Distances between these bays range from less than 200 meters
to greater than 20 kilometers. The first year of the study was a pre-restoration year, the bays were then restored during 2001, and 2002 was a postrestoration year. Each bay is partially encircled with drift fences and pitfall
traps. Salamanders at each bay were captured, batch marked, and released
between January and July, 2000-2002. Individuals were given visible implanted elastomer batch marks that designated the year and bay of capture,
the age (adult or juvenile), and whether it was captured entering or leaving
the wetland. Of the more than 4000 mole and marbled salamander captures,
fewer than 35 have been captured at more than one bay. Although traditional capture-recapture methodology can provide breeding population size
estimates and apparent survival rates, estimation of dispersal between bays
is more problematic. Therefore, in addition we are experimenting with the
use of amplified fragment length polymorphisms (AFLPs) to estimate relatedness among and within populations of mole salamanders and also of
marbled salamanders.

KINTER, CECILIA L.* and RICHARD N. MACK. Washington State University, lkinter@wsu.edu, Pullman, WA. Genetic sampling of emigrants
affects plant height and vigor in new ranges.
For two centuries, the tallest, most vigorous members of plant species have
often been reported in new, rather than native, geographic ranges of the
species. Attempts to evaluate whether these observations underlie a generalization about the performance of alien plants have produced equivocal
results. Where differences have been documented, they have been attributed
to environmental factors in the new range: greater resource availability, or
release from competition, predation, grazing, or parasitism, or to post-introduction evolution. Several hypotheses have been proposed, including
Enemy Release, Environmental Constraint, Historic Herbivore Pressure,
Optimal Defense, and Evolution of Increased Competitive Ability. None
of these hypotheses has provided a comprehensive explanation for the varied performance of alien plants. Alternatively, we asked whether the geographic source of immigrant populations affects subsequent performance
by comparing height and vigor of the annual grass Bromus tectorum L.
(cheatgrass, downy brome) from native European populations, highly invasive North America populations, and New Zealand populations which
are naturalized, but not invasive. In four common greenhouse experiments,
North American plants typically were tallest and most vigorous, followed
by European and lastly New Zealand plants (ANOVA, p , 0.05). These
differences in performance cannot be attributed to environmental or maternal effects, and so must be due to heritable differences among plants
from each region. Furthermore, these genetic differences are products of
the different locales in the donor ranges from which emigrant populations
were gathered. We found no evidence of the evolution of novel phenotypes.
Phenotypic trait values for both sets of introduced plants form discrete
subsets (Canonical Discriminant Function Analysis, p , 0.0002) of values
for the native range. Our results hold implications for prediction of whether
a species will become invasive in a new range, as well as for biosecurity,
quarantine regulations, herbicide efficacy, biocontrol, and the conservation
of biodiversity.
KINZIG, ANN P.1,2,* and CHARLES L. REDMAN.2 1 Arizona State University, Department of Biology, Tempe, AZ; 2 Arizona State University,
Center for Environmental Studies, Tempe, AZ. Agrarian landscapes in
transition: The case of central Arizona.
The patterns humans impose on the Earth through purposeful and inadvertent land-use change are fundamental determinants of local, regional, and
global ecological processes that ultimately influence the sustainability of
both biological and cultural landscapes, and thus human quality of life. The
introduction, spread, and abandonment of agriculture represents the most
pervasive alteration of the Earths environment during the past 10,000
years, affecting 2/3 of the Earths terrestrial surface. The transitions of
agrarian landscapes and life ways continue to take many forms, ranging
from abandonment to urban development to more intensified agriculture.
Our central objective is to understand what happens when humans impose
their spatial and temporal signatures on ecological regimes and must then
respond to the systems they have helped create, further altering the dynamics of the coupled system and the potential for ecological and social resilience. We are studying this question in a comparative context across six
different LTER sites, each with different agricultural histories and residing
in different biogeographic regions of the nation. In this presentation, we
give a broad overview of the objectives of the study and the sites involved,
and provide a narrative of agrarian transformations in central and southern
Arizona. In spite of being in an arid landscape with low precipitation, the
central and southern Arizona landscapes consist of nearly 2 million acres
of highly productive farmland today. Around the rapidly urbanizing area
of Phoenix, much of this farmland is being converted to residential settlements, while some is abandoned to desert conversion. We offer an historic
perspective on these conversionsfrom the pre-historic Native American
habitations to the historic Anglo occupationsand discuss some of the
implications for ecological processes.
KIRKMAN, KATHERINE* and ROBERT MITCHELL.* Jones Ecological
Research Center, Rt. 2 Box 2324, Newton, GA, USA. Are limited oldgrowth overstory systems useful in guiding restoration and conservation management of longleaf pine ecosystems?
Old growth canopies of longleaf pine (Pinus palustris) are useful in demonstrating the extreme longevity of the dominant species and in quantifi-

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183

cation of canopy disturbance patterns due to tropical storms and lightning

strikes. However, these rare stands may lack variation in sites and structure
that are critical to develop guiding principles for restoration and conservation management of the longleaf ecosystem. Based upon previous research, interactions with practicing land managers, and case studies, we
have developed a conceptual approach that involves the use of surrogates
that represent three major components of the ecosystem. In concert, these
surrogates can predict a diverse and ecologically functional ecosystem. All
are linked to critical structural and functional features of the system, as
well as previous land management histories. The presence of red-cockaded
woodpecker clusters implies the retention of maturing longleaf pine for
nesting cavities, as well as an open forest structure that is obtained with
frequent prescribed fire. Within its range, the presence of wiregrass (Aristida stricta or A. beyrichiana) suggests an absence of prior root disturbances that result in degraded floristic diversity. The presence of wiregrass
also reflects a history of frequent fire necessary for the plant community
to exist. The presence of an undisturbed ecotonal area between longleaf
pine-wiregrass uplands and depressional wetlands suggest that a high diversity of both plant and amphibians are likely to be present due to the
continuity of fire necessary to maintain this dynamic habitat. Larger, or less
fragmented woodland tracts often represent the more extensive environmental gradient in which longleaf once occurred and are sufficient in scale
to provide habitat for widely ranging wildlife species. Our premise is that
functional attributes of the system are more dependent on fire regimes,
canopy retention patterns and soil disturbance than the fact that a small
stand escaped the ax.
KIRWAN, MATTHEW L.* Duke University, Durham, NC,
mlk13@duke.edu. Dynamics of a Chesapeake estuarine forest and its
response to rising sea level.
To address the impact of rising sea level in a rapidly submerging Maryland
estuarine forest, 15 loblolly pines (Pinus taeda) were cored for dendroecological analysis. The study area is a pure stand of loblolly pine that
extends down an elevation gradient into surrounding marsh where dead
stumps and snags indicate a retreating forest margin. Although relative sea
level has risen considerably and there are dead trees at the forest-marsh
interface, there is no associated decline in ring width, making sea level
induced mortality unlikely. Instead, ring width is correlated positively with
annual precipitation and winter temperature and negatively with summer
temperatures. Although recruitment of new individuals was continuous between 1910 and 1930, there has been no more active recruitment except
for a small age class established immediately after regional drought. Because recruitment is failing in the present forest despite abundant seedlings
and an open canopy, recruitment ability appears to be limited by saturated
soils associated with periods of high sea level. The forest margin will
retreat stepwise, following storm induced mortality or continuously, following natural adult mortality. The position of the forest margin is then a
function of sea level position, but it represents the failure to recruit new
individuals, not the ability of adults to survive.
KITAJIMA, KAORU,1,4,* KIHACHIRO KIKUZAWA,2 DAVID ACKERLY,3 STEPHEN S. MULKEY1 and JOSEPH WRIGHT.4 1 University of
Florida, Gainesville, FL; 2 Kyoto University, Kyoto, Japan; 3 Stanford University, Stanford, CA; 4 Smithsonian Tropical Research Institute, Balboa,
Panama. Leaf-age effect on photosynthetic rates as a key parameter in
the theory of leaf longevity.
The cost-benefit theory of leaf longevity predicts that optimal leaf longevity
is shorter either when the initial photosynthetic rate is high or the rate of
its decline with leaf age is fast. The first prediction is well supported among
a wide range of plants species at both regional and global scales. In contrast, the second prediction, the link between the regression slope of photosynthetic capacity against leaf age, has been rarely shown. Here, we present strong support for both predictions from eight neotropical canopy tree
species ranging in mean leaf longevity from 29 to 400 days. Interestingly,
the parameter decreases with increasing leaf longevity more than predicted
by the theory. In other words, short-lived leaves die at the age their photosynthetic capacity reaches zero, while long-lived leaves senesce and abscise much earlier than the age when their photosynthetic capacity would
reach zero. We propose several possible explanations for this emergent

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property. Further, we demonstrate that the analysis of age-effects on photosynthesis opens a novel way to estimate daily photosynthetic gain from
instantaneous measurement of photosynthetic capacity.
KLAPER, REBECCA.* Science and Technology Policy Fellow, American
Association for the Advancement of Science, US EPA ORD/NCEA, Washington, DC. At the Crossroads of Genomics and Ecology: The Potential
for a Canary on a Chip.
The rapidly evolving field of genomics has provided new insight into not
only the study of human biology and medicine but ecology and environmental science. Genomics technologies measure the simultaneous changes
in the expression of hundreds of genes or proteins associated with complex
traits. In humans this has translated into new insights into learning and
memory, the development of disease, and the effects of toxins. Similarly,
in ecology and environmental science these technologies can provide unprecedented information as to biochemical changes associated with behaviors, the biochemical pathways affected by various toxins, the separate
effects of exposure to multiple stressors, and a direct comparison of responses of different species to environmental change. Eventually we may
be able to link health-related expression changes in other species, including
ecological indicator species, to changes in humans to provide a direct connection between human health and the health of other species in the same
environment. Like the canary in a coal mine, genomics of may provide a
canary-on-a-chip, or a technological tool to identify molecular changes that
could act as a warning before the significant decline of a population or
community of organisms. An ecologically-focused multi-species genome
project organized around ecological indicator species would provide the
basis for an early warning system for environmental exposure, responses
to anthropogenic changes and ecosystem decline that is more sensitive than
current measurements. This talk will provide a brief overview of the status
of the field, a description of the development of genomics technologies for
ecology and environmental assessment, and the current limitations with
regards to genomic data and funding in this field.
KLAUSMEIER, CHRISTOPHER A.* and ELENA LITCHMAN. Georgia
Institute of Technology, christopher.klausmeier@biology.gatech.edu, Atlanta, GA. Seasonal succesion in plankton communities.
Temperate lake plankton communities undergo succession of species over
the course of a season. There are broad regularities in the successional
trajectories between years and among lakes, but variation exists. Here we
use a periodically forced food web model to explore the dynamics of seasonal succession. A mathematical approximation that reduces the dynamics
to a series of jumps between equilibria facilitates the analysis. We find that
the model can accurately reproduce common successional patterns such as
the transition from a competitive phytoplankton species to a grazing resistant species after zooplankton become abundant, as well as predict novel
patterns. The model does not always produce identical dynamics from year
to year, but can exhibit regular and irregular multiyear cycles. More species
can coexist in the seasonal model than in the corresponding equilibrium
model, thus seasonal dynamics can explain some of the diversity of plankton communities.
KLEMOW, KENNETH M. Biology Department, Wilkes University,
Wilkes-Barre, PA, USA. Patterns of native and alien plant species in
two watersheds impacted by anthracite mining and urbanization.
Considerable interest has recently been focused on the role that human
impact plays on promoting the establishment and spread of invasive alien
species in ecosystems worldwide. To understand how mining and urbanization effect an increase in alien species, a detailed assessment of vegetation within 27 communities throughout two watersheds in northeastern
Pennsylvania was conducted in 2001. Patterns of native and alien species
were determined in each community to understand the degree to which
alien species benefit from disturbance, and whether some strata are more
impacted by aliens than others. Communities located on the mountain
slopes fringing each watershed have not been subject to human disturbance
since the early 1900s, and the plant communities there had low proportions
(,5% cover) of alien species. In contrast, communities impacted by urbanization showed much higher (.60% total cover) dominance by aliens.

Alien species were particularly well represented in herb and shrub layers,
much less so among saplings and trees. Communities recovering from surface mining were dominated by stress-tolerant species that were largely
native to Pennsylvania. The implications of these findings vis a vis efforts
to promote regional biodiversity of native species in human-impacted
ecoystems will be discussed.
KLIMAS, CHRISTIE,1,* MARK POTOSNAK,2 RAMESH MURTHY1 and
KEVIN GRIFFIN.3 1 Biosphere 2 Center, Columbia University, Oracle, AZ,
USA; 2 N.C.A.R., Boulder, CO, USA; 3 Columbia University, Lamont-Doherty Earth Observatory, Dept, of Earth and Environmental Sciences, Palisades, NY, USA. Light use efficiency in a non-nutrient limiting environment.
This experiment examined leaf level photosynthetic responses of Populus
deltoides trees grown in the Intensive Forestry Mesocosm at the Biosphere
2 Center, to light at different canopy positions under two CO2 treatments:
ambient (43 Pa) and elevated (120 Pa) CO2 conditions. A fully developed
shade (interior) and sun (exterior) leaf was chosen at three canopy levels
(high, middle, upper) for 5 trees in each CO2 treatment. PPF (photosynthetic photon flux) sensors were attached to branches adjacent to each leaf.
Light curves coupled with maximum photosynthesis (Asat) measurements
were done 3 times over a six-week experiment. After photosynthesis measurements, LMA and leaf nitrogen content were determined for the sample
leaves. Maximum rates of photosynthesis were significantly higher under
elevated CO2 conditions with a concurrent increase in daytime respiration.
Photosynthesis was significantly higher in middle and upper canopy and
exterior sunlit leaves under both ambient and elevated CO2 conditions.
Vertical position also had a significant effect on Rday with leaves in the
upper canopy having higher respiration values, and this effect was enhanced under elevated CO2. While elevated CO2 resulted in overall increased quantum efficiency (AQE) the effect of vertical position and shading on AQE showed no significant change between CO2 treatments. Midcanopy leaves showed a significantly higher AQE than upper and lower
canopy leaves in both CO2 treatments and this was mirrored by a lower
light compensation point. The leaves at the low canopy position under
elevated CO2 showed a significantly lower light compensation point than
the ambient low leaves. Nitrogen allocation also differed between the CO2
treatments with the ambient bay having higher percent leaf nitrogen content. These leaf level enhancements translated into greater accumulated
biomass in the elevated CO2 treatment. Elevated CO2 resulted in a significant improvement in light utilization irrespective of vertical profile and
incident light intensity.
KLINE, KERRY A.* and HORMOZ BASSIRIRAD. University of Illinois
at Chicago, Chicago, Illinois, USA. Maternal CO2 affects seed nitrogen
and seedling growth but not uptake and assimilation in wheat.
Seedling growth and vigor, two factors that impact seedling establishment
and plant community dynamics, are highly influenced by maternal as well
as current environmental conditions. We obtained seeds that were produced
under ambient and elevated CO2 from the Maricopa Free Air Carbon Dioxide Enrichment (FACE) site in Arizona and evaluated the CO2 effects
on the growth and physiology of the subsequent generation. We assessed
seed quality by determining carbon to nitrogen (C:N) ratio and the concentrations of calcium (Ca2+), potassium (K+), magnesium (Mg2+), ammonium (NH4+), nitrate (NO3-), phosphorus (P), and total nitrogen (N) in the
seeds. We also examined NO3- uptake capacity and whole-plant NO3- assimilation. Seed NO3- concentration was undetectable, but analysis of other
elements showed no significant changes in the concentrations of Ca2+, K+,
Mg2+, and P in response to elevated maternal CO2. Seed N and NH4+ concentrations were, however, lower in elevated compared to ambient produced
seeds. This lower seed N concentration may explain why eight-day old
seedlings produced from elevated maternal CO2 weighed significantly less
by at least 20% than ambient-produced seedlings regardless of the their
current CO2 conditions. Elevated maternal CO2 seedlings had a significantly
higher NO3- uptake rate than the ambient ones, but this effect was present
only if the growth CO2 was also elevated. We also examined whole-plant
NO3- reduction and found that elevated CO2 in either the maternal or current
environment had no affect on whole plant NO3- assimilation. The data
suggest that maternal CO2 does not alter NO3- uptake capacity or assimi-

lation, yet the early seedling growth is clearly stunted in offspring of high
CO2 plants. We believe that this maternal effect on the subsequent generation is mediated through an effect on total seed N concentration.
KLING, GEORGE W.,1,* KRISTI JUDD,1 MARTIN SOMMERKORN,2
KNUTE NADELHOFFER,2 EDWARD RASTETTER2 and LORETTA
JOHNSON.3 1 University of Michigan, Ann Arbor, MI; 2 Marine Biological
Laboratory, Woods Hole, MA; 3 Kansas State University, Manhattan, KS.
Transfers of carbon across the landscape: The controls on dissolved C
production in soils.
Much of the world9s plant production is returned to the soil within seasons
to decades. Labile fractions of this material are dissolved in soils and then
transported from uplands to lowlands and eventually deposited in the
oceans or exhaled to the atmosphere. Although this transported material is
important for regional to global carbon and nutrient budgets, our understanding of the integrated mechanisms responsible for this cycling, and our
ability to predict the sources and rates of production of dissolved materials
in soils, is rudimentary. We used pulse-labeling of 14C-CO2 added through
photosynthesis to intact plant-soil mesocosms and to field plots in order to
trace the movement of carbon through the plant-soil system of arctic tundra.
We found that the bulk of recent photosynthate lost from the plant is returned to the atmosphere through plant respiration in less than 10 days.
Recently-fixed plant carbon is also transferred immediately to roots, and
within 1-2 hours enters the dissolved pool in soils. The pool of dissolved
organic carbon (DOC) is most rapidly enriched with 14C from root exudation, followed closely by dissolved CO2. Dissolved CH4 labeled with 14C
is produced in the soil within hours of the label addition. Nutrient fertilization did not alter the rate of 14C transfer to the dissolved pool, but did
alter the magnitude of transfer for different C species. Dissolved organic
or inorganic carbon production dominated the dissolved soil pool, depending on the overlying vegetation type. Estimated transfer rates of C from
roots to DOC, CO2, and CH4 pools were high, and ranged from 5-25% per
day of the total pool size in the soils. We propose that this flux derived
from recently-fixed organic matter, versus from older organic matter, dominates the total production of dissolved material even in soils with high
organic matter content.
KLIRONOMOS, JOHN N.* University of Guelph, Guelph, ON, Canada.
Plant-microbe feedbacks in plant invasions. A synthesis of existing research and future prospects.
In this presentation, I describe a research study that focuses on the potential
for feedback responses between native and exotic plants and microbes to
determine plant invasiveness. The research study is currently being conducted in Southern Ontario, Canada using several native and exotic plants,
as well as several native and exotic mycorrhizal and parasitic fungi. Feedback interactions are being evaluated for all combinations of native and
exotic plants and fungi. Preliminary results indicate that the strongest negative feedbacks are exhibited when native plants are coupled with native
microbes, whereas the neutral and positive feedbacks are found when one
or both of the partners are exotic. Overall, rare plants display the strongest
negative feedback, whereas invasive plants typically have neutral feedback
responses. Feedback associated with fungal parasites is neutral to negative,
whereas in association with mycorrhizal fungi it is positive to negative,
depending on the source of the plants and fungi. Overall, these results
indicate that feedback responses between plants and microbes can contribute to plant invasiveness. These results will be discussed in the context of
other existing research, which should identify promising new directions for
research on plant invasion biology.
KLOEPPEL, BRIAN D.,1,* LISA MAZZARELLI,1 WAYNE T. SWANK,2
JAMES M. VOSE2 and THOMAS R. WENTWORTH.3 1 University of
Georgia, Coweeta Hydrologic Laboratory, Otto, NC, USA; 2 USDA Forest
Service, Coweeta Hydrologic Laboratory, Otto, NC, USA; 3 North Carolina
State University, Raleigh, NC, USA. Southern pine beetle impacts on
the ecosystem dynamics of a white pine watershed.
Southern pine beetle (Dendroctonus frontalis) most commonly affects loblolly, slash, and longleaf pine forests in the southern United States. However, the recent (1998 to 2002) drought conditions in the southern Appa-

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lachians including a 17% reduction in precipitation and a 34% reduction

in stream flow have predisposed white pine (Pinus strobus) trees to attack
by southern pine beetle. Several tree mortality pockets (. 2 ha each) in
the white pine watersheds (1 and 17) at Coweeta Hydrologic Laboratory
in North Carolina provide a great opportunity to study the southern pine
beetle and its effects on white pine watersheds. Therefore, the following
treatments were imposed. A treatment including cutting and removal of the
dead and infested trees was conducted in watershed 17 in October 2001.
The infested areas in watershed 1 remain untreated and the beetle- and
non-beetle impacted areas in watershed 1 were investigated further in this
study. We evaluated the changes in resource availability in beetle vs. nonbeetle infested areas including changes in light, soil water availability, and
soil and litter temperature. We found that in beetle-infested areas available
light increased 27 %, available soil water increased 13 %, mean soil temperature increased 0.9 C, but mean litter temperature was not significantly
different vs. non-beetle infested areas. Forest floor mass increased significantly (P , 0.05) after beetle-impact. In addition, after 15 months in the
field, white pine foliage decomposition exhibited a trend toward increased
decomposition in the beetle-impacted areas, but the difference was not
significant. The response to beetle-infestation was most dramatic in the
seedling and herbaceous layers. Seedling density increased two-fold and
seedling height was four times greater in beetle vs. non-beetle infested
plots. Herbaceous diversity was 30 % greater in beetle vs. non-beetle-infested areas. Overall, the likely long-term impact on white pine watersheds
will be a dramatic species change from nearly pure white pine forest stands
to mixed hardwood and white pine forests. This study also suggests some
of the ecosystem properties to measure to evaluate the impact of the exotic
insect, hemlock woolly adelgid, that is now impacting southern Appalachian hemlock trees.
KLUGER, EMILY C.* and RICHARD A. NIESENBAUM. Muhlenberg
College, Allentown, PA. The ecological impact of variation in host plant
quality on the geometrid herbivore, Epimecis hortaria.
The aim of this study has been to assess how the geometrid herbivore,
Epimecis hortaria, responds to different levels of plant defensive chemistry
and leaf nutritional quality that vary with light environment and leaf age
in populations of the host plant Lindera benzoin. Leaf chemical analysis
has revealed wide variation in plant volatile chemistry and leaf nitrogen
concentration in the host plant populations. Here we report on how E.
hortaria, the primary herbivore in these populations, responded to this variation. Field feeding trials showed that E. hortaria consuming leaves with
a greater proportion of volatile chemicals and lower nitrogen concentrations
actually gained more weight and consumed less plant material than those
feeding on leaves with less volatiles and more nitrogen. Those herbivores
that gained weight more rapidly also had faster developmental and pupation
rates. Dry weight measurements indicate that weight gain was independent
of water retention by caterpillars. Because individuals feeding on plants
with a greater proportion of volatile chemicals may be at greater risk of
attack by parasitoids, rapid development on this food source may be adaptive. Variation in rates of attack by parasitoid wasps, the interaction of host
plant leaf nitrogen content and volatile chemistry on feeding rates and
herbivore population biology, and the potential for nitrogenous plant defensive compounds and their detoxification by E. hortaria are considered.
KLUGH, KATRINA R.* and JONATHAN R. CUMMING. Department of
Biology, West Virginia University, Morgantown, WV. Variation in organic acid exudates among mycorrhizal species colonizing Liriodendron
tulipifera L. (yellow-poplar) in the presence of aluminum.
Acidic soils have phytotoxic levels of aluminum (Al) that have detrimental
effects on numerous plant physiological processes. Aluminum availability
is exacerbated by acidic deposition in the eastern United States. Liriodendron tulipifera (yellow-poplar) is an economically important tree species
in this region and is sensitive to Al. Root exudation of organic acids is one
mechanism of Al resistance in pants. Arbuscular mycorrhizal (AM) fungi
may also confer resistance, although the mechanisms involved are not
known. In the present study, the extent of Al resistance conferred by several
AM species was assessed and organic acid exudation was quantified to
determine if organic acid production was altered by AM fungi and Al.
Liriodendron tulipifera seedlings were inoculated with one of four AM

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fungal species (Glomus clarum, Acaulospora morrowiae, Glomus claroideum, or Paraglomus brasilianum) or a control inoculum and grown in
sand culture. Aluminum treatments of 0 and 50 mM Al were delivered in
solution. After eight weeks of treatment, biomass and root exudates were
measured. Organic acid profiles differed among fungi highlighting interspecific variation in carbon flux to the rhizosphere. Citrate, malate, and
oxalate concentrations were up to 6 times greater in the rhizospheres of
plants colonized by AM fungi compared to non-mycorrhizal plants. Fungi
conferring Al resistance maintained high organic acid concentrations under
exposure to Al. Rhizosphere citrate concentrations in plants colonized by
Al resistant G. clarum were not suppressed under Al exposure while plants
colonized by Al sensitive G. claroideum exhibited 50% reductions. Similarly, G. clarum colonized plants exhibited 30% reductions in malate rhizosphere production, while plants colonized by G. claroideum had 80%
reductions. Organic acid exudation differed among AM fungal species and
Al resistance of AM fungi appears to be associated with maintenance of
organic acid exudation.
KNAPP, LIZA B.,* JAMES H. FOWNES and ROBIN A. HARRINGTON.
University of Massachusetts - Amherst, Amherst, MA. Are invasive
woody plants released from ecophysiological constraints on photosynthetic capacity?
Invasive plant species may succeed in part due to release from tradeoffs
that constrain the performance of native species. Photosynthetic capacity
is generally constrained by interspecific tradeoffs between leaf N concentration and leaf mass per area (LMA). We compared interspecific and intraspecific trends in leaf physiology and morphology in four native and
four invasive woody species planted in open and understory plots. Leaf N
was positively correlated with mass-based, light-saturated net assimilation
(Asat), both among and within species. Leaf N was consistently higher in
invasives than in natives of similar life form and shade tolerance. We detected few significant differences in the Asatleaf N relationship between
invaders and their native counterparts. LMA and photosynthetic nitrogen
use efficiency (PNUE) were negatively correlated among species but positively correlated across treatments. Invaders did not consistently exhibit
lower LMA or higher PNUE than natives. The high photosynthetic capacity
of these invaders reflects high leaf N, rather than release from constraints
on PNUE.
KNEITEL, JAMIE.* Washington University, kneitel@biology2.wustl.edu,
St. Louis, MO. Resources and intermediate-consumer identity alters an
omnivore food web.
Omnivory has been increasingly incorporated into our understanding of
food webs and community structure. Models that address these food webs
make several predictions about what conditions facilitate coexistence between an omnivore and an intermediate consumer. These predictions include (1) the intermediate consumer must be a better competitor than the
omnivore, (2) the intermediate consumers can exclude the omnivore at low
resources, but will be excluded by the omnivore at high resources, and (3)
intermediate consumer abundance is lower in the presence of an omnivore.
Few empirical studies have taken into account the trait variation (competitive ability and predator tolerance) of intermediate consumers and how
this variation may alter community dynamics. I conducted a short-term
laboratory experiment using species that inhabit the modified leaves of the
pitcher plant, Sarracenia purpurea. These species included bacteria (basal
trophic level), five protozoa and a rotifer (intermediate consumers), and
larvae of the mosquito, Wyeomyia smithii (omnivore). Four replicates were
established for each of 32 communities which included 3 trophic levels, 2
resource levels, 5 intermediate consumers, and 2 densities of the omnivore.
Resources increased the growth rate and abundance of all species. The
intermediate consumers differentially affected both bacterial abundance and
Wyeomyia smithii growth. Three of the intermediate consumers significantly reduced bacterial abundance more than the omnivore. The presence
of two protozoan species resulted in an increase in omnivore growth rates.
The identity of intermediate consumers did not coincide with expected
outcomes: depending on the conditions, community dynamics converged
with different intermediate consumers or varied according to intermediate
consumer differences. The results of this study indicate the need to consider
trait variation (competitive ability and predator resistance) within a trophic

level because of their differential effects on the basal trophic level, the
omnivore, and the emergent community dynamics.
KNEPP, RACHEL G.,1,* JASON G. HAMILTON2 and EVAN H. DELUCIA.1 1 University of Illinois, Urbana, Il, 61801; 2 Ithaca College, Ithaca,
NY, 14850. Insect herbivory in a forest community exposed to free-air
CO2 enrichment.
Increased atmospheric CO2 concentration alters leaf structure and chemistry, but how these changes affect leaf herbivory in a complex multi-species
forest community is largely unknown. Leaf damage was quantified on hardwood seedlings planted in the understory of a loblolly pine plantation in
North Carolina. Seedlings were exposed to ambient (370 ppm) or elevated
(570 ppm) [CO2] with free-air CO2 enrichment (FACE). In 2001, insect
herbivores removed 51% less leaf area from seedlings growing under elevated compared to ambient [CO2] (f 5 9.33, P 5 0.0926). In 2002, a
drought year, there was consistently lower herbivory than in 2001, and there
was no significant effect of the treatment on the amount of tissue removed.
However, the number of leaves experiencing insect herbivory decreased
under elevated [CO2] (f 5 3.75, P 5 0.0245). The consistent reduction of
insect herbivory, at least in a wet year, suggests that elevated [CO2] may
change the population dynamics of affected insect species and their interactions with other plants and animals.
KNIGHT, TIFFANY M.,* ROBERT D. HOLT and MICHAEL BARFIELD. University of Florida, Gainesville, FL, USA. Can evolution by
natural selection rescue declining Trillium grandiflorum populations?
Rapid changes to the environment create novel selective pressures for species, which could cause them to decline towards extinction, unless they
rapidly evolve to a new phenotype. Although demographic analyses are
often used to project the rate of decline for many species of concern, these
analyses have not heretofore incorporated the potential for natural selection
to alter those projections. Here, we estimated the degree to which natural
selection on flowering time can rescue declining populations of the native
wildflower, Trillium grandiflorum, that are experiencing novel levels of
herbivory by white-tailed deer. We found that individuals flowering early
in the season were much more susceptible to deer herbivory than those
flowering later, and thus flowering time may be a trait that both would be
under strong selection and could alter the rates of decline of this species.
We employed two approaches to examine whether or not evolution of flowering time might rescue these declining populations from extinction. First,
we coupled demographic and selection analyses (elasticities and mean standardized selection gradients) to project change in the population growth
rate as mean flowering time changes. Second, we used individual-based
simulations to determine the probabilities of adaptation and extinction, assuming flowering time is a heritable, quantitative trait. We compared the
predictions of these two modeling approaches and relate these results to
natural Trillium grandiflorum populations at risk of extinction.
KNOPS, JOHANNES MH.1,* and SHAHID NAEEM.2 1 School of Biological Sciences, University of Nebraska, Lincoln, NE, US; 2 Zoology Department, University of Washington, Seattle, WA, US. The impact of elevated CO2, nitrogen and biodiversity on tissue quality and litter decomposition.
Increasing atmospheric CO2 concentrations, declining biodiversity and increased nitrogen deposition are changing productivity in many ecosystems.
These global changes also impact plant and litter quality and have the
potential to alter ecosystem nitrogen cycling. We examined nitrogen cycling
changes induced by changes in plant tissue quality and decomposition using
a three factorial experiment with elevated CO2 (ambient, 368 vs. elevated,
560 mol mol-1), nitrogen fertilization (unamended vs. deposition of 4 g N
m-2 yr-1), and species richness (1, 4, 9 or 16 species). We measured aboveground plant tissue and litter quality, e.g. carbon and nitrogen, soluble,
hemicellulose, cellulose and lignin fractions. In addition, we measured
aboveground litter decomposition over one and two year periods with insitu litterbags. Elevated CO2 decreased nitrogen concentrations by 9%.
Elevated nitrogen increased nitrogen concentrations by 7% while decreasing lignin concentrations by 16%. However, species richness had a much
larger impact than either CO2 or nitrogen fertilization, with higher species

richness decreasing tissue nitrogen concentrations by 18%, increasing cellulose by 20% and decreasing lignin concentrations by 16%. However, in
contrast we found only minimal differences in litter decomposition across
the treatments. Thus changes in tissue quality caused by elevated CO2,
nitrogen fertilization and changes in plant species richness do not result in
marked changes in litter decomposition rates or nitrogen released during
litter decomposition. The plant tissue quality response supports the hypothesis that plant communities produce different quality of biomass and
litter when resource availability or competition for nitrogen changes. However, these tissue quality changes have only a minor impact on litter decomposition and nitrogen cycling. Consequently, litter decomposition acts
as a negative feedback. Short term elevated CO2, nitrogen fertilization and
changes in species richness lead to changes in productivity and tissue quality. However, decomposition rates do not reflect these quality changes.
Thus, long-term the accumulation of limiting nutrients in litter will lead to
a negative feedback on productivity, diminishing the impact of changes in
biotic and abiotic changes on productivity.
KNOX, ROBERT G.,1,* FORREST G. HALL,2 KARL F. HUEMMRICH2
and JANNETTE C. GERVIN.1 1 NASAs Goddard Space Flight Center,
Greenbelt, MD; 2 University of Maryland, Baltimore, MD. Estimating terrestrial carbon exchange from space: How often and how well?
Data from a new space mission measuring integrated light-use efficiency
could provide a breakthrough in understanding global carbon, water, and
energy dynamics, and greatly improve the accuracy of model predictions
for terrestrial carbon cycles and climate. Over the past decade, Gamon and
others have shown that changes in photo-protective pigments are sensitive
indicators of declines in light-use efficiency of plants and plant canopies.
The requirements for integrated diurnal measurements from space need to
be defined, before a space mission can be formulated successfully using
this concept. We used tower-based CO2 flux data as idealized proxies for
remote measurements, examining their sampling properties. Thousands of
half-hourly CO2 flux measurements are needed before their average begins
to converge on an average annual net CO2 exchange. Estimates of daily
integrated fluxes (i.e., diurnal curves) are more statistically efficient, especially if the spacing between measured days is quasi-regular, rather than
random. Using a few measurements per day one can distinguish among
days with different net CO2 exchanges. Fluxes sampled between mid-morning to mid-afternoon are more diagnostic than early morning or late afternoon measurements. Similar results (correlation . 0.935) were obtained
using 2 measurements per day with high accuracy (65%), 3 measurements
per day with medium accuracy (610%), or 5 measurements per day at
lower accuracy (620%). An observatory in a geosynchronous or neargeosynchronous orbit could provide appropriate observations, as could a
multi-satellite constellation in polar orbits, but there is a potential trade-off
between the required number of observations per day and quality of each
observation.
KOCHSIEK, AMY E.* and JOHANNES M. KNOPS. University of Nebraska-Lincoln, Lincoln, Nebraska. Litter quality and decomposition in
irrigated versus dryland corn agroecosystems.
Agroecosystems within the Midwestern region of the US have lost a substantial amount of carbon over the last century. New and improved management schemes, such as no tillage, have the potential to increase carbon
pools and lead to a net carbon sequestration. One important change has
been the switch from dryland to irrigated systems, which has generally
been assumed to have a negative net impact on carbon sequestration. However, dryland corn systems are fertilized at the beginning of the growing
season with an adequate amount of nitrogen for the entire season assuming
average rainfall. In years with below average rainfall, this practice results
in over-fertilization. Therefore, there is the potential to better manage nitrogen additions by using irrigation. We examined the impact of irrigation
on tissue quality and litter decomposition for intensively managed corn
systems in eastern Nebraska during 2001, a year with below average rainfall. We found that corn litter had significantly higher percent nitrogen
present in the litter in dryland as compared to the irrigated corn. The magnitude of this increased varied from 10% to 100% higher depending on the
litter category, with the largest changes in stalks, 0.37% in irrigated versus
0.78% in dryland and corn cobs, 0.75% in irrigated versus 1.48% in dry-

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187

land. This corresponded with higher crop nitrogen use efficiency in irrigated corn fields. In addition, other studies that have shown that corn plants
accumulate nitrogen late in the growing season in stalks and cobs, before
the nitrogen is translocated into the developing corn seed. Dry periods
during the time of corn seed development lead to lower crop production
and incomplete translocation of this nitrogen into the developing corn
seeds. All litter categories decomposed 10% to 20% faster in the dryland
field, matching the higher tissue quality that we found in the initial senesced
plant categories. Thus, the ability to more precisely manage nitrogen addition can, in addition to increasing crop nitrogen use efficiency, also decrease decomposition rates. Thus, crop irrigation has the potential to lead
to higher rates of carbon accumulation in irrigated corn agroecosystems,
due to better nitrogen fertilizer management.
KOECHY, MARTIN* and KATJA TIELBOERGER. Potsdam University,
Potsdam, Brandenburg, Germany. Rainfall distribution and persistence
of populations of semi-desert annuals.
We used a spatially-explicit individual-based model to simulate the effect
of mean annual precipitation level (maP; low: 75 mm, medium: 90 mm,
high: 105 mm) and the shape of interannual distribution of precipitation
(more normal vs. right-skewed) on the performance of annual plant populations. Our model simulates soil moisture, seed germination, and plant
growth calibrated to the desert annual Filago desertorum (Asteraceae), assuming a homogenous environment located in the northern Negev desert
(Sede Boqer, Israel). We used two approaches for producing rain scenarios.
(1) We increased or decreased daily rain values of Sede Boqer to change
the shape of the maP distribution without changing the intraannual rain
pattern. (2) We reproduced the same maP distributions (shape and level)
by selecting years of data from several Israeli climate stations with different
intraannual rain patterns. We compared population performances for 20
simulations of 100 years for each scenario with an index based on the
percentage of years when seed production was greater than the 75-percentile under unmanipulated 1961-1990 Sede Boqer precipitation and subtracted the percentage of years when seed production was less than the 25percentile. Factorial ANOVA showed that performance differed significantly among approaches, maP levels, and distribution shape. Performance
always increased with maP (low: -0.31, medium: -0.07, high: 0.14). Performance at high maP was significantly higher under normal than under
skewed distribution for both approaches. This was reversed when maP was
low, but the difference was significant only for approach 1. Performance
at medium maP was significantly higher under normal distribution for approach 1 but did not differ between distributions under approach 2. The
results show that, subject to model assumptions, population performance
varies not only with maP but also with the shape of annual precipitation
distribution and intraannual variation of precipitation.
KOELLE, KATIA V.* and MERCEDES PASCUAL. University of Michigan-Ann Arbor, Ann Arbor, MI. Disentangling the roles of extrinsic and
intrinsic factors in nonlinear disease dynamics.
Population cycles are generally generated by the nonlinear interplay of
intrinsic feedback mechanisms and extrinsic environmental drivers, yet attempts to disentangle these two components from ecological time series
are rare. Exceptions include population models that incorporate seasonality.
We present a semi-parametric time series method that enables the separation of these factors and illustrate the approach with an application to disease dynamics. From a time series of number of cases, the method reconstructs the decay and duration of immunity. It also allows for temporal
variability in one of the parameters, the transmission coefficient, as the
result of environmental forcing. This forcing includes seasonal, interannual,
and long-term components. An application to cholera dynamics in Bangladesh identifies the roles of both temporary immunity and variable transmissibility. The seasonal transmissibility component re-establishes the
known annual pattern of the disease, while the interannual component
shows a correlation with ENSO events. The waning of immunity is straindependent. Disease patterns in recent decades show the interplay of straindependent immunity with climatic forcing. We propose the existence of
periods that are refractory to climatic events as the result of patterns of
immunity. Similar nonlinear time series models should be applicable to
other population studies to identify the respective roles of extrinsic and
intrinsic factors.

188

Abstracts

KOERNER, BRENDA A.* and JEFFREY M. KLOPATEK. Arizona State

University, Tempe, Arizona. Soil responses to water and nitrogen additions along an urban to rural gradient.
Over 50% of the world population lives in urban environments, and the
proportion of the world population in urban areas is expected to increase.
Urban cities present dynamic mesocosms in which to study the effect humans and land-use changes have on urban ecosystem processes. Urbanization can affect temperatures through the heat island affect, precipitation
through convection of heat, and nitrogen availability through increased nitrogen deposition. This has led to the question: how do urbanization factors
affect the movement of carbon through arid ecosystems? To address this
question, we are evaluating the effects of urbanization on the creosote bush
community along an urban to rural gradient. Water and nitrogen additions
were added to plots during the summer monsoons of 2001 and 2002, and
during the winter of 2001. Soil CO2 efflux was measured the day of treatment (prior to treatment), day after treatment and 7 days following the
treatment using an Infrared Gas Analyzer. Nitrate and ammonium availability was also determined concurrently with soil CO2 efflux measurements. During summer additions, soil CO2 efflux was significantly higher
in plots that received water and water-nitrogen additions compared to plots
receiving nitrogen or no treatment. During winter treatments, nearly no
response to any treatment was seen. Water treatment responses were also
twice as high under the canopy compared to interspaces. Additionally, the
soil CO2 efflux was higher at the urban site than at the suburban or rural
sites. Nitrogen availability may also help explain differences in soil CO2
efflux between sites.
KOIKE, FUMITO* and KOICHI MIKAMI. Yokohama National University, Yokohama, Kanagawa, Japan. Prediction of alien tree invasion to
various climax vegetations in a subtropical oceanic island.
Bischofia javanica (evergreen tree, Euphorbiaceae, Native in Ryukyu, south
China, Malaysia, tropical Australia) and Morus australis (deciduous tree,
Moraceae, Native in Ryukyu, south China) are invasive trees in the Pacific
Bonin Islands, where 71% of woody species are endemic. B. javanica is
already the dominant canopy tree in lower mesic forests, however, its ability
to invade into other habitats has not yet clear. In our previous research,
maximum tree height was the most important trait determining invasion
probability and final dominance in climax forest (Koike 2001). Maximum
height is quite sensitive to ground topography. We measured maximum
height of native and invasive species on various sites in Hahajima Island.
The island is the highest (460 m) and widest island in the Bonin Islands.
A statistical model to predict maximum height from topographic environment was obtained from various species. Solar radiation (dependent on
slope direction and shade by ridge), catchment area, direction of surrounding ridges intercepting wind, ground steepness, and altitude (temperature,
air humidity, etc.) were environmental variables. These values were calculated by a free GIS using 10 m mesh DEM. When all species were
combined, altitude showed negative and catchment area showed positive
effects on maximum tree height. B. javanica showed similar results although less sensitive to altitude. Map of species maximum height was
produced and invasive species were compared with those of native species.
Maximum height of B. javanica exceeded native species in valleys (current
dominant site) and even just below the summit. Shrubs with many endemic
plants are currently spreading under mesic environment at just below the
summit. In our prediction, B. javanica will dominate at this habitat in the
future, changing phisiognomy from shrub vegetation to forest, and probably
causing extinction of endemic shrub species.
KOLAR, CINDY S. U. S. Geological Survey, La Crosse, WI. Invasions
by freshwater non-indigenous species: interactions with other global
changes.
Study after study has shown the establishment of nonindigenous species
(NIS) and habitat degradation to be the most important anthropogenic impacts on freshwater ecosystems. Even though few NIS cause detectable
changes in the ecosystems they invade, large numbers of introductions
ensure that the global effects of NIS on native freshwater biodiversity and
ecosystem function would be large, even in the absence of other global
changes. In reality, however, the continued introduction and spread of NIS

in freshwaters will be accompanied by a multitude of other global changes.

In this presentation, I will summarize the nature of several major global
changes now underway with respect to freshwaters: globalization of commerce (commercial shipping, aquaculture, the aquarium and pond trade,
and bait trade), waterway engineering (building and removing of canals
and dams), land-use changes (siltation, eutrophication, and water withdrawl), climatic and atmospheric changes, and intentional stocking. It is
expected that globalization of commerce, waterway engineering, and the
intentional stocking of NIS will interact more strongly with NIS than other
global changes to increase the problem of NIS in the future. Given the
unpredictability and devastation that can ensue from biological invasions,
increased measures should be taken worldwide to reduce the spread of
aquatic NIS from these higher risk, and largely, more easily controllable
pathways. Predicting the identity, spread, or impact of potentially invading
species is a growing subdiscipline within invasion biology; researchers
should be encouraged to continue to develop quantitative, transparent, and
defendable models to predict aspects of biological invasions. Such tools
could be used not only to better understand invasion ecology, but could
also be the basis for developing best management practices or regulations
in the future to reduce the spread and impact of freshwater NIS.
KOLASA, JUREK. McMaster University, Hamilton, Ontario, Canada.
Complexity, system integration, and susceptibility to change.
Much of the research into complexity has examined the number of the
components, their interaction strength, or specific dynamic aspects such as
attractors or evolution. At the same time, it has been noticed that ensemble
properties may be largely unaffected by variations in the properties and
behavior of elements. This observation suggests that, as a system organizes;
a general property emerges which renders specifics of complexity (and
complicatedness) to decline in importance. I hypothesize that this is due to
system integration. Using a simple quantitative model, I examine the effect
of integration on the resistance of ecological systems to perturbation under
different permutations of vertical (complex) and horizontal (complicatedness) system structure. Preliminary analyses indicate that more integrated
systems are more robust to change of lower level components (but within
narrower environmental boundaries) than less integrated systems of the
same complexity. Furthermore, complexity per se appears to be less important than the specifics of the structure and the integration of the components. Scale, however, is important: behavior and attributes of a system
depend to a considerable degree on the level at which integration of subunits is expressed most. The findings seem to be in general agreement with
conclusions drawn from small world networks, perturbation studies, model
food webs, and many others. These observations imply the need for systematically different approaches to management and sustainability of ecosystems that differ in complexity, integration, and boundedness (e.g., young
vs old lake communities, or tropical rain vs boreal forests). The argument
is also made that sustainability does not depend on integration or complexity directly but rather on the completeness of the system structure included in the management. Integration and complexity, however, inform
the adaptive process of identifying the system structure and the best modes
to sustain it.
KOMAI, KATSUAKI and TADASHI HIBINO. Depertment of Social and
Environmental Engineering, Graduate School of Engineering, Hiroshima
University, Higashi-Hiroshima, Hiroshima, Japan. Resuspension and
physical properties of natural cohesive sediment in estuaries.
In semi-enclosed eutrophied estuaries, a fluid mud layer is observed above
the seabed, especially from summer to autumn. Since nitrogen and phosphorus absorb on the sediment particles of fluid mud in high concentrations,
the mechanisms of resuspension of cohesive sediment and fluid mud movement with disturbance by wave and tidal current are important to coastal
environments. Because the resuspension of cohesive sediment depends on
physical and chemical properties of sediments and bed shear stress by disturbance of wave and tidal currents, the generalization of the resuspension
mechanism is difficult determine. The purpose of this study is to clarify
the physical properties (water content, wet density and vane shear strength)
of natural cohesive sediment and the actual resuspension state with longterm observation in semi-enclosed estuary and hydraulic model experiments. Natural cohesive mud layers and fluid mud layers were sampled

with a bottom mud sampler without disturbing the sediment layers in Hiroshima Bay. The vertical profile of vane shear strength was measured. At
the observation point, time series of current speed and turbidity above the
seabed were observed continuously for one month when the fluid mud layer
was generated. The actual resuspension state under natural disturbance by
wave and current was confirmed. The resuspension of natural cohesive
sediment by wave or currents was investigated in a wave channel and a
circular channel with sampled sediments and salt water. In the experiment,
the physical properties of the sediment and bed shear stress by water flow
were confirmed, and suspended sediment concentration was measured.
From integration of suspended sediment concentration, the amount of resuspension was estimated.
KOO, KYUNG-AH,1,* WOO-SEOK KONG2 and CHONG-KYU KIM.3
University of Georgia, Athens, Georgia, USA; 2 KyungHee University,
Seoul, Korea, Korea; 3 KyungHee University, Seoul, Korea, Korea. Distribution of evergreen broad-leaved plants (trees and shrubs) and climatic factors in the Korean Peninsula.
1

The Korean Peninsula contains 132 species of evergreen broad-leaved trees

and shrubs(EBTS) which belong to 61 genera. The distributional patterns
of EBTS were categorized into seven groups on the basis of the number
of distributing sites, distributional altitudes and latitudes. Out of seven
groups, the cold-tolerant EBTS were common at groups 1and 2, along with
Empetrum nigrum var. japonicum, Diapensia lapponica subsp. obovata of
group 3. However, the warmth-tolerant EBTS were rich at groups 3, 4, 5
and 6. The northern distributional limit of warmth-tolerant EBTS includes
Is. Baeknyeong, Is. Daechung, Mt. Cheonma, Mt, Samak, Is. Ulreung, Is.
Dok, etc. The southern distributional limit of cold-tolerant EBTS includes
Mt. Jiri, Mt. Chiak, Is Ulreung, Is Cheju, etc. The lower distributional limits
of cold-tolerant EBTS in the groups 1 and 2 decreased as one moves toward
south. The upper distributional limit of warmth-tolerant EBTS in the groups
3, 4 and 5 decreased with increasing latitude. However, no clear distributional tendency is noticed in the groups 6 and 7. The warmth-tolerant EBTS
appear to show close relationship with the January mean temperature (48C) and January mean minimum temperature (-98C) than others. On the
other hand, the cold-tolerant EBTS seem to respond well to the August
mean temperature (198C) and August mean maximum temperature (268C)
than others.
KOPTUR, SUZANNE,1 JAMES R. SNYDER,2 MICHAEL S. ROSS,3
HONG LIU1 and CHRISTOPHER BORG.3 1 Department of Biological Sciences, Florida International University, Miami, FL, USA; 2 USGS Biological Resources Division, Big Cypress National Preserve, Ochopee, FL;
3
Southeast Environmental Research Center, Florida International University, Miami, FL, 33199. Effects of Key Deer herbivory after fire in pine
rockland understory.
We conducted a two-year experiment to detect and measure effects of Key
Deer grazing after fire (as part of a larger study investigating fire effects
on vegetation in Lower Keys pine rockland). Pine rockland is a fire-dependent successional habitat, with the diverse understory flora giving way
to hardwoods if the habitat is protected from fire for many years. Fire kills
all aboveground plant parts, stimulating resprouting in most species and
seed germination and establishment in some. New growth is more palatable
than surrounding vegetation, and Key Deer eat a variety of plants. We
hypothesized that selective herbivory by the deer has the potential to shape
the plant community. We built small exclosures (4 ft high field fence)
around circular 1 m2 plots in a blocked array in summer burn, winter burn,
and control units (exclosures were put up within several weeks of the
burns). Deer herbivory in open (non-exclosure) plots was substantially
greater in burned than control units, so differences between exclosure and
open plots were most dramatic in the burned units. Many plant species
attain greater heights inside exclosures, especially in burned units. There
was a significant interaction between exclosure treatment and season of
burn. Some herbaceous species were able to flower prolifically after fire
inside exclosures, whereas they were less abundant and flowered less when
deer had access to them. However, we detected no significant differences
in overall plant density with either exclosure treatment or season of burn.
Cover was estimated with a quantitative scale, and no significant differences were detected with exclosure treatment; however, season of burn did

Abstracts

189

affect cover of plants in the unit one year post-burn. Herbaceous plant
species richness was greatest inside exclosures in burned units, followed
by open plots in burned units, and lowest in unburned units (no significant
difference between exclosures and open plots). Normal interactions may be
exaggerated since Key Deer numbers are at an all-time high and much of
the pine rockland in the Key Deer Refuge has been long unburned, but
there is evidence from our study that selective herbivory after fire affects
plant morphology, reproduction, and species richness of the plant community.
KOSKI, MARCI L.* and BRETT M. JOHNSON. Colorado State University, Department of Fishery and Wildlife Biology, Fort Collins, CO, USA.
Interactive effects of climate change and eutrophication on growth of
a coldwater planktivore.
Anthropogenic environmental change will have direct and indirect effects
on aquatic ecosystems, including alteration of food web dynamics and trophic status of lakes and reservoirs. Climate change and eutrophication may
affect lentic characteristics such as water quality and clarity, primary and
secondary productivity, and zooplankton distributions that, in turn, can impact higher trophic levels. In this analysis we used an improved bioenergetics model fitted with a light-dependent functional response to predict
growth of a planktivorous fish (kokanee salmon, Oncorhynchus nerka) in
Blue Mesa Reservoir (BMR), CO. Inputs to the model included geographical location, kokanee foraging depth (diel vertical migration scenarios),
time of day, and limnological data such as secchi depth, zooplankton density, and temperature profiles. We first simulated yearling kokanee growth
using information collected from BMR in 2002 to predict growth under
current conditions. We simulated climate warming scenarios by increasing
epilimnetic temperatures by 2 and 4 degrees C. To simulate eutrophication
scenarios, we decreased water clarity and increased zooplankton density in
the epilimnion. We also created interaction scenarios by combining the
effects of both eutrophication and climate warming. We found that increasing epilimnetic water temperatures by 2 and 4 degrees C decreased kokanee
daily growth rate by 12% and 27% respectively; conversely, our eutrophication scenarios increased kokanee growth rate by approximately 30% due
to increased food availability. When simulated eutrophication conditions
were combined with warmer epilimnetic temperatures the benefits of higher
food availability were limited by the increased energetic costs of inhabiting
warmer water. We conclude that the interactive effects of climate change
and eutrophication may force kokanee to feed for longer periods of time
in the epilimnion, where temperatures are physiologically sub-optimal and
predation risk may be greater.
KOSLOW, JENNIFER M.,* JENNIFER A. RUDGERS and KEITH CLAY.
Indiana University, Bloomington, IN, USA. Endophytic fungi alter relationships between diversity and ecosystem function.
Recent studies have expanded the complexity of research on biodiversity
by investigating whether relationships between diversity and ecosystem
function hinge on the presence of widespread symbionts. Cool-season
grasses commonly harbor systemic, endosymbiotic fungi that enhance plant
resistance to herbivory, drought, and competition. We address whether
these endosymbionts modify associations between diversity, productivity
and invasibility. During a three year field experiment, we manipulated symbiosis in tall fescue grass (Festuca arundinacea). Plots lacking endophytes
exhibited a greater decline in primary productivity with increased plant
diversity than did plots with symbionts. In the greenhouse, we altered both
symbiosis in tall fescue and plant species richness. Invading, symbiotic
fescue achieved higher biomass than uninfected plants regardless of richness, but experienced a greater reduction in biomass as realized (harvest)
diversity increased. As many grasses harbor fungal endophytes, these clandestine symbionts may contribute more to the structure and function of
terrestrial communities than previously supposed.
KOTA, NATHAN L.,* RICK E. LANDENBERGER and JAMES B. MCGRAW. Department of Biology, West Virginia University, P.O. Box 6057,
Morgantown, WV, USA. Dispersal of Ailanthus altissima seeds into
varying environments: Is any area safe from invasion?
Ailanthus altissima (Tree of Heaven) is an invasive tree species from central China. Initially confined to highly disturbed urban areas, Ailanthus has

190

Abstracts

recently been found in relatively natural, undisturbed habitats. Female trees

can produce up to 300,000 wind-dispersed samaras, which may travel long
distances from focus populations. We examined seed dispersal by trapping
seeds in open fields, intact forests, and a recently harvested forest to characterize the dispersal kernel in these common, rural environments. Seeds
were found out to 100 m from isolated focus populations at all sites, and
seed densities showed strong declines with increasing distance; one intact
forest site, however, showed no relationship over distance and had the same
average density across the entire 100 m sampling interval. Examining seed
weight as a function of site and distance from focus populations, we found
that mean weight varied among all sites (F569.12, p,0.0001) but showed
no consistent statistical relationship with dispersal distance (F50.5365,
p50.46). Heavy seeds traveled just as far, on average, as light seeds; however, the effect of distance depended on site (F56.71, p,0.0001), with one
intact forest site showing an increase in mean seed weight at greater distances (R250.12 p50.01), and a harvested forest site showing the opposite
effect (R250.05, p50.02). Ailanthus focus populations vary in their dispersal properties in significant ways, but our results show clearly that longdistance dispersal is frequent in both open and forested environments. This
pattern makes control of this noxious invasive plant challenging. Nevertheless, control may become increasingly important with expected increases
in timbering over the next several decades.
KOVACIC, DAVID A.,1 ANTHONY J. KRZYSIK,2 MICHAEL P. WALLACE,1 JEFFREY J. DUDA,3 D. CARL FREEMAN,4 JOHN H. GRAHAM,5 JOHN C. ZAK6 and HAROLD E. BALBACH.7 1 University of
Illinois, Champaign, IL; 2 Prescott College, Prescott, AZ; 3 U.S. Geological
Survey - Biological Resources Division, Western Fisheries Research Laboratory, Seattle, WA; 4 Wayne State University, Detroit, MI; 5 Berry College, Mount Berry, GA; 6 Texas Tech University, Lubbock, TX; 7 U.S.
Army Construction Engineering Research Laboratory, Champaign, IL. Soil
mineralization potential as an indicator of ecological disturbance.
Military reservations contain some of the largest unbroken landscapes remaining in the United States. Their vast area and diverse vegetative cover,
provide refuge for many rare plant and animal species. These same characteristics also make these lands ideal locations for military training. The
combined fates of plants, animals, and continuing military training rely on
sustainable ecosystem management. The development of a set of readily
measured ecological indicators would help managers determine the integrity of various ecosystems on military lands. Early identification of at-risk
habitats could prevent their loss or further degradation. Our objective was
to determine if soil N dynamics could be used as an indicator of ecosystem
integrity. In mature N limited forests, competition for ammonium released
through mineralization is intense and soil inorganic N is low. Under such
conditions populations of nitrifying bacteria are also low. Following disturbance and interrupted plant uptake, more N may be found in the soil.
Populations of nitrifying bacteria increase under such conditions, and nitrate may become the dominant form of inorganic N present. We studied
9 forest sites at Ft. Benning, GA, representing a gradient of military training disturbance. We hypothesized that following incubation low disturbance
soils would produce the highest levels of ammonium, while high disturbance soils would produce low levels of ammonium and the highest levels
of nitrate. After a 4-week incubation, soil N exhibited significant trends
across the three disturbance levels. Average mineralized soil ammoniumN was 1073, 728, and 240 mg m-2 in the low, moderate and high training
intensity soils, respectively. Average soil nitrate-N formed was 277, 597,
and 1503 mg m-2 in the low, moderate, and high training intensity soils,
respectively. Soil mineralizable N may be an effective ecological indicator
for determining ecosystem condition on military training lands.
KRAMER, KAREN A.* and ANDREW NORTON. Colorado State University, Ft. Collins, Colorado. Integrating intensive grazing and biological control for the management of Dalmatian toadflax.
Intensive grazing, as well as arthropod biological controls have been applied separately against an array of weed pests infesting rangelands in the
western United States. Although these management alternatives have provided documented benefits, we have little information on the compatibility
of these two techniques. To gain better understanding in this area, the
interactions between intensive goat grazing and the biological control agent

Brachypterolus pulicarius on Dalmatian toadflax infested rangeland in Colorado were examined. Both the impact of goats and the biological control,
on weed performance and population dynamics were examined, as well as
the impact of intensive grazing on biological control agent population dynamics and performance. Research plots were situated in short grass prairie
at an elevation of ca. 6,500 feet at the Nature Conservancy Phantom Canyon preserve north of Fort Collins, CO. Our results clearly indicated that
goats will feed on Dalmatian toadflax, and that goat grazing has the potential to interact with at least one of the biological controls of this weed.
Goat grazing altered the morphology of this plant, reduced the number of
fruits produced, and may alter the flowering phenology as well. In addition,
grazing appears to have altered the relationship between flower density and
beetle density in our plots.
KRAUS, JOHANNA M.* University of Virginia, Charlottesville, VA.
Changes in subsidized versus intrinsic resource availability at a pondterrestrial edge.
The movements of nutrients and energy among habitats, also known as
subsidies, can influence the distribution and abundance of consumers. For
example, web-building spiders aggregate at the edge of bodies of water to
catch emerging aquatic insects. The increased abundance of consumers in
subsidized versus unsubsidized populations has been shown to suppress
intrinsic prey density at terrestrial interfaces with streams, rivers and marine
systems. The role of subsidies in the communities near ponds, however, is
virtually unknown. This study examines changes in density of a common
and abundant predator at the pond edge, the wolf spider (Aranea: Lycosidae), with changes in terrestrial prey and aquatic subsidies over time and
space. I measured patterns of aquatic insect emergence, ground arthropod
activity and density, and insect herbivore abundance at two fishless ponds
located in the southern Appalachians. I trapped once a week at 0, 1 and 2m from the pond edges, from June through mid-August 2002, along six
transects radiating from each pond. Emergence rates differed over the summer and between the two ponds. Over 90% of the emerging insects were
flies (Diptera). Wolf spiders may obtain a large proportion of their diet
from these flies when they emerge in high numbers. In future work, I will
examine this trophic link experimentally using population manipulations
within mesocosms and stable isotopes as a tracer of diet. The results of
these studies will identify the role of subsidies in shaping arthropod community structure temporally and spatially at a pond-terrestrial edge.
KRAUSE, ANN,1,* KENNETH FRANK,1,2 DORAN MASON,3 ROBERT
ULANOWICZ4 and WILLIAM TAYLOR.1 1 Michigan State University,
Department of Fisheries and Wildlife, 13 Natural Resources Blg., East Lansing, MI; 2 Michigan State University, Department of Counseling, Educational Psychology, and Special Education, East Lansing, MI; 3 NOAAGreat Lakes Environmental Research Laboratory, Ann Arbor, MI; 4 University of Maryland, Chesapeake Biological Laboratory, Solomons, MD.
Intuitive compartments: The other half of trophic structure in food
webs.
Food webs are representations of the feeding interactions among organisms
in a community. When examining the structure or organization of these
interactions, most food web studies focus on trophic levels, where measures
quantify the number of energy transfers, and organisms are grouped based
on the similarity of their interactions. In contrast, compartments are underutilized and are defined as groups of organisms where interactions within the group are stronger than interactions between groups. In this study,
we applied a method from the social networking science to identify compartments within an established food web of Chesapeake Bay, USA. This
method has distinct advantages over previous methods in that organisms
are assigned to only one compartment, the number of compartments is not
identified a priori, and the results are tested for significance. Using this
method on different weightings of interactions of the same Chesapeake Bay
food web, we found that the results of some of the weightings were greater
than expected by chance (a 5 0.05). The connectance, or realized density
of interactions relative to the potential density of interactions within compartments was higher than the overall connectance, and the connectance
between compartments was lower than either within or overall. Depending
on the weighting of the interactions, the analysis identified two to six compartments within the web. Compartments contained organisms across sev-

eral different trophic levels and represented groups of organisms found in

similar habitats, such as the pelagia or benthos. Using a multidimensional
scaling technique based on the results, we arranged compartments and their
associated organisms in relation to each other. This graphical representation
gave us an intuitive understanding of the compartmental structure of the
food web. Identifying and understanding compartmentalization in food
webs is necessary to ecological theories such as the relationship between
stability and diversity in real food webs.
KRENZ, CHRISTOPHER,* MAXINE L. CHANEY and WAYNE R.
WOOD. Oregon State University, Corvallis, OR. Does spatial variation
in barnacle and mussel recruitment affect rocky intertidal community
dynamics?
Variation in the input of new individuals into populations is predicted to
have important effects on the interactions that occur between species. For
most marine animals the input of new individuals, termed recruitment, occurs when planktonic larvae are delivered to a population by ocean currents, and they metamorphose into the adult form. Recruitment of barnacles
and mussels to the rocky intertidal zone varies dramatically along a 600km
section of the U.S. west coast. Barnacle and mussel recruitment is one to
two orders of magnitude higher at sites along the Oregon coast than at
sites along the northern California coast. We conducted a combination of
experimental and observational studies at two high recruitment sites in
Oregon and two low recruitment sites in California to test the prediction
that the strengths of interspecific species interactions are correlated with
rate of recruitment. At the high barnacle and mussel recruitment sites in
Oregon, new recruits quickly filled available space causing intense competition for space. Predation by snails on mussels decreased the intensity
of competition between mussels and barnacles. In contrast, there was little
competition for space at the low barnacle and mussel recruitment sites in
California. Predation by snails on mussels in California had no effect on
the outcome of competition between mussels and barnacles, because of the
lack of competition for space. Overall, the strength of interspecific interactions was positively correlated with recruitment regime. This is strong
empirical evidence in support of the hypothesis that recruitment rates have
important effects on community dynamics.
KREPS, TIMOTHY A.* and DAVID M. LODGE. University of Notre
Dame, Notre Dame, IN. Impacts of the invasive crayfish Orconectes
rusticus on lake energy pathways.
Invasive species are one of the most important drivers of change in freshwater communities and ecosystems. Invasive crayfish can have particularly
strong impacts due to their high biomass and ability to impact numerous
trophic levels through omnivory. In north temperate lakes, the introduction
of the rusty crayfish, Orconectes rusticus, results in a reduction in the
diversity and abundance of littoral zone macrophytes and macroinvertebrates. However, very little is known about impacts on higher trophic levels. We predict that due to reductions in invertebrate and small littoral fish
abundance, fish that are able to prey on crayfish must depend much more
heavily on crayfish and thus, the littoral zone for food. Here we examine
resource use by fish, which are good integrators of food webs. Using carbon
stable isotopes (d13C), we compared the relative dependence of fish on
littoral and pelagic production in lakes with (n 5 7) and without (n 5 6)
O. rusticus. Littoral vs. pelagic primary consumers were collected to provide average d13C values for energy sources derived from these habitats.
d13C values from muscle samples from several common fish species were
compared to the littoral and pelagic d13C endpoints as an estimate of dependence on these energy sources. No effect of O. rusticus on energy
source existed for bluegill, pumpkinseed, or rockbass. However, as predicted, in lakes with O. rusticus, crayfish predators such as yellow perch,
smallmouth bass and walleye demonstrated a greater dependence on littoral
sources. Therefore, the introduction of O. rusticus increases the importance
of crayfish as an energy pathway to higher trophic levels. However, because
they are large and well defended (large chelae) O. rusticus are generally
less available to a number of fish species than the organisms that they
replace.

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191

KRZYSIK, ANTHONY J.,1,* DAVID A. KOVACIC,2 MICHAEL P. WALLACE,2 JOHN H. GRAHAM,3 JOHN C. ZAK,4 JEFFERY J. DUDA,5
JOHN M. EMLEN5 and D. C. FREEMAN.6 1 Prescott College, Prescott,
AZ, USA; 2 University of Illinois, Urbana-Champaign, IL, USA; 3 Berry
College, Mount Berry, GA, USA; 4 Texas Tech University, Lubbock, TX,
USA; 5 U.S. Geological Survey - BRD - WFRC, Seattle, WA, USA;
6
Wayne State University, Detroit, MI, USA. Robust multivariate approaches for developing ecological indicators to classify landscapes on
a military disturbance gradient.
Nine research sites were selected in Sand Hills physiography of Fort Benning Georgia that represented a disturbance gradient of military training
activities; three sites each in High, Medium, and Low disturbance classes.
High sites have a high level of current mechanized infantry training activities. Medium sites had a past history of military training activities, but are
only lightly used at present. Low sites have not experienced military training activities. Seven Ecological Indicator Systems (EISs) were separately
analyzed with Discriminant Analysis (DA) and Non-metric Multidimensional Scaling (NMS) to extract seven suites of environmental variables
that best characterized this disturbance gradient. These groups were called
Ecological Indicator Guilds, because of their respective functional and defining response to the disturbance gradient. Two EISs, General Habitat
Parameters and Woody Ground Cover Community were capable of separating the three disturbance classes with only a single discriminant function
(DF). The High sites were readily separated from the Low and Medium
sites by DF1 while the Low and Medium sites were separated by DF2 in
three EISs: Ant Community, Soil Chemistry, and Nutrient Leakage. The
Microbial Community EIS was able to separate only the High from the
Low and Medium sites on DF1. The Developmental Instability EIS, using
multiple characters from three perennial plant species, was unable to consistently classify the disturbance gradient. NMS was effective at ordinating
the relative multivariate spatial relationships among all nine research sites.
A-Horizon Soil Depth and Soil Compaction were identified as simple and
reliable ecological indicators to quantify landscape disturbance.
KUBISKE, MARK E.,1,* EVAN MCDONALD,1 PAULA MARQUARDT,1
JOHN KING,2 DON ZAK,3 KURT PREGITZER,2 BILL HOLMES3 and
DAVID KARNOSKY.2 1 US Forest Service, Rhinelander, WI; 2 Michigan
Technological University, Houghton, MI; 3 University of Michigan, Ann
Arbor, MI. Does elevated CO2 alter growth allometry of northern tree
species?
In two separate experiments, various planting arrangements of aspen (Populus tremuloides), birch (Betula papyrifera) and maple (Acer saccharum)
trees were grown in elevated CO2 in factorial combination with soil fertility
(using open-topped chambers), and atmospheric ozone (using the Aspen
FACE facility). Trees were harvested after at least three growing seasons
under treatments, and dissected into dry mass of leaves, wood, coarse roots
and fine roots (,2 mm diam). In both experiments, elevated CO2 significantly increased total tree and organ biomass. The CO2 effect on growth
was augmented by high N soil and eliminated by elevated O3. There were
few significant effects of CO2, soil N, or O3 on biomass allocation among
different plant organs. Regression analysis indicated that such effects were
driven by differences in tree size and were not true shifts in growth allometry due to CO2, soil N or O3. In open-topped chambers, aspen in monoculture were significantly smaller than aspen grown in combination with
maple, whereas the reverse was true for maple. In FACE, aspen trees in
monoculture were significantly larger than those grown with maple. In both
experiments, aspen trees grown in monoculture had a larger proportion of
biomass in stems and a smaller proportion of biomass in roots compared
to those grown in combination with maple. In open-topped chambers, maple grown in combination with aspen had larger proportion of biomass in
leaves whereas those in monoculture had larger proportion in roots. Regressions indicated that differences in biomass allocation among planting
arrangements represented real shifts in growth allometry. Results from both
experiments indicate that elevated CO2, elevated O3, and soil N availability
have minimal effect on growth allometry whereas competitive interactions
with other species are very influential in altering growth allometry.

192

Abstracts

KUDEYAROV, V N.,1,2 K Y. BIL,1,2,* V M. SEMENOV,1 S A. BLAGODATSKII,1 R MURTHY,2 G A. BARRON-GAFFORD,2 E G. DEMYANOVA1 and L HANDLEY.2 1 Russian Academy of Science, Puschino, Russia;
2
Biosphere 2 Center, Columbia University, Oracle, AZ, USA. Effect of
elevated CO2 concentration on soil respiration and above and belowground biomass productivity.
Above and below ground biomass of a coppice cottonwood (Populus deltoides Bartr.) tree system grown under 40, 80, and 120 Pa CO2 was excavated after 3 years of growth in the Intensive Forestry Mesocosm at the
Biosphere 2 Center. Soil respiration at natural moisture level was rather
low and did not differ significantly between plots with different levels of
atmospheric CO2. This may be attributed to the variability of soil moisture
content. Soil respiration at the optimal moisture conditions (basal respiration) for the 120 Pa CO2 was significantly higher than for the 80 or 40 Pa
CO2. The same relationship was found for substrate-induced respiration
(SIR). The greatest increase in respiration induced by moistening was observed for 120 Pa CO2. The 120 Pa mesocosm also had the highest microbial biomass. Results reveal that respiration activity of soil and microbial
biomass in the 120 Pa mesocosm were higher than in the other two mesocosms. Total tree biomass (above+belowground) between the 40 and 80
Pa CO2 mesocosms was significantly different.
KULA, ABIGAIL R.,* DAVID C. HARTNETT and GAIL WT. WILSON.
Kansas State University, Manhattan, KS. Mycorrhizal symbiosis and insect herbivory in tallgrass prairie.
Mycorrhizal symbiosis regulates plant composition and diversity and may
influence plant-herbivore interactions in grassland communities. We conducted greenhouse and field studies to determine the effect of mycorrhizal
fungi on multiple parameters of grasshopper herbivory on tallgrass prairie
plants. We manipulated mycorrhizal colonization of eight co-occurring species growing simultaneously in greenhouse microcosms to examine mycorrhizal effects on grasshopper feeding and plant response to herbivory.
We also grew single species in caged pots to study the role of mycorrhizal
dependence in nymphal development of mixed- and grass-feeding grasshoppers. In field transects of long-term fungicide-treated and control plots,
we surveyed patterns of insect feeding intensity on plants growing with
and without mycorrhizal fungi. Our results indicate no strong effects of
mycorrhizal symbiosis on grasshopper feeding or development. Although
grasshoppers were located more frequently on mycorrhizal plants, feeding
levels on mycorrhizal vs. non-mycorrhizal plants in microcosms did not
differ significantly. Very few factors of grasshopper development were affected by mycorrhizal responsiveness of the host plant, and more frequently, developmental differences were based on natural feeding specificity (i.e.
grass vs. forb). Small differences in feeding on mycorrhizal vs. non-mycorrhizal plants occurred in the field, and the differences were very similar
to the differences in burned vs. unburned sites, indicating that feeding
patterns may be principally influenced by plant community composition.
Mycorrhizal symbiosis positively affected plant re-growth following herbivory. Depending upon the level of herbivory and plants dependence on
the symbiosis, some grazed plants in mycorrhizal microcosms had higher
biomass than their non-defoliated counterparts. This is the first study to
examine mycorrhizae-herbivore interactions of multiple co-occurring plant
species displaying a range of mycotrophy.
KULAKOWSKI, DOMINIK* and THOMAS T. VEBLEN. University of
Colorado - Boulder, Boulder, CO. The persistence of quaking aspen (Populus tremuloides) in west-central Colorado.
Quaking aspen may be the most important deciduous tree in the subalpine
forests of the Rocky Mountains. There has been substantial concern that
aspen has been declining during the 20th century in the western landscape
due to conifer encroachment associated with fire suppression, as well as
other causes. We used a Geographic Information System to compare an
1898 map of vegetation and fires of the Battlement Mesa Forest Reserve,
Colorado to a modern map of present forest cover types in order to assess
long-term changes in extent of quaking aspen. Based on this comparison,
a larger portion of the current landscape is dominated by quaking aspen
relative to the late 19th century, before extensive burning in this area.
During this time period aspen was persistent over most of its extent, even

in the absence of fire. Fires of the late 19th century also increased aspen
cover in stands that were previously dominated by spruce and fir. The
increase in aspen cover occurred primarily at lower elevations. At higher
elevations, where the life history traits of spruce and fir are favored, aspen
has been replaced by the conifers in a limited area. However, the total area
where spruce and fir have replaced aspen is small in comparison to the
area where aspen has increased or has persisted. These findings suggest
that the net effect of large severe disturbances during and after the late
19th century increased aspen cover in the Battlement Mesa area relative to
the reference period that preceded these disturbances. Where the successional replacement of aspen by conifers is occurring today, such a trend
may be a return to conditions more typical of the reference period, prior
to Euro-American settlement. The long intervals between natural disturbances in these ecosystems result in a broad amplitude of natural ecological
patterns.
KULKARNI, MADHURA V.* University of Maryland Center for Environmental Science, Appalachian Laboratory, Frostburg, MD, USA. Contrasting controls on denitrification in a natural and an artificial wetland in the Appalachian mountains.
The denitrification dynamics of soils from a wetland in a high-elevation
reclaimed mineland and a natural, high-elevation wet meadow were studied
through late spring and summer. Soils from the contrasting wetlands had
very different denitrification rates as determined in anaerobic slurry incubations with varying levels of added nitrate and dextrose. The reference
site (natural wetland) soils denitrified at significantly higher baseline (no
amendments) and potential (high dextrose and nitrate amendments) rates
than the mine soils. The baseline denitrification rates scaled areally to
38600 mg N2O-N m-2 hr-1 (SD 5 9800) and 33000 mg N2O-N m-2 hr-1 (SD
5 5000) in the natural and artificial wetlands, respectively. The potential
denitrification rates were 67700 mg N2O-N m-2 hr-1 (SD 5 19000) and
27200 mg N2O-N m-2 hr-1 (SD 5 7700), respectively. Furthermore, denitrification patterns were shown to be controlled by differing factors at each
site, though anoxia-related factors played a large role at both sites. Temperature was the only other factor established to have a relationship with
denitrification dynamics at the mine site while temperature, nitrate, and
dextrose were all shown to have relationships with those of the natural
wetland. Baseline and potential denitrification rates changed over the spring
and summer sampling season at both sites. These results suggest that artificially created wetlands in Appalachian reclaimed minelands may exhibit
lower denitrification rates and less sensitivity to biogeochemical controls
than natural wetlands in the same environment.
KULMATISKI, ANDREW,* KAREN BEARD and JOHN STARK. Utah
State University, Logan, Utah. Plants, soils, and alternative steady states
in plant communities.
Many weed species have invaded a large portion of western North America
degrading agricultural and rangelands. Land managers lack a predictive
knowledge of the ecological and soil factors that determine where different
weeds can establish and whether there are soil and vegetation conditions
that make sites resistant to weed invasion. Recent research on plant-soil
feedbacks suggests a novel and promising approach for the study of invasive plants. We present evidence that soil conditions (biota) play a more
significant role in the outcome of plant-plant competition between native
plant species and exotic invasive species than has been appreciated. We
use the invasive weed Centaurea diffusa (diffuse knapweed) and the soils
of central Washington State as a test system. We determined the relative
importance of soils against more traditional, better-studied factors that have
been invoked to explain invasibility (water-use, seed rain, light, interspecific competition). We present data from a field survey revealing alternative
steady state populations of native and weedy plants in previously-tilled and
never-tilled soils. We also present data from two years of plant growth in
720 experimental plots that isolate seed rain, light, plant competition, and
soil disturbance effects of tillage. We found that knapweed distribution was
limited to tilled soils due largely to seed dispersal and release from competition from live plants. We also present data demonstrating a strong plantsoil feedback that precludes the coexistence of native plants and knapweed.
Fungal community effects were relatively small, but exerted significant
plant community level changes on both knapweed and native plant cultivated soils.

KUMMEL, MIROSLAV* and STEPHEN W. SALANT. University of

Michigan, Ann Arbor, MI. The economics of partner selection in ectomycorrhizal mutualism.
The objective of this paper is to determine whether mutualistic partner
choice in the context of the C for N trading between plants and fungi can
be the mechanism causing both co-occurrence of fungal partners of differing effectiveness on a single host and changes in composition of fungal
partners along light or carbon availability gradients. We investigated the
question both as plants-choose-fungi and as fungi-choose-plants. We constructed a model where growth of the choosing organism is dependent on
the exchange rate of C and N it receives from its trading partners. In the
model, the trading partners are characterized by their C for N exchange
functions. The choosing mutualist selects the composition of its trading
partners that maximize its growth. The results of the model are dependent
on the shape of C for N exchange functions of the trading partners. In the
plants-choose-fungi system both co-occurrence of several ectomycorrhizal
fungal (ECM) species on single host plants and changes in ECM composition along light gradients are possible when fungi have accelerating C
for N exchange functions. In the fungi-choose-plants system changes in
ECM composition on plants along light gradient arise from equivalent nonlinearity in the plants C for N exchange function. Comparison of the
plants-choose-fungi and fungi-choose-plants systems reveals conflicts of
interest among the mutualists. The model is discussed in light of available
empirical evidence.
KUNKLE, JUSTIN M.,* MICHAEL B. WALTERS and RICHARD
KOBE. Michigan State University, Dept. of Forestry, East Lansing, Michigan, USA. Association of root morphology and gas exchange with
northern temperate tree species landform affinity.
Greater use efficiency and access to water may be two general adaptive
mechanisms to low soil water availability. Contrasting glacial landforms
with differences in water holding capacity (outwash-low, ice contact-moderate, moraine-high), and dominant vegetation, in Northwestern Lower
Michigan provide an ideal system to develop a mechanistic understanding
of the association between plant traits and plant performance. First-year
seedlings of eight tree species (in order of increasing site moisture affinity,
(Quercus velutina, Quercus alba, Quercus rubra, Prunus serotina, Acer
rubrum, Acer saccharum, Fraxinus Americana, Betula alleghaniensis) were
transplanted across these landforms. In their third year, leaf gas exchange
and soil moisture were measured monthly, and seedlings were completely
excavated early-summer and fall to obtain growth, size and morphological
characteristics. 2002 had a dry growing season with July-September precipitation of 113 mm vs. a 30-year average of 318 mm. Soil moisture
decreased from moraine to outwash sites and all sites were lowest in July.
Maximum seedling rooting depth differed among species (p , 0.0001) and
was highest for drought tolerant oaks, and lowest for black cherry and red
maple. Rooting depth varied among sites (p , 0.0001) with outwash seedlings deepest rooted, however, contrary to expectations, outwash seedlings
had the lowest root depth extension between summer and fall harvests.
Photosynthesis estimated at common transpiration rates of 0.25 mmol H2O/
m2/s (Aw) differed among sites (p , 0.01) with outwash seedlings generally having higher values for all species. Aw also differed among species
(P, 0.001) but not in the order expected with drought tolerant oaks having
lower values, and thus lower water use efficiency than the other species.
Aw was negatively related to rooting depth on ice contact and moraine
sites indicating lower water use efficiency for more deeply rooted species,
however, there was no clear relationship for outwash sites.
KURSAR, THOMAS A.,1,* BETTINA M.J. ENGELBRECHT1 and MELVIN T. TYREE.2 1 Department of Biology, University of Utah, Salt Lake
City, Utah; 2 USDA Forest Service, PO Box 968, Burlington, Vermont.
Soil moisture release curves for two tropical forests having similar
rainfall but distinct tree communities.
While primary productivity, plant distribution and plant diversity in tropical
rainforests are correlated with water availability, the mechanisms that link
plant productivity, distribution and diversity with differences among tropical rainforest soils in their ability to supply water to plants are not well
understood. We compared two sites in Panama, BCI and Rio Paja, that

Abstracts

193

have very similar rainfall but different plant communities. The Rio Paja
site has plants that are typical of much wetter forest. Soil water release
curves were obtained from about -0.1 MPa to -9 MPa. We found that the
Rio Paja site actually has a much more porous soil than does the BCI soil.
Soil moisture release data alone are not sufficient to explain the observed
distribution and diversity patterns. Nevertheless differences among soils in
their abilities to supply water to plants likely help determine distribution
and abundance in tropical forests. A comparison of moisture release data
with other commonly used measures of soil water availability such as gravimetric water content, indicate that moisture release data provide a better
understanding of the capacity of these soils to supply water to plants.
KWIT, CHARLES,1,* DOUGLAS J. LEVEY,1 CATHRYN H. GREENBERG,2 SCOTT F. PEARSON,3 JOHN P. MCCARTY,4 SARAH SARGENT5 and RONALD L. MUMME.5 1 University of Florida, Gainesville,
FL; 2 US Forest Service, Asheville, NC; 3 Department of Natural Resources, Olympia, WA; 4 University of Nebraska, Omaha, NE; 5 Allegheny College, Meadville, PA. Spatial and temporal variation in fruit production
and consumption in a southeastern U.S. landscape.
Since 1994 we have been gathering data on fleshy fruit production in 5
habitat types at the Savannah River Site (South Carolina, USA) by counting
fruit monthly in replicate (n510-13 per habitat type) 20 x 50-m plots.
Fleshy fruit production, which exhibits dramatic spatiotemporal variation,
typically begins in May, peaks in September and October, and decreases
throughout the winter months. In most habitats, fleshy fruit biomass is
similar to that of hard mast. Most fruit is eaten by omnivorous birds, but
removal rates, proportions, and patterns differ among plant species. Most
fruit consumption occurs in late fall and winter, indicating that winter residents, not migrating birds, rely most heavily on fruit resources. Indeed,
the distribution of some winter residents is related to available fleshy fruit
biomass, and consumption rates of certain winter-fruiting species suggest
that fruits of such species are especially important during cold winters. In
addition to examining ecological relationships between birds and fruits, our
long-term data also provides information for land managers to predict fruit
production in a variety of habitats.
KYLE, SEAN C. * and MARK E. RITCHIE. Department of Biology,
Syracuse Univeristy, 130 College Place, Syracuse, NY, USA; 2 USFS
Rocky Mountain Research Station, C/O Lincoln National Forest, Sacramento Ranger District, Cloudcroft, NM, 88317. Effects of spatial arrangement and climate on the colonization and extinction probabilities
of Utah prairie dog colonies.
1,2,

1 1

The Utah prairie dog (Cynomys parvidens) is currently listed under the
federal Endangered Species Act as threatened. This fossorial species inhabits colonies in semi-arid grasslands and shrublands in the southwestern
quarter of Utah. These colonies may be widely spaced and separated by
areas of varied permeability. Colony-level extinctions and colonizations are
regular events. We examined the role of spatial arrangement of colonies,
colony size, and climate in these events based on annual count data from
over 700 colonies from 1992 to 2002 collected by the Utah Division of
Natural Resources and weather station climate data. Extinction probabilities
are negatively correlated with the average number of prairie dogs in the
colony (P , 0.0001), elevation (P 5 0.0002), annual precipitation from
the previous year (P 5 0.0003), and the number of other colonies within
a 2 mile radius (P 5 0.0016). Colonization probabilities are higher on
private land than public land (P 5 0.0034). Colonization probabilities are
positively correlated with the number of colonies within a 2 mile radius
(P 5 0.0006), the average number of prairie dogs at the nearest colony (P
, 0.0001), and annual precipitation from the previous year (P 5 0.0036).
Colonization probabilities are also negatively correlated with the percentage of frost-free days from the previous year(P 5 0.0136). These results
will benefit the recovery of this threatened species by helping to identify
and rank areas for establishment of new colonies and focusing recovery
efforts in areas that have the highest probability of persistence.
LABRAM, JILL A.* and CRAIG R. ALLEN. South Carolina Cooperative
Fish and Wildlife Research Unit, Clemson University, Clemson, SC. Predictors of habitat use by herpetofauna at multiple scales at the Savannah River Site.
Understanding patterns of habitat selection at different scales is important
for managing and conserving species. It may be insufficient simply to

194

Abstracts

maintain a landcover for a species if required microhabitat components

(e.g., amount of woody debris, ground cover, litter depth) are not provided
within that landcover. We sampled herpetofauna for three seasons (fall
2001, spring 2002, and fall 2002) using pitfall-drift fence arrays, funnel
traps, PVC pipes, coverboards, and visual sightings at the Savannah River
Site. We measured twenty-one habitat variables in 35 sites within seven
landcovers to determine which variables at what scales best predicted the
occurrence of herpetofauna. We used logistic regression to build models
that predicted the presence or absence of eight abundant species (two salamanders, two anurans, three lizards, and one snake). Significant models
included between two and ten variables and r2 values ranged from 0.130.67. The percent litter type was positively associated with the presence of
all species. At least one landcover type was useful in predicting presence
or absence of all species except Tantilla coronata. The presence of the
lizards, Sceloporus undulatus and Scincella lateralis, was associated with
greater percentage of ground shade. Ambystoma opacum, Gastrophryne
carolinensis, and Scincella lateralis were predicted by the presence of
coarse woody debris. Models that incorporate broad-scale elements such
as landcover, together with appropriate fine scale variables, better predict
herpetofauna occurrence than models at single scales.
LACEY, EMERSON S.,1 LAWRENCE M. HANKS1 and JOCELYN G.
MILLAR.2 1 University of Illinois at Urbana-Champaign, Urbana, IL; 2 University of California-Riverside, Riverside, CA. Mate location by the longhorn beetle, Neoclytus acuminatus.
We report evidence that the longhorn beetle, Neoclytus acuminatus, have
both a contact pheromone and a long-range pheromone. Female beetles
have cuticular hydrocarbons that males respond to by mounting the female
when the male antennae come in contact with the female. We have identified two active hydrocarbons as 7Me-C27 and 9Me-C27. Field experiments
show that male beetles produce a long-range attractant that the female
beetle recognizes and follows to the male.
LADD, DAVID G.* and NAOMI CAPPUCCINO. Carleton University, Ottawa, Ontario, Canada. Effective dispersal of the invasive perennial vine
pale swallow-wort Vincetoxicum rossicum.
Invasive plants pose a threat to local flora in many locations around the
world. One such member of the milkweed family is pale swallow-wort
Vincetoxicum rossicum (Kleo.) Barb. Introduced to the northeastern United
States and southern Ontario, pale swallow-wort has come to dominate various localities forming monocultures in old fields and urban parklands. We
investigated the effective dispersal of V. rossicum hypothesizing that smaller seeds would travel greater distances but would be less likely to germinate
and survive. In early April of 2002, seeds that had over-wintered on the
ground were collected, weighed and planted in an old field. Seeds were
individually marked and monitored through the spring, summer and early
fall until the first frost and again the following spring. Later in the same
year, sticky seed traps were placed in a different old field at various distances from a seed source population of plants. Distances of individual
seeds were recorded and masses measured in the laboratory. Seed mass
was a predictor of germination and dispersal distance. Combining these
two measures allowed us to calculate the effective dispersal of V. rossicum.
Future considerations include how best to measure rates of rare long distance dispersal events and their importance to the spread of swallow-worts.
LADEAU, SHANNON L.* and JAMES S. CLARK. Program in Ecology,
Duke University, Durham, NC. Maternal and paternal fecundity of trees
growing in an elevated CO2 atmosphere.
Fecundity plays a key role in determining the species composition of forest
communities While ecological theory predicts fundamental trade-offs
among timing of fecundity, tree longevity, productivity and height, these
relationships are not well understood due to the difficulty of estimating
fecundity parameters in closed stands. Under elevated atmospheric CO2
concentrations, the biodiversity of forests will depend on both changes in
growth rates that confer competitive advantage to some species over others,
and on reproductive success. In addition, although male reproductive effort
is often overlooked in estimates of tree fecundity, pollination can be a
fundamental limit on seed production. Data presented here are from the

FACTS-I site at Duke Forest, NC. Central towers in each of the six rings
allow for direct observation of individual tree fecundity. This study demonstrates that loblolly cone and seed production are greatly enhanced by
CO2 fumigation over five years. Elevated and ambient trees both experienced low fecundity in 2001, but re-established the fumigation effect in
2002. Maturation rates are increased in the elevated rings, with 30% of the
trees producing both male and female cones in 2002 (vs. 15% in ambient
rings). In 2002, more than 40% of the elevated trees produced male strobili,
over twice that in the ambient rings. These data suggest that the maturation
rate of loblolly pine has been increased under CO2 fumigation. Changes in
ontogeny, as implied by this study, may have important implications for
lifespan and tree cycling in forests.
LAFOREST, JOSEPH H.,1,2,* DANIEL A. HERMS1 and PIERLUIGI BONELLO.2 1 The Ohio State University, Department of Entomology, Wooster, OH, United States of America; 2 The Ohio State University, Department of Plant Pathology, Columbus, OH, United States of America. Urban
soil management and its effects on insect herbivory.
Fertilizer and mulch are commonly applied to urban soils degraded by
construction activities in order to ameliorate nutrient deficiencies and stimulate tree growth. Two competing hypotheses address effects of nutrient
availability on tree resistance to insects. If stress weakens plant defense,
fertilization may reduce herbivory by increasing tree vigor. On the other
hand, if there exists an allocation trade off between growth and defense,
fertilized trees will grow faster, but be less resistant to herbivores. In a twoyear field study, we tested these competing hypotheses by quantifying the
effects of four soil management treatments (fertilization, mulching with
composted yard waste, mulching with composted bark / manure blend, and
bare soil control) applied in factorial combination to soil types (topsoil,
inverted subsoil) on microbial activity, nutrient cycling, as well as growth
phenolic chemistry, and insect resistance of paper birch (Betula papyrifera).
Trees in topsoil grew faster than those in subsoil regardless of soil treatment, but were less resistant to insects as indicated by higher larval growth
rates of forest tent caterpillar [Malacosoma disstria], whitemarked tussock
moth [Orgyia leucostigma], yellow-necked caterpillar [Datana ministra]
and fall webworm [Hyphantria cunea]. Fertilization of subsoil plots increased tree growth but decreased foliar phenolic content, and insect resistance, with the larval growth of some species increasing to levels observed
on trees in topsoil. There was a significant negative correlation between
total phenolic content and tree growth. These results are consistent with
the hypothesis that faster growing trees are less insect resistant due to a
physiological tradeoff between growth and chemical defense.
LAHTI, MEGAN,1,* CHERYL SWIFT1,* and KATIE ROSS.2 1 Whittier
College, Whittiier, CA; 2 University of Virginia, Charlottesville, VA.
ENSO event effects on Southern California riparian plant communities.
Highly variable rainfall typical of Mediterranean climates results in large
year to year fluctuations in stream discharge. In the 1998 ENSO event Los
Angeles County received almost three times the normal annual precipitation. The majority of this rainfall occurred in a three month period extending from January to March, and stream discharges increased significantly
during this period. We sampled the plant community along two reaches of
the San Gabriel River in 2002 in order to examine the effects of this increased discharge on plant community structure. Each of the two reaches
had been sampled previously in 1995. Reaches were sampled by setting
out a series of 100 m 2 plots within 5 m of the thalwag, 5-20 m from the
thalwag and greater than 20 m from the thalwag. The community changed
significantly as a result of the ENSO event. The number of stems/ha was
significantly higher in 2002, but the stems were smaller since a significantly
larger proportion of stems was less than 3 cm in diameter as compared to
1998 where a larger proportion of stems were greater than 8 cm in diameter.
There were also significantly more stems per individual in 2002 suggesting
that the increase in stems may be the result of resprouting from fallen
stems. These changes in community structure were most apparent closer to
the stream. The results of this study suggest that the 1998 ENSO event
changed the structure of the riparian plant community significantly and that
Southern California riparian plant communities respond to disturbance by
resprouting from existing fallen individuals and then undergoing a period

of self thinning as light becomes limiting until a more open forest composed of larger trees establishes.
LAKE, FRANK K.* Oregon State University, Corvallis, Oregon, USA.
Traditional ecological knowledge: Native perspectives on fire ecology,
historical landscape changes, and forest restoration.
Native peoples of Northwestern California possess Traditional Ecological
Knowledge (TEK) of fire ecology and historical landscape changes related
to forest composition, structure, and function. TEK is being integrated with
fuels reduction projects to restore and conserve biodiversity and productity
of degraded forest habitats in the Klamath-Siskiyou ecoregion. Interviews
of native elders on fire ecology, management, and implications for reinstating cultural burning practices will be discussed. On the ground fuels
reduction and prescribed burning projects that reflect the integration of
TEK with forest restoration principles and criteria will be highlighted.
LAKE, JEFFREY K.,1,* STEPHEN P. HUBBELL1,2 and W. JOHN
KRESS.3 1 University of Georgia, Athens, GA; 2 Smithsonian Tropical Research Institute, Balboa, Panama, Panama; 3 National Museum of Natural
History, Washington, DC. Lineage diversification under neutral theory:
An empirical test.
Classical community ecology theory rarely incorporates considerations
over evolutionary time periods or considers predictions about evolutionary
divergence of taxa. A new theory of biodiversity and biogeography, however, is unique in that it presents a number of testable hypotheses concerning lineage divergence and persistence (Hubbell 2001). Under this neutral
theory, individuals have equal per capita probabilities of birth, death, and
dispersal, and speciation is modeled as the probability of a speciation event
per individual birth. Because abundant lineages have more births per unit
time and are expected to have longer evolutionary lifespans, abundant lineages are expected to produce more daughter lineages than rare lineages
simply by chance. Older lineages on average should be more abundant than
recently derived ones. The theory, under the point mutation model of speciation, also predicts a fractal relationship between lineage number and time
depth within a monophyletic clade. In contrast, other models of speciation,
such as random fission speciation or peripheral isolate speciation, do not
show a single fractal-scaling region. We explore this and other theoretical
predictions of neutral theory and test them using a recently published phylogeny of family Costaceae in combination with extensive range data for
the family, gathered from herbarium specimens and collections databases.
Using empirical data from Costaceae, we examine the relationships between lineage age and range size, measure species richness in individual
clades as a function of clade age, and compare these observations.
LAMAR, WILLIAM R.* and JAMES B. MCGRAW. West Virginia University, Morgantown, WV. Investigation of population dynamics of Eastern Hemlock (Tsuga canadensis) using remote sensing.
Our goal was to investigate the population dynamics of Eastern Hemlock
(Tsuga canadensis) using remote sensing. We emphasize the advantages of
using remotely sensed data to collect large, spatially explicit databases.
Large-scale, low elevation aerial photographs of a naturally occurring population of Eastern Hemlock was collected in early spring 1997, 1998, and
1999. Using automated spectral and spatial segmentation procedures, the
population was divided into individual units. To ensure compatibility of
multi year imagery, an automated reconciliation procedure was applied to
the data sets following segmentation. Estimates of matrix population model
parameters were obtained from the size structured, multi year imagery.
Fertility estimates were calculated as a function of the distance and size of
adult trees (at time a) from canopy "births"(at time a+1). Survival, growth,
reproduction, population growth rates, and size specific sensitivity values
were compared for the 1997-98 and 1998-99 aerial image pairs. The matrix
population models using aerially obtained parameter estimates were compared to models using estimates from traditional ground collected measurements.

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195

LAMBRECHT, SUSAN,1,4,* MICHAEL LOIK,1,4 DAVID INOUYE2,4 and

JOHN HARTE.3,4 1 University of California, Santa Cruz, CA; 2 University
of Maryland, College Park, MD; 3 University of California, Berkeley, CA;
4
Rocky Mountain BIological Laboratory, Gothic, CO. How are the rates
and resource costs of plant reproduction affected by global warming?
Under predicted patterns of climate change, the distribution of plant species
is expected to shift. The capacity to reproduce is one factor contributing
to the potential of a species to persist within its current range or to disperse
to new ranges as climate changes. To address how plant reproduction may
be influenced by global warming, we have used an ongoing experiment in
a Colorado subalpine meadow that employs infrared warming to simulate
future radiative forcing near the soil surface. We tested the hypothesis that
allocation of resources to reproduction may be partially explained by responses to earlier snowmelt in the spring, which leads to increased exposure to episodic freezing events, and decreased soil moisture in the summer.
For two species that exhibit reduced rates of reproduction, the niveal emergent Erythronium grandiflorum, and summer-flowering Delphinium nuttallianum, we measured the water costs of reproduction as well as foliar gas
exchange rates and properties. For D. nuttallianum, floral transpiration rates
were greater in the warmed plots than in the control plots. Furthermore,
leaf water potential, photosynthesis, and stomatal conductance were significantly lower in the heated plots as compared to the control plots. However, warming did not produce similar effects for E. grandiflorum, for
which increased exposure to spring freezing events damages the reproductive structures, precluding subsequent flower and fruit production. Understanding the mechanisms that reduce reproduction for these species while
producing no response in others exposed to the warming experiment is
critical for discerning how the abundance and distribution of species will
be altered by climate change.
LAMONTAGNE, JALENE M.1 and ROBYN L. IRVINE.2,* 1 University
of Alberta, Edmonton, Alberta, Canada; 2 University of Calgary, rlirvine@ucalgary.ca, Calgary, Alberta, Canada. Sage grouse in space: What
measure of spatial heterogeneity is most meaningful in assessing population dynamics?
Sage grouse populations (Centrocercus urophasianus) in Utah are in decline. Sage grouse populations in Utah are estimated to occupy only 50%
of the habitat they once did and their abundance is approximately half what
it was in 1850. The strutting grounds (leks) for sage grouse are distributed
throughout the state and these leks are situated at a wide range of vertical
(approx. 1300 to 2800 meters above sea level) and horizontal (up to 1400
km apart) locations. Despite the increasing recognition of the role of habitat
heterogeneity and environmental variability in driving population cycles,
the role of altitude in conjunction with the role of the linear separation
distance between populations has not been explored. Sage grouse in Utah
are predicted to be listed as endangered within a year. It is therefore important to understand what drives the dynamics of the spatially separate
populations scattered around the state to allow management plans to address the decline. We modeled the population dynamics of 150 sage grouse
populations, restricting our analysis to populations with a minimum of 10
years of count data, and we assessed the spatial autocorrelation among leks
using non-centered correlograms. Linear distances (latitude and longitude),
vertical distances (altitude), and absolute distances (the hypotenuse distance
of the altitude and latitude/longitude triangle) were assessed within the
structure of the correlogram to infer spatial patterns of different types of
population dynamics. We will present results for the effects of the different
distance types on correlations in population fluctuations and the spatial
patterning of population dynamics. We will also comment on the management implications associated with different spatial locations of populations.
LAMPHERE, BRADLEY A.* University of North Carolina - Chapel Hill,
lamphere@email.unc.edu, Chapel Hill, NC. Connecting multi-scale
movement patterns to population dynamics in a stream fish, Cottus
bairdi.
Long moves, which represent the tails of a movement distribution, have
great impact on population dynamics. Unfortunately, the infrequency of
long moves makes them difficult to observe in the field for most species.
As a result, to study the extent of and cues to movement, I selected a

196

Abstracts

stream fish noted for its sedentary habits, the mottled sculpin, Cottus bairdi.
If movement beyond normal home range is significant and scale-dependent
in C. bairdi, it likely is for more vagile species as well. Over 43% of
individually-marked recaptures moved 20 m or more. Moreover, the proximal cues to reach-scale moves (20 2 180 m) differed from shorter moves,
particularly during the spring breeding season. While moves during summer exhibited the negative density dependence expected of a territorial
species, moves which occurred in the spring showed the opposite trend
(i.e., conspecific attraction). Next, I manipulated C. bairdi density and
breeding habitat availability during summer and spring to determine whether the conspecific attraction was (1) a direct response to C. bairdi being
present, or (2) an indirect result of C. bairdi being attracted to breeding
habitat. In the absence of direct information on true dispersal events, ecologists often use extrapolations from mark-recapture data to estimate the
shape of the movement distribution9s tail. Alternatively,genetic data can
provide a more realistic estimate of the tail9s shape, beyond the range of
traditional mark-recapture. I plan to use microsatellite DNA loci to (1) infer
dispersal extent in this population and (2) make quantitative comparisons
between the genetic data and several new and existing models for extrapolating mark-recapture data. Together, the mark-recapture, experimental,
and genetic data indicate that even an exceptionally sedentary fish can
exhibit complex and scale-dependent movement patterns at several levels.
These results will directly aid efforts to use C. bairdi in biomonitoring and
more generally enhance our understanding of the proximal causes and population-level consequences of movement.
LANCASTER, JILL,1 BARBARA J. DOWNES2,* and PAUL REICH.2
1
University of Edinburgh, Edinburgh, Scotland, Scotland; 2 University of
Melbourne, Melbourne, Australia, Australia. Linking landscape patterns
of resource distribution with models of aggregation in ovipositing caddisflies.
Spatially-explicit measures of dispersion were employed to detect how any
aggregation of individuals is influenced by the dispersion of resource patches. For two species of caddisfly (Hydrobiosidae) in two stream stretches,
we mapped the spatial distribution of potential and exploited patches (emergent rocks as oviposition sites), and enumerated egg masses. As documented previously, egg masses of Ulmerochorema and Apsilochorema
were aggregated on individual rocks and both species showed velocityspecific preferences for oviposition sites: oviposition sites of Ulmerochorema occurred most often in fast flowing water; those of Apsilochorema in
slow water. Point pattern analysis was used to describe the spatial pattern
of emergent rocks, and to test hypotheses about how site selection behaviour of females influences dispersion. Emergent rocks were clumped and
equally abundant in both stretches. Our perception of how caddisflies deposit eggs into the landscape varied with the neutral landscape model. The
most informative NLM compared the spatial pattern of oviposition sites
with the underlying pattern of emergent rocks, while constraining the random selection of rocks to reflect species velocity preferences. Oviposition
sites of Ulmerochorema were clumped spatially, over and above the background pattern; those of Apsilochorema were over-dispersed spatially.
LANDENBERGER, RICK E.,* JAMES B. MCGRAW, TIMOTHY A.
WARNER and TOMAS BRANDTBERG. Department of Biology, West
Virginia University, Morgantown, WV, USA. Aerial census of Haleakala
silverswords using high resolution color-infrared imagery.
High spatial resolution remote sensing offers largely untapped potential for
censusing and monitoring rare plant populations. The Haleakala silversword is a federally listed threatened species whose natural range is restricted to the highest elevations on Maui. With its distinctive foliage set against
the volcanic background, the species provides an excellent test of the capabilities of color-infrared remote sensing to provide a spatially explicit,
individual-based approach to monitoring. We used a helicopter-mounted,
high spatial resolution digital camera system with a color-infrared filter to
image a series of permanent census plots. Ground-based censuses from
2001 were compared to photointerpretation and automated computer classification of super-high resolution color-infrared imagery. Errors of omission and commission occurred in both methods, although at low rates.
Photointerpreters and automated classification accurately estimated silversword size in aerial images, with higher accuracy in the larger size classes.

Although imperfect, both photointerpretation and classification effectively

differentiated between living and dead individuals. Relative to photointerpreters, flowering individuals presented challenges for the classification.
Although not as accurate as localized ground-based censuses, high resolution aerial censuses provide a rapid partial population census over larger
areas, which may be adequate for monitoring rare plants growing in barren
landscapes.
LANDIS, FRANK,* THOMAS GIVNISH and ANDREA GARGAS. University of Wisconsin-Madison, Madison, WI. Arbuscular mycorrhizal
fungi and understory plant diversity in Wisconsin oak savannas.
Arbuscular mycorrhizal fungi (Glomeromycota, AMF) are ubiquitous in
most terrestrial plant communities, and a number of researchers have shown
that AMF can affect plant community structure and composition in different systems. Furthermore, glomalean fungi are obligate biotrophs, obtaining
all their carbohydrates from their photosynthetic partners. Combining these
facts creates an interesting question: How do AMF influence plant community composition and structure along steep light gradients? This situation
exists in Wisconsin oak savannas, which are among the most species-rich
and endangered plant communities in the upper Midwest. Scattered savanna
oaks create steep light gradients over short distances, and recent research
has shown that these light gradients, along with soil texture patterns, influence the distribution of understory plant species. Our study looks at the
relationships between plant diversity, glomalean diversity and light, and
soil N and P concentrations. One hundred eight 1-m2 plots were established
in three remnant oak savannas in Wisconsin. Plant species and percent
cover were tallied repeatedly, and soil cores were taken to determine the
species of AMF present. Soil nutrient levels were measured, and hemispheric photography was used to estimate the light regime. Data were processed via NMS ordination and a simple structural equation model. The
results show a positive correlation between AMF and plant species richness, and surprising interactions among the abiotic factors, AMF composition, and plant community composition.
LANGDON, CHRIS,1 WALLACE S. BROECKER,2 DOUGLAS E. HAMMOND,3 EDWARD GLENN,4 KEVIN FITZSIMMONS,4 STEVEN G.
NELSON,4 TSUNG-HUNG PENG5 and IRKA HAJDAS.6 1 Biosphere 2
Center of Columbia University, Oracle, AZ; 2 Lamont-Doherty Earth Obs.
of Columbia University, Palisades, NY; 3 Dept. of Earth Sciences, Uni. of
S. California, Los Angeles, CA; 4 Environmental Research Laboratory, Uni.
of Arizona, Tucson, AZ; 5 NOAA-AOML, Miami, FL; 6 IPP ETH Hoenggerberg, Zurich, Switzerland. Effect of elevated CO2 on the community
metabolism of an experimental coral reef.
The effect of elevated pCO2 on the metabolism of a coral reef community
dominated by macroalgae has been investigated utilizing the large 2650 m3
coral reef mesocosm at the Biosphere-2 facility near Tucson, Arizona. The
carbonate chemistry of the water was manipulated to simulate present day
and a doubled CO2 future condition. Each experiment consisted of a 1-2
month preconditioning period followed by a 7-9 day observational period.
The pCO2 was 404663 matm during the present day pCO2 experiment and
658659 matm during the elevated pCO2 experiment. Nutrient levels were
low and typical of natural reefs waters (NO3- 0.5-0.9 mM, NH4+ 0.4 mM,
PO43- 0.07-0.09 mM). As previously observed, the increase in pCO2 resulted
in a decrease in calcification from 0.04160.007 to 0.00660.003 mol
CaCO3 m-2 d-1. Net community production (NCP) and dark respiration did
not change in response to elevated pCO2. Light respiration measured by a
new radiocarbon isotope dilution method exceeded dark respiration by a
factor of 1.260.3 to 2.160.4 on a daily basis. The 1.8-fold increase with
increasing pCO2 indicates that the enhanced respiration in the light was not
due to photorespiration. Gross production (GPP) computed as the sum of
NCP plus daily respiration (light+dark) increased significantly (0.2460.03
vs. 0.3260.04 mol C m-2 d-1). However, the conventional calculation of
GPP based on the assumption that respiration in the light proceeds at the
same rate as the dark underestimated the true rate of GPP by 41-100% and
completely missed the increased rate of carbon cycling due to elevated
pCO2. We conclude that under natural, undisturbed, nutrient-limited conditions elevated CO2 depresses calcification, stimulates the rate of turnover
of organic carbon, particularly in the light, but has no effect on net organic
production.

LANGLEY, SUSAN K.,1 CECIL C. FROST,2 JOHN E. FELS3 and THOMAS R. WENTWORTH.3 1 Georgia Southern University, langley@gasou.edu, Statesboro, GA; 2 North Carolina Plant Conservation Program, Raleigh, NC; 3 North Carolina State University, Raleigh, NC. Landscape fire ecology: Defining fire compartments and a topographic index
of fire probability.
Fire compartments are a central concept of landscape-level fire ecology. A
fire compartment is a unit of the landscape having continuous fuel and no
natural fire breaks, such that an ignition would be likely to burn the entire
compartment barring changes in weather or fuel moisture. We hypothesized
that presettlement (potential/natural) distributions of fire-dependent longleaf pine (Pinus palustris) communities could be accurately mapped using
a new environmental variable, presettlement fire compartment size. We
quantify the average fire probability index (AFPI) for any point within a
fire compartment using topographic features and compartment size. GIS
layers with watershed information were used with slope-class maps derived
from DEMs to map fire compartments in the Uwharrie National Forest,
North Carolina. The AFPI for points in this landscape was predicted using
a GIS and an algorithm that included distances to firebreaks and prevailing
wind direction during fire season. AFPI values were assigned to one of five
topographic fire regime (TFR) classes, from class 1 (the most fire protected
areas) to class 5 (the most fire exposed areas). TFR classes were comparatively assessed using 209 witness trees, mapped in a GIS layer from
county survey plats of the study area ca. 1700-1800. Trees were assigned
to vegetation fire regime (VFR) classes based on species characteristics.
The most fire refugial species were assigned to VFR class 1 and the most
fire-dependent species were assigned to class 5. Correspondence between
VFR and TFR classes was 86%. Our hypothesis that presettlement firedependent communities could be accurately mapped using fire compartment size was generally supported by our results. Our weighting scheme
also appears to be realistic for this landscape. The AFPI for a particular
site may be the most ecologically meaningful environmental variable for
understanding and managing remnant natural areas to maintain original
levels of biodiversity in fire-dependent landscapes.
LANZA, JANET,1,* MARJORIE STEPHEN,1,2 KARA A. DAVIS1 and
MYLINDA L. TERRY.1 1 University of Arkansas at Little Rock, jxlanza@ualr.edu, Little Rock, AR; 2 Arkansas Baptist Middle School, Little
Rock, AR. Nectar-borne amino acids increase energy reserves in and
reproduction by monarch butterflies.
Nectar, the reward many plants provide for pollinators, contains water, sugar, and amino acids. Although cabbage white butterflies (Pieris rapae) and
monarch butterflies (Danaus plexippus) prefer sugar-amino acid nectar
mimics over sugar-only nectar mimics, the nutritional role of nectar-borne
amino acids in pollinators is poorly studied. In order to determine if nectarborne amino acids increased adult energy reserves, the number of egglaying days, total egg production, and hatching rate, monarch butterflies
were fed either sugar-only or sugar-amino acid nectar mimics of Lantana
camara. Energy reserves were measured on newly emerged adult butterflies
and unmated butterflies (5 day and 7 day old) fed sugar-only and sugaramino acid nectars. Males, but not females, accumulated energy reserves
more quickly when fed sugar-amino acid nectars than when fed sugar-only
nectars. In order to measure the effect of amino acids on reproduction,
mated females were placed in individual cages with a sprig of milkweed
on which to lay eggs. Eggs were counted daily and the hatching rate was
measured for twenty eggs from each butterfly each day. Compared to butterflies fed the sugar-only nectar, butterflies that were fed sugar-amino acid
nectar produced more eggs and tended to lay eggs on more days. The amino
acids did not affect hatching rate. These results demonstrate that nectarborne amino acids contribute to the nutrition of monarch butterflies.
LAROCQUE, GUY R., ROBERT BOUTIN, DAVID PARE, GILLES
ROBITAILLE and VALERIE LACERTE. Natural Resources Canada, Laurentian Forestry Centre, Ste-Foy, Quebec, Canada. Modelling the soil carbon and nitrogen cycles in three eastern Canadian forest types.
A process-based model was developed to simulate the soil carbon and
nitrogen cycles in sugar maple (Acer saccharum Marsh.), balsam fir (Abies
balsamea (L.) Mill.) and black spruce (Picea mariana (Mill.) B.S.P.) forest

Abstracts

197

types in eastern Canada. The development and calibration of the model are
part of the ECOLEAP project that involved the establishment of field stations and laboratory incubation microcosms for collecting data on the carbon and nitrogen pools and fluxes in the litter, organic and mineral layers
of the soil in these forest ecosystems. Litter samples were also incubated
on several sites along a climatic gradient to monitor site-specific litter decomposition rates for a complete annual cycle. Soil temperature was monitored on a daily basis. The model simulates the carbon and nitrogen transfers among the different pools, as well as the amount of CO2 respired. For
the organic and mineral layers, the carbon was partitioned into active, slow
and passive pools. An Arrhenius-type function was used to model the effect
of temperature on mineralization rates. The simulations conducted over a
period of 50 years indicated that the different carbon pools will decrease
significantly with a gradual 38C increase in temperature, while the amount
of CO2 respired will increase. However, the results also indicated that the
decrease in carbon pools may be offset by an increase in litter production.
LARRIMER, AUDREY K.* and BRIAN C. MCCARTHY. Ohio University, Athens, Ohio. Biological diversity associated with bigtooth aspen
(Populus grandidentata) clones in a mixed-oak landscape.
Environmental heterogeneity is a commonly proposed mechanism for the
maintenance of species diversity. Biotic factors, including forest overstory
composition, may contribute to variability in resource availability and environmental conditions, thereby leading to increased understory species
richness. The objective of this study was to examine the ability of the clonal
species Populus grandidentata to alter resource availability and environmental conditions beneath its canopy, thereby increasing the environmental
heterogeneity of the forest landscape as a whole. Twenty P. grandidentata
clones were located in southeastern Ohio, and adjacent control sites delineated. Resource availability and environmental factors were quantified in
each clone and compared to control sites. Soil pH was found to be significantly greater in P. grandidentata clones than in adjacent control sites (P
, 0.05). Percent soil organic matter was significantly less in P. grandidentata clones than in control sites (P , 0.01). No differences in soil
nitrate, phosphorus, moisture, or light were detected between treatment and
control plots. The results suggest that P. grandidentata alters certain environmental conditions beneath its canopy, possibly through differences in
leaf chemistry. A vegetation survey currently underway will evaluate the
relevance of microevironmental variation to the composition and structure
of understory communities inside and outside of P. grandidentata clones.
Observations suggest that P. grandidentata clones may increase forest beta
diversity by increasing environmental heterogeneity.
LARSON, KATHERINE C. University of Central Arkansas, klarson@mail.uca.edu, Conway, AR. Pollinator visitation to the exotic japanese honeysuckle: Impacts on fruit set and vegetative growth.
The exotic Lonicera japonica requires pollen from another individual plant
for successful fruit set, and thus within its invaded range in the southeastern
US, it is completely dependent on native pollinators for seed production. I
quantified much lower insect visitation rates to L. japonica than to other
simultaneously flowering species, thus suggesting that fruit set in L. japonica was pollen limited. Experimental hand pollinations showed that fruit
set could be increased from the natural control level of 17% to 78% when
an abundance of pollen from another individual was applied to receptive
stigmas. Japanese honeysuckle also reproduces vegetatively through horizontal shoots that root at the nodes. To determine if resources for fruit
development were diverted to additional vegetative growth when pollination was low, I contrasted experimental plants getting supplemental pollination with plants having low levels of pollination. Allocation to vegetative
growth was quantified in these two groups of plants. The potential impacts
of pollination success on the rate of vegetative growth and invasiveness of
L. japonica is discussed.
LASSEN, KARI E.* and E WILLIAM HAMILTON. Washington and Lee
University, Department of Biology, Lexington, VA. Competitive interactions between the endangered sink-hole species Helenium virginicum
and Boltonia asteroides when exposed to variable water depths.
Helenium virginicum (Virginia sneezeweed) is an herbaceous plant from
the family Asteraceae and is classified as a threatened species federally and

198

Abstracts

as endangered by the state of Virginia. It is found in seasonally flooded

sink-hole ponds in 30 sites in Virginia and 1 site in southern Missouri. In
sites where Helenium is found Boltonia asteroides (Asteraceae) also occurs
in high numbers. To help identify important ecological variables that influence the recovery of Helenium three experiments were conducted. The first
was a germination experiment in which it was determined that Helenium
germinates when soils are at field capacity but does not germinate when
submerged under water due to anoxia. Boltonia germinates under all conditions (e.g. flooding, under a standing water column and at field capacity).
The second experiment quantified photosynthetic tolerance of each species
when subjected to water logging and submergence. Chlorophyll content,
chlorophyll fluorescence (Fv/Fm), and net photosynthesis were used to quantify effects on the light reactions and Calvin cycle, respectively. Helenium
had reduced but positive photosynthetic rates when waterlogged and submerged and increased chlorophyll a:b ratios. Boltonia was not photosynthetically tolerant to water logging and submergence and had reduced chlorophyll concentrations. The third experiment was a competition experiment
designed to determine potential interactions that determine the distribution
of Helenium and Boltonia in sink-hole ponds. Biomass accumulation in
Helenium when competing with Boltonia was significantly lower than intraspecific competition biomass and only when plants were submerged could
Helenium out-compete Boltonia. These experiments suggest that in order
to successfully implement a recovery strategy for Helenium water levels
and hydro-periods must be considered and or managed in suitable Helenium
habitats.
LATTY, ERIKA F.1,2,* and DAVID J. MLADENOFF.2 1 Hollins University,
Roanoke, VA; 2 University of Wisconsin-Madison, Madison, WI. Considering multiple temporal and spatial scales in determining reference
conditions for Great Lakes forest restoration.
In establishing restoration goals it is difficult to choose both a single time
period applicable to all management needs and a geographical extent that
accurately characterizes system dynamics. To avoid these pitfalls requires
an approach that integrates multiple data sources spanning several temporal
and spatial scales. The goal of our study was to describe the natural range
of variation in northern-hardwood and hemlock-hardwood forests of the
northern Great Lakes states, USA, using historical data and empirical field
data at two spatial scales, that of the landscape and the forested stand.
Determining reference conditions based on landscapes free of all human
influence is not a possibility, thus historical data provided an understanding
of forest parameters prior to Euro-American settlement. Still, it is not clear
that forest characteristics as determined by historical analyses are resilient
enough to withstand current conditions and endure through future surprises.
Therefore we synthesized existing, current data on the structural components, spatial characteristics, and biogeochemical processes of remnant oldgrowth forests with historical data sources to establish baseline variability.
Using this method we suggest a set of reference conditions that bracket a
range of possible outcomes in order to guide restoration goals. These reference conditions are then compared to the range of variability detected in
the same parameters measured in managed forests of the region. For some
parameters, such as net nitrogen mineralization rates, management practices
resulted in conditions similar to expectations based on the established reference conditions. However, forest management significantly altered other
parameters, like the amount of coarse woody debris. Although, our analyses
were limited to specific forest types within a limited geographic region, we
suggest that our approach has wider application as it provides a conceptual
framework useful for managing dynamic forested ecosystems that are vulnerable to uncertainties due to future human influences.
LAU, JENNIFER. UC Davis, jalau@ucdavis.edu, Davis, CA. Ecological
and evolutionary responses of native species to novel community members: Direct and indirect effects of invasion.
When a novel species enters a community, it can have both direct and
indirect effects on native species within that community. For example, plant
invaders can affect native plants both directly, via competition, or indirectly
by altering the native species9 interactions with other organisms within the
community. My work addresses the ecological and evolutionary consequences of the invasion of an exotic plant and its effects on a co-occurring
native via competition and through the native plant9s interactions with its

herbivores. The exotic plant Medicago polymorpha increases levels of herbivory on the native plant Lotus wrangelianus by increasing densities of
legume-feeding herbivores. Phenotypic selection analysis revealed that Lotus individuals in the presence of Medicago experience more intense selection for resistance to weevil folivory than Lotus individuals in the absence of Medicago. Evolutionary responses to direct competition with the
Medicago were also observed. The present study uses a two-by-two factorial design where both herbivore presence and Medicago presence are
manipulated to determine the relative roles of indirect and direct effects of
an invader on both the ecology and evolution of native L. wrangelianus.
LAUGHLIN, KAREN D.,1,* ALISON G. POWER1 and ALLISON A.
SNOW.2 1 Cornell University, Ithaca, NY; 2 Ohio State University, Columbus, OH. Fitness of wild Cucurbita pepo after introgression of transgenic resistance to zucchini yellow mosaic virus.
One risk of genetically modified (GM) virus-resistant plants is the introgression of transgenic virus-resistance into closely related wild plant populations. If disease pressure limits the wild plant population, then transgenic
virus resistance could increase the fitness of the plant under virus infection
and potentially increase the plants invasive ability. We used a cultivated
squash (Cucurbita pepo) engineered for resistance against zucchini yellow
mosaic virus (ZYMV) and a conspecific wild squash to study the effect of
a virus-resistance transgene on wild plant fitness. To mimic introgression
of the virus-resistance transgene into a wild population, we crossed the GM
squash with the wild squash and then backcrossed transgenic progeny into
wild squash for two successive generations. We compared the fitness of
wild individuals, transgenic backcross individuals and non-transgenic backcross individuals under ZYMV infection and healthy conditions. We measured total male and female flower production, fruit and seed number, and
dry biomass to quantify reproductive and vegetative fitness. Under healthy
conditions, wild, transgenic backcross and non-transgenic backcross individuals showed no significant difference in fitness measurements. Under
ZYMV infection, transgenic backcross individuals showed significantly
higher fitness than wild individuals. Non-transgenic backcross individuals
also showed significantly higher reproductive fitness than wild C. pepo,
though not as high as transgenic backcrosses. These results suggest that
wild C. pepo fitness would increase with introgression of transgenic
ZYMV-resistance if wild C. pepo populations are limited by ZYMV pressure. They also indicate that other conventional crop genes would contribute ZYMV-resistance to wild populations.
LAURENT, EDWARD J., * JOSEPH P. LEBOUTON, MICHAEL B.
WALTERS2 and JIANGUO LIU.1 1 Department of Fisheries & Wildlife,
Michigan State University, East Lansing, MI, USA; 2 Department of Forestry, Michigan State University, East Lansing, MI, USA. Evaluating the
effects of landcover classification parameters on landscape indices used
to investigate wildlife-habitat relationships.
1,

Many studies of wildlife distribution patterns are based on land cover maps
created from satellite images. While research into the relationships between
wildlife distributions and land cover classes have been instrumental for
increasing knowledge of species occurrence patterns, descriptions of landscape heterogeneity vary with the grain of analysis, the classification system in use, and the variability of spectral information employed. Further,
land cover classes often do not accurately classify the landscape in ways
that many species respond. For these reasons, an examination of classification parameters on map accuracy is badly needed. To address these concerns we used a grain representative assessment and inventory protocol
(GRAIN) to survey sample locations (N 5 198) for forest bird species in
the Upper Peninsula of Michigan. Pixels overlapping survey plots were
used as seed pixels in a habitat analysis by iterative classification procedure
(HABICLASS) to create spectral signatures that described survey plots.
These signatures were then used in supervised image classification to classify maps of species occurrence. Parameter values employed in supervised
classification were incrementally modified to assess their influence on map
accuracy. Map accuracy was affected by parameter values in different ways
for each species. Parameter values resulting in the highest map accuracy
for species occurrence were used to classify maps of land cover specific to
each bird species. Landscape indices previously documented as having influence on the study species occurrence or life history were calculated for
each land cover map and compared.

LAWLER, SHARON P.* University of California, Davis, CA, USA.

Should ecologists throw away their petri dishes and aquaria?
Understanding of the limitations and benefits of microcosm studies is essential to their use in ecology. Microcosms are sometimes too small to
allow full expression of processes that may be important at larger scales
(e.g., migration). Scale transitions usually prevent quantitative extrapolation
of microcosm results. They are prone to artifacts including wall effects,
build-up of toxins, chemical imbalances, sampling depletion, and isolation
from environmental and biotic conditions that were of unforeseen importance. Note that careful examination of isolation effects and other artifacts
may advance theory by revealing overlooked factors. On the positive side,
creative microcosm experiments can provide powerful evaluation of theory
and mechanisms, yielding testable, if qualitative, predictions for large systems. Well-known advantages of microcosm research include replicability,
control, ease of manipulation and sampling, and the opportunity to observe
population dynamics. Like any experimental system, microcosms vary in
levels of realism. Minature worlds (e.g., phytotelmata, rock pools) provide
elements of realism missing from laboratory microcosms, such as realistic
levels of environmental heterogeneity, disturbance, dispersal, and variation
in community structure. Given these advantages, natural microcosms are a
better choice for testing some aspects of ecological theory, especially if the
robustness of theory to a variable world is a concern. However, it is more
difficult to prove mechanisms or to dissect signal from noise when environmental and biotic conditions are less controlled. Some kinds of theory
are more easily tested in laboratory microcosms than natural microcosms.
Examples include theories about environmental noise and resonance, and
theory pertaining to evolutionary ecology. In general, fears that microcosm
results will mislead science have not been borne out. Instead, microcosms
have served as an essential link in the scientific cycle of observation, development of theory, and experimental tests that lead to new observations
and refinement of theory.
LAWSON, DAWN M.,1,* PAUL H. ZEDLER2 and DAVID K. REILEY.2
U.S. Navy, Southwest Division Naval Facilities Engineering Command,
San Diego, California, USA; 2 University of Wisconsin, Institute of Environmental Studies, Madison, Wisconsin, USA. Patterns of recruitment
and mortality in mixed coast live and Engelmann oak woodlands in
southern California.
1

Two species are dominant in lower elevation oak woodlands of extreme

southern Californiathe widespread Quercus agrifolia (coast live oak) and
the much rarer Q. engelmannii (Engelmann oak). Concern about the future
of the rarer species is a special case of a more general concern about oak
woodlands. In this study we resampled plots in the Engelmann oak phase
of southern oak woodland originally sampled ten years earlier. We observed
increases in the saplings of both species on a subset of the plots, but the
increase was substantially greater for coast live oak. Previous studies have
shown that Engelmann oaks were less successful in seedling establishment
but had a higher survival into the larger size classes, apparently because
of greater ability to survive in gaps. In addition spatial analysis using ARCINFO proximity analysis revealed a significantly greater tendency for live
oak saplings to recruit in patches than Engelmann oaks such that live oak
saplings are predicted to be less likely to recruit effectively to the canopy.
The long term data suggest a more complex pattern of survival and growth
in which growth was greater in gaps, but a greater number of saplings were
found under the outer canopy of large trees than in either gaps or the inner
canopy. These new data suggest that factors allowing the coexistence of
these two species may vary over time and are probably more complex than
previously believed.
LAYMAN, CRAIG A.* and KIRK O. WINEMILLER. Texas A & M University, Department of Wildlife and Fisheries Sciences, 210 Nagle Hall,
College Station, TX, 77843-2258. Piscivore exclusion experiments predict impacts of commercial fishing on fish assemblages in tropical river
lagoons.
Mediating effects of over-harvesting top predators depends on a clear understanding of how species interactions propagate through ecological communities. Due to complex direct and indirect interactions among species,
such effects are especially difficult to predict in species-rich communities.

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199

Characteristics used to group species, and thus generalize ecological interactions, can aid in constructing predictive models in speciose food webs.
We have demonstrated how size could be used to predict a trait-mediated
response of multiple prey species (n.50) to piscivore (n.8) exclusion in
the Cinaruco River, a lowland neotropical river. Here we review results of
our small-scale experiments, and examine if they can be scaled up to predict larger-scale effects of commercial fishing. Lagoons (1-30 km2) are
dispersed throughout the Cinaruco River floodplain landscape, and many
are exploited by commercial netters. This results in a large-scale pattern in
which some lagoons support robust populations of large-bodied piscivores
(e.g., piranha and peacock cichlids), whereas others have had piscivore
populations greatly reduced. Fish assemblages in netted lagoons are dominated by relatively larger prey fishes (. 40 mm), including Moenkhausia
af lepidura and Bryconops caudomaculatus, the two species that responded
most significantly in piscivore exclusion experiments. Assemblages in unnetted lagoons are dominated by a group of smaller (, 40 mm) species,
none which were found to respond to piscivore exclusion. Results of extensive stomach content analyses (n.3,500) revealed prey fishes .40 mm
were the most common prey of piscivorous fishes (i.e., those piscivores
targeted by netters), providing additional evidence that observed differences are due to piscivore removal. Together, these results suggest (1) removal
of piscivores has significant cascading effects on fish assemblages, and (2)
prey size, as was demonstrated in exclusion experiments, can be used to
predict large-scale assemblage patterns in a speciose tropical river food
web.
LEACH, ANDREA D.* and CHRIS J. PETERSON. University of Georgia,
Athens, GA. Microsite vegetation and environmental differences in salvaged and unsalvaged wind-disturbed forest.
Intense winds are an important component of the disturbance regime in
eastern North American forests, disturbing thousands of hectares annually
and creating patches (or microsites) that are differentially favorable for
germination of seeds and survival and growth of seedlings and saplings.
As a result, the abundance, arrangement, and composition of microsites
may in large part determine patterns of regeneration. While natural wind
disturbance creates characteristic patterns of microsite abundance and composition due to treefall pits and mounds and downed tree crowns, salvage
logging creates distinct microsite types, such as slash piles and skid trails,
that may not have close natural analogs. We established 380 0.5 m2 circular
plots across salvaged and unsalvaged wind-damaged forest in which we
measured percent cover of herbaceous vegetation, height of tree seedlings
and saplings, soil moisture and temperature, and percent canopy openness
of five different microsite types (treefall pit, treefall mound, downed tree
crowns, bare soil areas, and patches of Vitis rotundifolia). We found that
percent canopy openness and soil temperature differed between post-windthrow treatments (ANOVA, p,0.001) with microsites in salvaged areas
having both more light and higher temperatures. Soil moisture and soil
temperature differed between microsite types (ANOVA, p,0.001). Soil on
treefall mounds had the lowest moisture levels and highest temperatures
contrasted with pits having the wettest soils with the lowest temperatures.
Canonical Correspondence Analysis using log-transformed environmental
variables demonstrated that soil moisture and temperature most strongly
influenced species composition of microsites. We conclude that microsite
type is an important factor in determining environmental conditions and
vegetation and may overshadow whether the disturbed forest was salvagelogged.
LEAKEY, ANDREW,1,* PHILLIP DAVEY,2 DAMIAN ALLEN,3 ALISTAIR ROGERS,4 EVAN DELUCIA,1 BERT DRAKE,5 RAMESH MURTHY6 and STEVE LONG.1 1 University of Illinois, Urbana, IL; 2 University
of Essex, Colchester, Essex, UK; 3 BASF Plant Science, Research Triangle
Park, NC; 4 Brookhaven National Laboratory, Upton, NY; 5 Smithsonian
Environmental Research Center, Edgewater, MD; 6 Columbia University
Biosphere 2, Oracle, AZ. How will leaf respiration respond to growth
under elevated [CO2] in three diverse tree canopies?
Terrestrial plant respiration is a large component of the global carbon cycle,
releasing 10 times more carbon per annum than fossil fuel combustion.
Some previous studies have shown an inhibition of leaf mitochondrial respiration under future elevated [CO2] (-18 %), which would be sufficient to

200

Abstracts

increase sequestration of carbon into biomass and partially offset rising

atmospheric [CO2]. However, the nature of the direct and indirect responses
of respiration to growth at elevated CO2 remains contentious. We used
high-resolution differential oxygen analysis to assess respiration without
the potential experimental artifacts arising from traditional infra-red gas
analysis of CO2 flux. We present data on the following diverse experimental
forest canopies that have developed under elevated CO2 in the field: Pinus
taeda at FACTS-1, Duke Forest, NC; Populus deltoides at Biosphere 2,
AZ; and Quercus myrtifolia & Quercus geminata at the Smithsonian opentop chamber facility, FL. We found: (A) no direct effect of elevated [CO2]
on respiration in any species at any time, but; (B) variation between 0 23 % greater respiration after long-term growth at elevated [CO2]. Further
data is presented assessing rates of respiration in canopies grown at elevated [CO2] in relation to leaf photosynthetic, carbohydrate and nitrogen
status.
LEAR, JENNIFER A.* and MARK J. BUTLER. Old Dominion University,
Norfolk, Virginia. Interactions between juvenile Caribbean spiny lobsters and Caribbean reef octopuses in the Florida Keys.
Juvenile Caribbean spiny lobster (Panulirus argus) and the Caribbean reef
octopus (Octopus briareus) may simultaneously compete for resources and
interact as predator and prey within hard-bottom habitats in the Florida
Keys, FL (USA). Lobsters chemically detect the presence of octopuses and
avoid shelters from which these cues emanate, thus altering patterns of
lobster shelter use. We conducted surveys of juvenile lobsters and octopuses occupying artificial shelters at 18 locations in the Florida Keys during
spring 2002 and 2003 and summer 2001, 2002 and 2003. Our findings
show that juvenile lobsters do not cohabitate in shelters with octopuses,
however there was no correlation between the numbers of octopuses and
juvenile lobsters on a site. We then used mesocosm experiments to investigate why the strong behavioral interaction between octopus and lobster
is not reflected in local population structure. First, we investigated whether
juvenile lobsters reach a size refuge from octopus predation. There was no
correlation between octopus weight and the size of juvenile lobsters that
they consumed; thus large juveniles were just as readily consumed as small
juveniles. Second, we examined the effects of alternative prey, lobster size,
the presence of lobster conspecifics, and initial residency on shelter competition between lobsters and octopuses. The presence of alternative prey
and conspecifics reduced predation on lobsters and their susceptibility to
displacement from a preferred shelter. Although octopuses are effective
predators of lobster and can alter their use of dens, the presence of alternative prey and lobster conspecifics mediates predation and shelter use,
which may explain why there is no relationship between lobster and octopus abundance in the field.
LEAVITT, PETER R.,1,* BRIAN F. CUMMING,2 JOHN P. SMOL,2 MEL
REASONER,3 REINHARD PIENITZ4 and DOMINIC A. HODGSON.5
1
University of Regina, Peter.Leavitt@uregina.ca, Regina, SK, Canada;
2
Queens University, Kingston, ON, Canada; 3 Mountain Research Initiative Coordination Office, Bern, n/a, Switzerland; 4 Universite Laval, Quebec, QC, Canada; 5 British Antarctic Survey, Cambridge, n/a, UK. Climatic control of UV radiation impacts on lakes.
UV radiation (UVR) damages most biota, yet little evidence exists for its
long-term impacts on natural ecosystems. Here, we used paleoecological
techniques at three low elevation lakes to show that ecosystem production
was depressed 10-fold by UVR during the first millennium of lake existence. Likewise, analysis of a lake near treeline showed that algal production declined 10- to 25-fold both early in the lake history and during the
last ca. 4000 years when inputs of UVR-absorbing dissolved organic matter
(DOM) declined despite constant nutrient levels since ca. 10,000 14C yr BP.
This rapid (-1.25%/yr) sustained (.600 years) suppression of ecosystem
production arose from directional climatic change that reduced DOM inputs
and occurred despite initial reservoirs of photoprotective DOM that were
typical of most boreal lakes. Hence, we conclude that many lakes may be
vulnerable to order-of-magnitude declines in production arising from future
climate-DOM-UVR interactions.

LEBOUTON, JOSEPH P.,* EDWARD J. LAURENT, MICHAEL B. WALTERS and JIANGUO LIU. Michigan State University, East Lansing, MI.
Regional and local effects of white-tailed deer herbivory on vegetation
structure and composition in northern hardwood forests.
Many studies of northern hardwood forests in the U.S. link high levels of
white-tailed deer herbivory with low seedling and sapling densities and
altered herbaceous community composition. We hypothesized that the juxtaposition of vegetation patches on the landscape is a critical factor affecting local white-tailed deer densities and their subsequent effect on the
vertical structure and species composition of forest plant communities. We
tested this hypothesis across a regional winter deer density gradient (0-20
deer/km2) using a network of 143 vegetation plots in a 500,000ha study
region in the Upper Peninsula of Michigan. Within each plot we sampled
herbaceous vegetation, three size categories of tree seedlings and saplings
(,0.25m tall, 0.25-1.5m tall, and .1.5m tall-10cm dbh), and canopy composition. We compared forest vertical structure and community composition
data to regional white-tailed deer winter population indices from 1956 to
present, controlling for forest management history. We also compared present vegetation to landscape composition indices around each plot, including edge density, landscape connectivity, and distance to key white-tailed
deer habitat types. Seedling and sapling density and species composition
varied in response to regional white-tailed deer winter population density,
forest management history, and the landscape composition around each
stand. Plots within high winter deer density regions and local landscapes
favorable to high winter deer densities (e.g., stands near winter deer yards)
exhibit less vertical structure and fewer seedlings, saplings, and canopy
recruits than plots within low winter deer density regions and/or in local
landscapes that are far from key deer habitat. In contrast, understory community composition showed little response in species richness but a large
response in community evenness to regional deer density and local landscape composition. Species avoided by white-tailed deer (e.g., mat-forming
Carex species) had much higher dominance within high winter deer density
regions, regardless of local landscape composition.
LEE, EUN JU,* IN SUNG KIM and YONG-JOO CHO.
ejlee@plaza.snu.ac.kr, Seoul National University, Seoul, Seoul, S. Korea.
Macronutrient input by pollen in two pine stands in south-east Korea.
This study was carried out to examine macronutrient input by pollen in
two naturally regenerating pine stands in south-east Korea. Durham gravity
pollen collectors were used to measure pine pollen deposition and macronutrients in pine pollen were analyzed. In 1998, pine pollen deposition
began just before April 18 and lasted for about two weeks. Two species of
pine differed in the timing of polen release, with P. rigida beginning pollen
release two days earlier than that of P. densiflora. Total pine pollen deposition was a bit different over the two sampling sites; 27.5 kg/ha in the
mature stand, 17.7 kg/ha in the young stand. The values for nutrient deposition by pine pollen are 549 g/ha N, 78 g/ha P, 240 g/ha K, 45 g/ha S,
22 g/ha Mg in the mature site and 353 g/ha N, 51 g/ha P, 151 g/ha K, 27
g/ha S, 14 g/ha Mg in the young site, suggesting that nutrients in pine
pollen contributes to forest nutrient cycling. Total basal area of mature pine
stand was 20 times higher than that of young pine stand but pine pollen
deposition was only 1.5 times higher in the mature pine stand. Macronutrient deposition by pine pollen is concentrated temporally in spring. Although the annual contribution of nutrient mass by pollen is small compared to that of litterfall, the rapid turnover rate of pollen nutrients combined with episodic deposition suggests that pollen may play a disproportionate role in temperate pine forest nutrient cycling.
LEE, SARAH C.1,* and BRIAN R. SILLIMAN.2 1 University of North
Carolina-Chapel Hill, Chapel Hill, NC; 2 Brown University, Providence, RI.
Competitive displacement of the pulmonate snail Melampus bidentatus
by a sympatric salt marsh gastropod.
Although the importance of biological interactions in structuring salt marsh
plant communities has recently been established, few studies have explored
the role of interspecific interactions in controlling salt marsh animal distributions. In southern marshes, the pulmonate snail Melampus bidentatus
is most abundant in Juncus roemerianus in the high marsh, while another
gastropod, Littoraria irrorata, dominates the lower marsh zone (dominated

by short-form Spartina alterniflora). Both snails feed primarily on dying

marsh grass, fungi, and microalgae, suggesting that exploitative competition with Littoraria could limit the lower distribution of Melampus. Alternatively, Melampus may not be abundant in the short Spartina zone because there is less plant cover and thus elevated desiccation stress. We
tested these competing hypotheses by manipulating Littoraria presence and
solar radiation levels in short Spartina over two years in two marshes.
Reducing light levels and removing Littoraria induced Melampus immigration and significantly increased local densities. Littoraria removal had
a much greater effect than shading, suggesting that snail presence, not
desiccation stress, is the primary factor limiting Melampus from the short
Spartina zone. Conversely, addition of Littoraria to the Juncus zone, where
Melampus naturally occur in high densities, induced emigration and significantly decreased local Melampus densities. Further experiments suggested that Melampus actively avoids Littoraria because grazing periwinkles significantly reduce: 1) available detritusdecreasing Melampus
growth rates, 2) Spartina stem densityincreasing desiccation stress, and
3) successful eggmass attachment using cues present in Littoraria mucus.
Our results demonstrate that periwinkles can competitively exclude Melampus from highgrowth habitats and suggest that patterns of snail zonation
in these marshes are largely determined by biological interactions.
LEFEVRE, KARA L.* and F. H. RODD. University of Toronto, Toronto,
Ontario, Canada. Influence of habitat disturbance on bird-mediated
seed dispersal in a neotropical rainforest.
The effects of anthropogenic habitat disturbance on ecosystem processes
are not well documented. We are investigating the influence of forest clearing and plantations on seed dispersal patterns in the lower montane rainforest of Tobago, West Indies. Data concerning changes in the community
of avian dispersers, diet composition, and foraging ecology in disturbed
versus intact rainforest will be presented, along with results of experiments
to quantify fruit removal rates and patterns of seed deposition. Implications
of these changes for seed dispersal and plant demography will be discussed.
LEFFLER, A. JOSHUA,* MICHAEL S. PEEK, RONALD J. RYEL and
MARTYN M. CALDWELL. Ecology Center, 5205 Old Main Hill, Utah
State University, Logan, Utah, USA. Hydraulic redistribution: The role
of shoots.
Hydraulic redistribution, previously termed hydraulic lift, is the movement
of water from soils of high water potential to soils of low water potential
through the root system of a plant. Hydraulic redistribution has been demonstrated in over 80 taxa from desert to tropical ecosystems and is thought
to be a purely physical process that can occur whenever active portions of
root systems are in soils of different water potential. Hydraulic redistribution likely takes place when stomata are closed and the soil-plant-air
continuum is decoupled from the atmosphere. Given this, aboveground
plant tissue should play little role in redistribution. To test this hypothesis,
we injected a 2H label into the lower rooting zone of a Bromus tectorum
(cheatgrass) monoculture and examined d2H in the upper soil. Our injections were performed twice, once in the late growing season for B. tectorum
and again after the aboveground tissue was completely senesced. During
both sampling periods the aboveground tissue was either left intact or removed. We found a significant difference between the water signature in
upper soils collected from our deep injection plots and background samples
of the soil water for both clipped and unclipped plots in the early sampling
period and for the unclipped plots in the later sampling period. In the plots
that differed significantly from background samples, 4 to 6% of the water
in the upper soil was derived from the water we injected into the deep
rooting zone. The similar fraction of upper soil water derived from the
deep injections (ca. 6%) in clipped and unclipped plots during the early
sampling period demonstrate that aboveground tissue is not necessary for
hydraulic redistribution to occur. Although we observed lower quantities
of redistributed water during the later sampling date, our data do suggest
that roots of dead plants can continue to serve as conduits for water movement through the soil.

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201

LEHMAN, ELIZABETH, ALAN C. LOVE, ELLA L. INGRAM* and

KELLY MYER POLACEK. Indiana University - Bloomington, Bloomington, IN. An ecological example of incorporating inquiry into non-laboratory classrooms: Exploring organisms.
Inquiry approaches to teaching science are well established as effective
techniques to engage students in the process of science and increase student
learning. We most often think of inquiry as typical of laboratory classroom
setting, although inquiry approaches are not limited to this context. We
describe methods of incorporating inquiry into non-laboratory classrooms,
using an ecology activity as an example. We focused our activity on having
students ask and evaluate questions. We observed class meetings when the
activity was scheduled and when a non-inquiry activity was scheduled, and
recorded classroom dynamics using a tool designed to assess inquiry-focused behaviors. During the inquiry activity, we observed a greater diversity of student behaviors in the classroom as compared to the non-inquiry
activity. Also, we observed more inquiry-appropriate behaviors of the instructor and an increase in students practicing inquiry skills during the
activity. The two most dramatic differences between inquiry-based and
non-inquiry-based class meetings were 1) students passively listening to
the instructor half as frequently during the inquiry activity, and 2) students
asking each other questions eight times more frequently during the inquiry
activity. The major lesson learned from the assessment of this activity is
that students can successfully practice inquiry in a non-laboratory classroom. We describe refinements of the presented teaching activity and summarize strategies for incorporating inquiry into discussion sections or learning groups.
LEIBOLD, MATHEW A.,1 BERNADETTE PINEL-ALLOUL,2 GINETTE
METHOT2 and CHRISTOPHER F. STEINER.3 1 Section of Integrative Biology, University of Texas - Austin, Austin, Texas, U.S.A.; 2 Departement
de Sciences Biologiques, Universite de Montreal, Montreal, Quebec, Canada; 3 Department of Ecology, Evolution, and Natural Resources, Rutgers
University, New Brunswick, New Jersey, U.S.A. Effects of ecosystem
productivity on temporal species-turnover and biodiversity in plankton.
Previous work has shown that ecosystem productivity can affect biodiversity in different ways at different spatial scales. At the local level there is
often a unimodal (humped) relationship with productivity whereas the effects are monotonically increasing at larger regional scales. Models of
community assembly indicate that one important mechanism could involve
greater likelihood of heteroclinic invasion cycles at high productivity. This
occurs as food chains build up to support larger populations of organisms
at higher trophic levels that become invasible by competitors at lower trophic levels resistant to the resident predator. We examined species turnover
in the composition of zooplankton from year to year to see if we could
find a relationship with lake productivity in two different regional biotas
in Canada. Consistent with our assembly models, we found higher temporal
turnover of zooplankton at higher productivity but could not assess if this
was due to collapsing food chains due to lack of sufficient data on other
trophic levels. Nevertheless these results indicate that temporal turnover
can be associated with productivity and may explain why diversity patterns
show distinct scale-dependent patterns with productivity.
LEICHT, STACEY A.,* JOHN A. SILANDER and CYNTHIA S. JONES.
University of Connecticut, Stacey.Leicht@uconn.edu, Storrs, CT. Comparative ecology of an invasive bittersweet species and its native congener.
A powerful approach to the understanding of invasive species comes
through comparing invasive species with native congeners. In this way, we
may better understand traits that provide invasive species an advantage in
the landscape. Celastrus orbiculatus was introduced into the eastern United
States from East Asia in the late 1800s as a horticultural plant and has
become highly invasive in the mid to late 1900s. The native C. scandens
ranges from the eastern part of the United States to the Midwest, and is
on the decline in the Northeast, where some states list it as threatened. We
compared germination, development, growth and life-history traits of these
species grown from seed in competition at three densities in a common
garden. Our factorial design included two soil types (varying both in pH

202

Abstracts

and texture), two light levels (shaded and unshaded) and the opportunity
to climb (staked or unstaked). In the first year of growth, C. orbiculatus
had greater relative growth rates, a higher tolerance of shade and later leaf
abscission than C. scandens. C. orbiculatus also showed greater responses
in several morphological measures of growth than C. scandens in high
density treatments. These results may help explain the decline of C. scandens in the Northeast.
LEISHMAN, MICHELLE.* Macquarie University, Sydney, NSW, Australia. Predicting invasiveness from plant traits: the role of disturbance
type.
One of the major threats to global biodiversity is invasion by exotic plant
species. Invasive plants can affect ecosystem structure and function by
reducing native species richness and modifying ecosystem processes via
altered water or fire regimes. Many studies have attempted to relate invasiveness to plant traits, with limited success. It has become increasingly
apparent that the interaction between the invader and the target community
is critical. I present data on a wide range of traits of native and invasive
species from sites subject to different disturbance types: nutrient and water
enrichment, nutrient enrichment, physical disturbance and grazing. Specific
leaf area (SLA) of invasive exotic species was consistently higher than
SLA of non-invasive exotic and native species, regardless of disturbance
type. In sites subject to physical disturbance or grazing, invasive species
were more likely to be herbs and grasses, while in sites subject to nutrientenrichment, exotic species were more likely to be climbers or trees. There
were no significant differences in seed mass between native and invasive
species, for any disturbance type. Differences in dispersal mode between
natives and invasives depended on disturbance type. Thus different plant
traits contribute to exotic species success under different disturbance types,
with the exception of SLA which was consistently higher for invasive species. High SLA may be associated with the superior ability of invasive
species to capture and maintain space, with potential for rapid growth being
a key component. I present data on leaf trait (SLA, Nmass, Pmass) relationships
of invasive species compared with natives and show that the relationships
have a common axis but that invasives have consistently higher values of
these traits than native species. This suggests that there may be consistent
functional relationships of invasive plant species.
LEMASSON, BERTRAND H.,1,* JAMES W. HAEFNER,1 MARK BOWEN2 and KATHY FRIZZELL.2 1 Utah State University, birdy@biology.usu.edu, Logan, Utah, U.S.A.; 2 Bureau of Reclamation, Denver, Colorado, U.S.A. Parameter estimation for the behavioral algorithms of a fish movement model.
An individual-based movement model was statistically validated using extraction efficiency experiments conducted in the Tracy Fish Collection Facility (TFCF) in Tracy, CA. This work is part of the ongoing conservation
efforts of the state of California and the Bureau of Reclamation in reducing
the loss of endangered and economically important fish species in the lower
San Joaquin river. The model incorporates individual fish behavior and
swimming physiology with simulations of fluid flows in a physical model
of the TFCF. Previous work used existing literature values for many behavioral parameters. Predicted extraction efficiency for 6 common species
in the TFCF ranged between 0.66 and 0.83; sensitivity analysis quantified
the effect of initial fish energy level and fish avoidance behavior on predicted efficiency. Model predictions were within the experimental error for
most species, although statistical variability is high for measured efficiency.
To more precisely test and parameterize the model, new validation and
parameter estimation experiments were conducted using digital video taping of fish behavior in an experimental flume constructed by the Bureau
of Reclamation, Denver, Colorado. The flume is a plexiglas chamber with
an array of vertical louver slats and controls to vary bypass velocity ratio.
We tested Rainbow Trout(mean size 98 mm) and Splittail(mean size 151
mm) using two approach velocities and louver configurations. Using images recorded every 0.1s and video tracking software, we estimated parameters of fish obstacle avoidance behavior from frequency distributions of
turning angles and accelerations. Avoidance behavior is a change in the
variance of these two parameters between time steps. To investigate the
effect of different types of obstacles on fish behavior, we compared avoidance behavior near the louver array and near the opposite solid wall. We

report these values and the results of extraction efficiency simulations based
on new flume fluid dynamics and the refined parameter estimates.
LENNON, JAY T.* and LIZA E. PFAFF. Dartmouth College, Hanover, NH.
Microbial constraints on the flow of terrestrial subsidies in lake ecosystems.
Terrestrial ecosystems deliver large quantities of dissolved organic matter
(DOM) to nearby lake ecosystems. The energetic importance of these subsidies for higher trophic levels remains unclear, but should be determined
in part by the microbial processing of DOM. In this study we investigated
how varying sources and supplies of terrestrial DOM influence microbial
metabolism. First, in laboratory regrowth experiments, we found that gross
bacterial productivity (GBP) was higher on beech-, hemlock-, and oakderived DOM than pine-, maple-, and birch-derived DOM. These differences could be explained in part by variation in C : P ratios between DOM
sources. In contrast, bacterial respiration (BR) was less affected by different
DOM sources. Second, using field mesocosms, we found that DOM supply
was also important in regulating bacterial metabolism. For example, net
bacterial productivity (NBP 5 GBP - BR) was highest when subsidies were
added as single, large pulses instead of smaller, but more frequent DOM
inputs. It was only under these pulsed conditions that NBP was affected
by different DOM sources. Our results suggest that the importance of terrestrial subsidies in lake ecosystems may be influenced by forest cover and
hydrologic regimes that govern DOM loading rates.
LENSING, JANET R.* and DAVID H. WISE. University of Kentucky,
lensingjr@yahoo.com, Lexington, KY. Initial response of leaf-litter arthropods to altered rainfall predicted by models of climate change.
Activities of arthropods in leaf litter affect rates of litter decomposition and
nutrient cycling. The community of arthropods and microorganisms is
strongly affected by abiotic factors, particularly changes in moisture. Thus,
changes in rainfall predicted to occur with global climate change will likely
affect the functioning of this component of the forest ecosystem. Models
predict increased rainfall and increased severity of drought as two outcomes
of climate change. We constructed rainout shelters to cover 8 large fenced
plots randomly assigned to either a high- or low-rainfall treatment (irrigated
with extremes approximately +/- 30% of long-term mean). This rainfall
regime mimics extreme high and low periods of rainfall during the past
107 years for Kentucky. We conducted the rainfall manipulations from June
through November 2002. Arthropods were sampled by sifting the leaf litter
in October 2002. Samples revealed that total arthropod densities were ca.
2x higher in the high-rainfall plots (F1,4 5 8.05, P 5 0.047) due to an
approximate doubling of spider densities (F1,4 5 42.7, P 5 0.003). Collembola, mites, millipedes, centipedes, ants, and other insects did not differ
significantly between treatments (all P . 0.21). These results are intriguing
since differences have emerged after only 6 months of manipulations but
only in one of the major arthropod groups. Undoubtedly more patterns will
be revealed as the experiment continues.
LENTILE, LEIGH B.,1,* FREDERICK W. SMITH2 and WAYNE SHEPPERD.3 1 Colorado State University, lentile@cnr.colostate.edu, Ft. Collins,
CO; 2 Colorado State University, skip@cnr.colostate.edu, Ft. Collins, CO;
3
USDA Forest Service, Rocky Mountain Research Station, wshepperd@fed.fs.us, Ft. Collins, CO. Fire effects and fire scar formation following the Jasper Fire, Black Hills National Forest, SD.
Fire histories in ponderosa pine (Pinus ponderosa var. scopulorum) forests
are often reconstructed from fire scar and tree origin information. Fire frequency and fire size are inferred from dates of fire occurrence as recorded
as scars on individual trees. However, little is known about rates of fire
scar formation in relation to fire behavior and stand conditions. We examined fire scar formation and ponderosa pine regeneration patterns in
areas of different fire severity 2-years following the large (40,000 ha.)
Jasper fire of 2000 in the Black Hills of SD. We measured rates of mortality, fire scar formation and ponderosa pine regeneration in areas of low,
moderate and high fire severity. We examined 2600 live trees within areas
of low and moderate fire severity for evidence of fire scar formation. Dead
cambium on a significant portion of tree circumference where the tree had
live cambium and a live crown was taken as evidence of fire scar formation.

Two years post-fire, tree mortality was 6%, 24 % and 100% in areas of
low, moderate and high fire severity. Fire scar formation was detected in
23% and 43% of surviving trees in low and moderate severity areas. Regeneration densities were 7000 seedlings/ha in areas of low severity,
40% lower in moderate areas, and we observed no regeneration in high
fire severity areas. Fire scar formation and regeneration patterns following
this contemporary fire provide a context for interpretation of historic fire
regimes based on dendrochronological and historical records. We propose
that the landscape pattern of post-fire cohorts and fire-scarred trees that
resulted from the Jasper fire is indicative of mixed severity fire regime.
LENTZ, AMANDA J.1,2 and KAREN M. KESTER.1 1 Virginia Commonwealth University, Richmond, Richmond, VA, USA; 2 Virginia Polytechinical and State University, Blacksburg, VA, USA. Postemergence experience impacts fitness of the insect parasitoid Cotesia congregata Say
(Hymenoptera: Braconidae).
Although learning is hypothesized to improve fitness, the direct effects of
experience on ovipositional decision-making have not been tested. We investigated the effects of postemergence experience with host or novel
plants on clutch size and sex ratio allocation by Cotesia congregata Say.
Wasps that emerged on one of two host plants produced clutches with a
higher proportion of females when offered the same plant at oviposition.
Wasps that emerged on one of two novel plants oviposited larger clutches
in the presence of a relatively unattractive novel plant, irrespective of postemergence experience, and produced clutches with higher proportions of
females on either plant than wasps with no plant experience. Differential
effects of postemergence experience on clutch size and sex ratios may
reflect prior adaptation to host plants and facilitate local adaptation to plants
utilized by abundant hosts.
LEONARD, NORMAN E.* University of New Orleans, New Orleans, LA,
70148. Invasive Sapium sebiferum detritus breaks down five times faster than detritus from native Acer rubrum.
Chinese tallow (Sapium sebiferum) is an invasive exotic tree found in riparian zones in the southeastern United States, where it may out-compete
native red maple trees (Acer rubrum). The decay rate of Chinese tallow
leaves in streams has not been reported although they may comprise much
of the leaf material entering a water body where the species is dominant.
Information on leaf breakdown is necessary to assess the impacts of altered
allochthonous inputs on aquatic faunal communities. During the summer
of 2002 I compared the decay rate of Chinese tallow leaves and red maple
leaves in streams. Air-dried leaves were bundled into 40 leaf packs of
known mass (20 of each species) and placed in a stream on 25 July 2002.
Four leaf packs of each species were removed from the stream after 3.5,
7, 28, 35, and 56 days. The leaf material was rinsed, dried and weighed to
assess mass loss rates. The processing coefficients (k) for the two species
were significantly different (p50.000), with values of k5-0.0166 for Chinese tallow and k5-0.0033 for red maple. Aquatic invertebrates rinsed from
the leaf packs were identified and counted. Initially S. sebiferum leaves
supported greater abundances of aquatic invertebrates than A. rubrum
leaves; however, as tallow leaves lost mass more rapidly, only maple leaves
supported invertebrate assemblages by the end of the study. Streams with
riparian zones invaded by Chinese tallow may experience significant changes in the quantity, quality, and persistence of allochthonous resources, and
these changes may affect the diversity and abundance of in-stream consumers.
LESLIE, HEATHER.* Oregon State University, leslieh@science.
oregonstate.edu, Corvallis, OR. Environmental influences on fitness in
the intertidal barnacle Balanus glandula.
In recent decades, ecologists have documented variation in recruitment
among local populations, and attributed it to a number of mechanisms at
multiple scales. Variation in reproduction and thus larval supply are necessary prerequisites for variation in recruitment, but have received considerably less attention. Nonetheless, these population parameters have important implications for population and community persistence, as well as
for management and conservation. To investigate the relative importance
of micro- and mesoscale environment on growth and reproduction in the

Abstracts

203

intertidal barnacle Balanus glandula, I manipulated conspecific density at

two rocky intertidal sites in Oregon, USA. The sites were chosen to bracket
a documented gradient in oceanographic conditions, which in turn has led
to variation in intertidal community structure and dynamics. At each site,
I cleared three plots on wave-exposed, mid intertidal benches (+ 2m). Once
barnacle recruitment occurred, one plot of each set was randomly selected
as the thinned treatment and the other as natural. For the next eight months,
I maintained the thinned treatment. The mean number of barnacles/cm2 in
the natural plots at the low productivity site was significantly higher than
at the high-productivity site, due to variation in recruitment and/or survival
of juveniles. On average, thinned barnacles at the high-productivity site
grew most rapidly and attained the largest size. Individual allocation to
reproduction was greater at the high-productivity site. On a population
scale, the proportion of brooding individuals/100 cm2 was negatively related to barnacle density. This resulted in a unimodal association between
total brood mass/cm2 and conspecific density, suggesting that barnacles
living at intermediate densities may contribute disproportionately to the
regional larval pool.
LETOURNEAU, DEBORAH K.* and SARA G. BOTHWELL. University
of California, Santa Cruz, CA. Bottom-up effects on redwood understory
vegetation in natural and logged light gaps.
Redwood forests are Californias most unique and majestic natural habitats.
They are also severely light-limited communities, with very low plant species diversity in old growth forests. We hypothesized that plant and arthropod diversity would be greatest in medium-sized canopy gaps, which represent natural levels of disturbance, and that lower biodiversity would occur
in both small canopy gaps and selectively logged forest gaps. Comparing
five replicates of each gap size, we measured light availability, plant species
richness, and arthropod species richness. Treefall gaps of moderate canopy
gap size had significantly higher plant and arthropod diversity. Preliminary
analyses of three trophic levels suggest that light drives a bottom-up diversity cascade.Although consistent with the intermediate disturbance hypothesis, general patterns of predatory arthropod abundance and diversity
also support the notiont that trophic interactions may feed back to limit
herbivores and plants in larger gaps with the highest light levels.
LEUNG, BRIAN,* JOHN DRAKE and DAVID M. LODGE. Dept. Biol.,
Univ Notre Dame, Notre Dame, IN, USA. Predicting invasions: Propagule pressure and the gravity of the Allee effect.
Invasions by non-indigenous species impose large environmental and economic costs. In order to prevent invasions and target monitoring efforts
most effectively, we need to forecast locations at the greatest risk of new
invasions. To accomplish this, we need to estimate propagule pressure (inoculum size) and consider population processes, possibly including Allee
effects. Here, we develop a method to estimate the probability of establishment, based on survival analysis and maximum likelihood techniques.
We demonstrate theoretically the validity of this approach, considering environmental heterogeneity, estimation error, and non-linearity. We then apply this method to zebra mussel (Dreissena polymorpha) invasions of
Michigan inland lakes. We fitted our model using presence-absence data
between 1992-1996 and propagule pressure estimates from gravity models.
Using our fitted model, we estimated the probabilities of establishment and
demonstrated that Allee effects were present in the zebra mussel system.
We validated our model using invasion data from 1997-2001, which was
not used to parameterize the model. Using the validation data set, we correctly predict up to nine times as many invasions as the null (random)
model. Further, the Allee model assigned average probabilities 4.5 times
higher for lakes that became invaded compared to uninvaded lakes, whereas
the non-Allee model only predicted probabilities two times as high. Thus,
our model demonstrates the importance of considering the Allee effect and
improves predictions of future invasions.
LEVINE, JONATHAN M.1,* and MARK REES.2 1 UCLA, Los Angeles,
California, USA; 2 Imperial College at Silwood Park, Ascot, Berkshire, SL5
7PY. Annual rainfall fluctuations favor rare plant persistence in invaded habitats.
From a traditional conservation biology perspective, environmental fluctuations are regarded as detrimental to persistence, reducing long-term av-

204

Abstracts

erage growth rates and increasing the probability of extinction. By contrast,

numerous coexistence models from community ecology suggest that for
species with dormancy, environmental fluctuations may be essential for
persistence in competitive communities. We developed stochastic population and community models to examine the effects of annual rainfall fluctuations on the persistence of endangered annual plants competing with
exotic grasses on the California Channel Islands. We found that in multispecies models, despite both types of species being favored by the same
high rainfall environments, differences in germination biology and dormancy caused the rare annual plants to perceive variation in their environment differently than their exotic grass competitors. This allowed the rare
plants to benefit from annual fluctuations in the environment, yielding predictions contrary to the prevailing ideas for their management. Moreover,
persistence was maximized in systems with negative temporal autocorrelation and a relatively low probability of a wet year. Results suggest that
the yearly rainfall fluctuations in southern California may be essential to
the persistence rare species in exotic plant dominated habitats.
LEVINE, MIA T.* and KEN N. PAIGE. University of Illinois at UrbanaChampaign, Urbana, IL. Direct and indirect effects of extreme drought
on compensation following herbivory in Ipomopsis aggregata (Polemoniaceae).
Compensation following herbivory is now appreciated as an important element of plant defense; however, variation in compensation under naturally
stressful conditions has yet to be evaluated directly. During Arizonas worst
drought on record, a typically overcompensating population of Ipomopsis
aggregata severely undercompensated through female function. We tested
two factors that potentially contributed to this atypical pattern of undercompensation: 1) The direct effect of extreme water stress on the capacity
to compensate (following the typical single bout of herbivory) and 2) The
indirect effect of water-stress on compensation through drought-associated
ungulate browsing. We assessed the direct effect of water stress using a 2
x 2 fully factorial experiment with treatments: +/- supplemental water, +/clip (simulating the typical single bout of herbivory). We found a significant clip x water interaction for female fitness, suggesting that water limits
compensation through female function. Simulated herbivory had a larger
detrimental effect on female fitness in the nonwatered plants than on the
watered treatment plants. Moreover, watered plants equally compensated
(similar seed production in unclipped and clipped treatments) while waterstressed plants undercompensated (fewer seeds in clipped treatment)
through female function. In the second experiment, we examined the indirect effect of drought on compensation through ungulate browsing by
comparing 1) unbrowsed plants (caged), 2) clipped plants (simulating typical single bout of herbivory, caged), and 3) unprotected natural controls
(open to natural browsing levels). The natural controls suffered unprecedented levels of herbivory, resulting in severe undercompensation. We conclude that although water was an important limiting factor for compensation during the drought year, the overriding factor determining fitness
achieved through compensation was the indirect effect of drought
drought-associated ungulate browsing. Implications for plant-animal interactions under future climate change are discussed.
LEVRI, EDWARD P.,* CAROLYN ITLE, SHANE LUNNEN, JUSTIN
WINTERS and ROBERT PLATT. Division of Mathematics and Natural
Science, Penn State - Altoona, 3000 Ivyside Park, Altoona, PA, USA. The
effect of a trematode parasite on the response of a snail to light and
gravity.
Parasites often influence the behavior of their hosts in ways that increase
the transmission rate of the parasite. The New Zealand snail, Potamopyrgus
antipodarum, is commonly infected with the trematode Microphallus sp.
Infected snails must be eaten by waterfowl for transmission of the parasite
to occur. Previous studies from Lake Alexandrina, South Island, New Zealand, have demonstrated that infected snails are more exposed to predators
on top of rocks in the early morning, when waterfowl are feeding, than
uninfected snails. Infected snails then move to safer locations beneath rocks
when non-host predators (fish) begin to feed later in the day. The mechanism by which the parasite alters behavior is not clear. These experiments,
conducted in the field and in the lab, were designed to determine the effect
of the parasite on specific behaviors. Here we examine the geotactic and

phototactic behaviors of infected and uninfected snails. Snails were placed

in behavioral arenas. In the geotactic experiments the arenas were moved
into a vertical position, and snails were collected after four minutes and
separated by whether they moved up or down. In the phototactic experiments, snails responded to a directional light source and were separated
after four minutes by their angle of orientation. All snails were measured
and dissected to determine gender, brooding condition, and parasitism. All
groups of snails were found to be positively geotactic, however, infected
snails responded significantly less than any other snail group. Snails were
also found to be weakly positively phototactic, however, there were not
enough infected snails to determine if the parasite influenced this behavior.
These results may help to explain how the parasites influences the behavior
or the snail in a time-specific manner.
LEWIS, DANNY* and ROBERT F. DENNO. University of Maryland, College Park, Maryland, US. Tidal disturbance decouples predator-prey
interactions and promotes outbreak dynamics.
Physical disturbance has been proposed as a factor mediating the effects
of predators on herbivore populations. This study assessed the differential
impact of tidal inundation on the spatial distributions of arthropod predators
and prey and the possibility that such disturbance creates temporary spatial
refuges for herbivores and promotes outbreaks. We examined this issue on
a mid-Atlantic salt marsh where planthoppers and their invertebrate predators are the dominant players in the terrestrial food-web. Planthoppers
often reach outbreak densities in low marsh habitats characterized by greater tidal inundation. We hypothesized that three of the most abundant predators of planthoppers, the lycosid spider Pardosa littoralis, the sheet-web
building spider Grammonota trivitatta, and the mirid egg predator Tytthus
vagus, would be more adversely affected by rising tidewaters than are
planthoppers. We measured densities of planthoppers and predators during
different phases of the tide cycle at different elevations within and across
habitats. Prey were most abundant in low-elevation marsh habitats and their
distribution was unaffected by tidal inundation. In contrast, all three predator taxa shifted their distribution toward higher elevations during extremely high tides. Spatial shifts in predator distribution created a potential refuge for prey at low elevations, where indeed prey outbreaks are most frequently observed. Thus, even temporary refuges created by tidal disturbance can have important consequences for the coupling of predator-prey
interactions and outbreak dynamics of insect herbivores.
LEWIS, DAVID B.,1,2,* LINDA B. STABLER3 and CHRIS A. MARTIN.3
Center for Environmental Studies, Arizona State University, Tempe, AZ,
USA; 2 Biology Department, Arizona State University, Tempe, AZ, USA;
3
Plant Biology Department, Arizona State University, Tempe, AZ, USA.
Ecological stoichiometry of horticulture: Consequences of pruning and
water for plant nutrient use efficiency.
1

We used an experiment to test the hypothesis that factors encouraging plant

growth reduce nutrient use efficiency (NUE), measured as carbon : nitrogen
(C:N) and carbon : phosphorus (C:P) ratios. In an arid, urban ecosystem
(Phoenix, AZ), replicate individuals of both Texas sage (Leucophyllum frutescens) and oleander (Nerium oleander) were treated with high vs. low
irrigation crossed with no pruning vs. pruning once per 6 weeks. Foliar
tissue chemistry was measured seasonally, including damp, cool and dry,
hot periods. From our hypothesis, we predicted that (i) NUE is reduced by
pruning, (ii) NUE is reduced by irrigation, (iii) low-water conditions offset
the reduction of NUE caused by pruning, and (iv) any irrigation X pruning
interaction is only evident during drought. Prediction (i) was partially supported. Pruning reduced C:N of Texas sage and C:P of oleander. Prediction
(ii) was fully supported, as irrigation reduced both nutrient ratios of both
species. Prediction (iii) was also supported. The reduction in Texas sage
C:N caused by pruning was offset by seasonal drought, and the reduction
in oleander C:P caused by pruning was offset by withholding irrigation.
Prediction (iv) was supported by oleander. For this species, the reduction
in NUE caused by pruning could be offset by withholding irrigation, but
this phenomenon was more pronounced during the natural drought condition of July than during the moist monsoon and winter rain period. Because
varied horticultural practices are widespread throughout urban ecosystems,
these plant-level modifications of NUE may profoundly effect ecosystem
level material cycling.

LEWIS, GREGORY,* HOLLY GARRETT, BRANNON ANDERSEN and

KENNETH SARGENT. Furman University, Greenville, SC, USA. Concentrations and fluxes of dissolved nitrogen and sulfur in montane forested watersheds in South Carolina.
Because of rapid urbanization, atmospheric deposition of nitrogen and sulfur to forested ecosystems in the southeastern United States may increase
in coming decades. In anticipation of such changes, the purpose of our
study was to establish baselines for stream chemistry in forested watersheds
in northwestern South Carolina. The specific goals of our study were (1)
to determine spatial and temporal variability in dissolved nutrient concentrations in montane forested watersheds in South Carolina, and (2) to estimate annual effluxes of dissolved nutrients from one of those watersheds.
We analyzed nutrient concentrations in stream water from four watersheds
during June 2002-January 2003. Concentrations of total dissolved nitrogen
(TDN), nitrate-nitrogen, and dissolved organic nitrogen (DON) ranged from
0.05-0.30, ,0.01-0.10, and ,0.01-0.28 mg N/L, respectively. Nitrate-N
concentrations exceeded DON concentrations in only about half of all samples. Sulfate-S concentrations ranged from 0.31-0.74 mg/L. In all samples,
sulfate contributed more to total anion charge than did nitrate. Over the
period June 2001-January 2003 in the forested Middle Saluda River watershed, both nitrate-N and DON concentrations were highest during the
growing season and lowest during the dormant season. Sulfate-S concentrations showed the opposite pattern. We estimate that the watershed lost
about 1.1 kg TDN ha-1 yr-1 and 2.5 kg sulfate-S ha-1 yr-1 in stream water.
Preliminary budget calculations suggest that the watershed retains about
75% of the atmospheric N deposition and about 55% of the atmospheric
S deposition (wet plus dry in both cases) it receives. In conclusion, dissolved N and S concentrations in streams of forested watershed in South
Carolina are low in comparison to regions receiving high levels of atmospheric N and S deposition. At present, forested watersheds in South Carolina apparently have the capacity to retain much of the N and S they
receive by atmospheric deposition.
LEWIS, J D.,1,* ANDREW MARTIN,1 GREG TURNER,1 JEANNINE
RICHARDS,1 KEVIN L. GRIFFIN2 and DAVID T. TISSUE.3 1 Calder Center, Fordham University, Armonk, NY; 2 Lamont-Doherty Earth Observatory, Columbia University, Palisades, NY; 3 Texas Tech University, Lubbock, TX. Plant age and nitrogen supply interactively affect photosynthetic responses to flowering in Xanthium strumarium.
The transition from vegetative to reproductive growth in annual plants may
alter net photosynthetic rates through changes in the relative balance between carbohydrate source and sink tissue. However, these effects may be
mediated by nitrogen supply and by plant age at flowering. In addition to
the direct effects of nitrogen supply on net photosynthetic rates, both nitrogen supply and plant age at flowering may indirectly affect net photosynthetic rates through effects on source:sink balance. In this study, we
examined the interactive effects of nitrogen supply and plant age at flowering on photosynthetic responses to the transition between vegetative and
reproductive growth. Four cohorts of Xanthium strumarium were planted
at 14-d intervals and grown at one of three nitrogen supplies. Plants were
simultaneously induced to flower by switching the photoperiod from 18 h
to 13 h beginning 70 d after germination of the first cohort. Plant size
significantly decreased with decreasing plant age at the onset of flowering.
Developmental rates, as measured by the number of main stem leaves at
the onset of flowering, did not significantly vary between the high and
medium nitrogen treatments, but were 30% lower in the low nitrogen treatment. Net photosynthetic rates during vegetative growth did not significantly vary with plant age, but decreasing nitrogen supply reduced net
photosynthetic rates 30% on average from the highest to the lowest nitrogen
supply. The onset of flowering was associated with a 30% and 20% reduction in net photosynthetic rates in the high and medium nitrogen treatments, respectively, but there was no significant change in the low nitrogen
treatment. Decreases in the relative balance between source:sink tissue associated with increasing age at flowering were associated with smaller relative responses of net photosynthetic rates to the transition between vegetative and reproductive growth. These results suggest that both carbohydrate source:sink balance and nitrogen concentration affect photosynthetic
responses to developmental changes.

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205

LEWIS, KRISTIN C.* and FAKHRI A. BAZZAZ. Harvard University,

Cambridge, MA. Evolution of defense chemistry in Alliaria petiolata
since introduction into North America.
Invasive species threaten the integrity and function of ecosystems across
the globe. However, prevention of invasion has proved elusive because of
the few commonalities among invasive species that might help predict invasiveness. One characteristic common among invasive plant species is
escape from specialist herbivores in their new range. It has been predicted
that plants released from herbivory will adjust their resource allocation to
reduce investment in defense, particularly when defenses are costly. Such
adjustment could increase resources available for activities such as growth
and reproduction, which might in turn contribute to competitive ability and
rapid spread. Alternatively, release from specialists and continued pressure
from generalists in the new range could increase investment in defenses if
these mainly deter generalists. Using an invasive crucifer, Alliaria petiolata,
with costly defenses (glucosinolates), we are investigating whether and how
this species has shifted resource allocation between chemical defenses and
other fitness-related traits. We have compared growth, herbivory, defense
chemistry, and reproduction in field populations in New England and in
Hungary to determine whether invasive US populations differ significantly
from native-range European populations. Using herbivore exclusions, we
can also distinguish between constitutive (inflexible) defenses and herbivore-induced defenses. These data will provide insight into the changes
that have occurred since introduction of these plants and the role of defense
chemistry in competitive ability and invasiveness. The potential for herbivory in the new range and flexibility in resource allocation could become
useful screening tools to predict which species may become invasive when
introduced outside their home range.
LEWISON, REBECCA1,* and SELINA HEPPELL.2 1 Nicholas School of
the Environment and Earth Science, Duke University, Beaufort, NC, US;
2
Department of Fisheries and Wildlife, Oregon State University, Corvallis,
OR, US. Predicting recovery patterns in short and long-lived organisms.
Life history theory has been used to evaluate the effect of different demographic profiles on population growth rates. These comparisons have
served to place organisms into broad categories long vs. short lived,
high vs. low fecundity (effort), delayed or early reproduction. Life history
theory also makes predictions about a populations ability to recover from
a collapse, such as one mediated by severe environmental or anthropogenic
factors. How useful are life history characterizations to projecting the recovery process? Can we predict post-disturbance population trends for species of conservation concern? Large, historical pulse disturbances that have
occurred in the past 30 years provide a natural experiment to test these
predictions. To test the utility of demographic profiles to predict recoveries,
we construct a standard population model for each species based on all
available demographic data. Using time series data (pre and post disturbance), we then compare model-predicted and observed recovery trends
for a range of short and long lived species. For cases where the demographic model projects a recovery, but none was observed, we explore the
likelihood alternate hypotheses, such as inaccurate demographic estimates,
Allee effects, or compensatory mortality from another source. Our comparisons suggest the utility of demographic profiles to predict recovery may
be species specific, as there were no clear trends in the success of the model
to capture the post-disturbance trajectories.
LI, BAI-LIAN. University of California, Riverside, CA. Biological and
physical foundations of allometric scaling revisited.
Allometric scaling laws have recently received much attention in ecology,
especially the work done by West and collaborators by noting the transport
of materials on the fractal-like networks within living bodies. Here we reanalyzed their work and found that their model and theory are biologically
and physically not sound, and mathematically inconsistent, even under their
own assumptions. We also showed that their results could be derived or
explained by the two fundamental biological and physical principles (constancy of energy flux per unit surface area and the existence of minimal
levels of the volume-specific metabolic rate for supporting life) that characterize the primary process of energy consumption by an organism from

206

Abstracts

its external environment, without involving the knowledge about transport

materials and optimization within a living body. In addition, we predicted
and explained the exponential decrease of leaf area index with elevation,
the change of animal body geometry and the breakpoint of the allometric
scaling. Since the process of energy consumption by an organism from the
external environment is much more fundamental than the transport of matter in a living body, our approach should be more general and robust, and
in particular, this approach is logically more coherent and connected when
linking an individual organism to population, community, ecosystem and
landscape patterns and processes.
LI, HARBIN,* KARI MINKKINEN, ZHENGQUAN WANG and CARL
TRETTIN. Uncertainty in scaling up carbon estimates in peat soils of
Finland: Comparison among four methods.
Policy or decision making often requires extrapolating information across
scales (i.e., scaling up) because most management problems are occurring
at large scales, but much of our knowledge and data is accumulated at
small scales at which research activities of most disciplines take place. In
the process of scaling up, errors from data and models may inevitably get
propagated into results. Uncertainty is a fundamental characteristic of modeling (or scaling) because errors, caused by incomplete data, limitations of
models, and lack of understanding of underlying processes, exist in every
aspect of modeling. Thus, uncertainty of large-scale estimates must be studied and quantified as an integral part of scaling to guarantee the adequacy
and reliability of the scaling results. In this study, we calculate carbon
estimates in peat soils of Finland based on data from intensive sampling
at the plot level, quantify uncertainty of the estimates, and examine effectiveness of different uncertainty analysis techniques. The scaling algorithm
used is the mathematical expectation with aggregated area data of vegetation types. The four methods of uncertainty analysis compared are: probability theory, Taylor series approximation, Monte Carlo simulation, and
sequential partitioning. The key error sources considered are: peat depth,
peat bulk density, peat carbon content, and total areas of vegetation types.
The results suggest that uncertainty of the carbon estimates is high and
must be reduced to be useful to policy-making and that peat depth and
bulk density show the highest relative contributions to the uncertainty of
the total carbon estimates. All four methods of uncertainty analysis work
well in this relatively simple scaling exercise, indicating that, for complex
situations, sequential partitioning is a promising method given that models
can be divided into independent compartments to deal effectively with uncertainty assessment.
LI, J. -H.,1,* R. B. LINKE,2 A. MADRITSCH,2 M. H. MEISTER,2 C. R.
HINKLE3 and B. G. DRAKE.1 1 Smithsonian Environmental Research Center, Edgewater, MD 21037, USA; 2 University of Vienna, A-1090, Vienna,
Austria; 3 Dynamac Corp., Kennedy Space Center, FL 32899, USA. Photosynthetic acclimation of scrub-oaks to long-term elevated CO2: A
summary of seven years.
The Florida scrub-oak ecosystem has been exposed in situ to elevated atmospheric carbon dioxide concentration (Ca) (350 mmol mol-1 above ambient) in large open-top chambers since May 1996. The photosynthetic
responses of Quercus myrtifolia willd. and Quercus geminata Small, the
dominant and codominant species, have been studied at physiological and
biochemical levels on detached branches in the lab and on intact leaves in
the field during the last seven years. Photosynthetic acclimation has been
species and season specific, occurring in Q. myrtifolia only initially, then
later in Q. geminata only, and not occurring in young leaves for both
species and during senescence for Q. myrtifolia. Discrepancies on acclimation have also occurred comparing measurements made on predawn excised branches in the lab with intact leaves in the field. Maximum rate of
carboxylation (Vcmax) and maximum rate of potential electron transport
(Jmax) measured indoor were 21% lower than that measured outdoor in
August and September 2002. For Q. myrtifolia, significant acclimation was
indicated from outdoor measurements but not from indoor measurements
at the later stage. Photosynthetic acclimation in Q. myrtifolia may be caused
by a limitation of carbon sink through feed-back inhibition since elevated
Ca significantly increased sugars and starch contents but did not affect leaf
nitrogen, total soluble protein and Rubisco contents per unit leaf area. Large
variations arise between species, exposure period, season and excised vs.

intact measurements highlight the importance of long-term elevated Ca

studies with native ecosystem as well as measurement approaches and
methods.
LICHTER, JOHN,1 STACY BARRON,1 ADRIEN C. FINZI2 and WILLIAM H. SCHLESINGER.3 1 Bowdoin College, 6500 College Station,
Brunswick, ME, USA; 2 Boston University, Boston, MA, USA; 3 Duke University, Durham, NC, USA. Soil carbon sequestration and turnover after six years of CO2 enrichment at the Duke Forest.
After three years of CO2 fumigation at the Duke Forest Free-air CO2 Enrichment (FACE) experiment, a significant accumulation of carbon associated with the elevated CO2 treatment was detected in the forest floor, but
not in the mineral soil. Now after six years, the forest floor under the
elevated CO2 treatment has continued to accumulate carbon at a steady rate
of 57 gC m-2 yr-1 greater than the control. However, there are still no
detectable changes in the carbon content of the mineral soil. Increased
carbon inputs to the forest floor associated with CO2-enhanced primary
productivity do not entirely account for the increase in forest-floor carbon
storage, suggesting that the rate of decomposition may be declining under
the treatment. Estimates of decomposition based on measured carbon pools
and inputs are consistent with reduced decomposition under the treatment
(Ambient CO2: k 5 0.345 (+0.020); Elevated CO2: k 5 0.273 (+0.012); n
5 3, p 5 0.036). Although there is no indication of a change in litter
chemistry, decomposition rates may decline if microbial activity is constrained by nutrient availability. Our data suggest that the CO2-induced
growth enhancement at the Duke Forest FACE experiment will begin to
attenuate as nitrogen availability becomes progressively more limiting with
declining decomposition.
LIEBHOLD, ANDREW M.1,* and OTTAR N. BJRNSTAD.2 1 Northeastern Research Station, USDA Forest Service, 180 Canfield St., Morgantown, WV, USA; 2 Depts of Entomology and Biology, 501 ASI Bldg.,
Pennsylvania State University, University Park, PA, USA. Roles of climate and dispersal in the synchronization of insect outbreaks over
large spatial scales.
Outbreaks of phytophagous forest insects are often spectacular, causing
defoliation of millions of hectares of forests. A common characteristic in
these outbreaks is the phenomenon of spatial synchrony among populations
separated by as much as 1,000 km. By comparing patterns of synchrony
among insect species exhibiting varying degrees of dispersal capabilities
we are able to draw inferences about the relative role of dispersal vs. weather (the "Moran effect") as causes of spatial synchrony. Our analyses utilized
historical aerial survey records of damage caused by the spruce budworm,
the western spruce budworm, the larch budmoth, the mountain pine beetle,
the forest tent caterpillar, and the gypsy moth. Results suggest that weather
is a more dominant cause of synchrony but the mechanisms by which
weather function are often difficult to identify. Furthermore, geographical
variation in density dependence may be critical to understanding how stochastic weather affects population synchrony. Finally, we discuss the
emerging field of complex spatial dynamics, its theoretical basis, methods
for its identification in natural populations, and likely roles of weather.
LILL, JOHN T.1,2,* and ROBERT J. MARQUIS.2 1 George Washington
University, Washington, DC, USA; 2 University of Missouri-St. Louis, St.
Louis, MO, USA. Leaf ties as oviposition sites for forest arthropods.
Most efforts to explain variation in the local structure of insect herbivore
communities have focused on plant quality factors (e.g., nutritional and
secondary chemistry), given their demonstrated impacts on components of
herbivore fitness. However, for many, if not most insects, host plants also
provide a "home" for their resident herbivores, sheltering them from the
elements and attack from natural enemies. We experimentally investigated
the influence of leaf shelters made by leaf-tying caterpillars on colonization
via oviposition for forest arthropods of white oak (Quercus alba). We created artificial leaf ties by clipping together adjacent leaf pairs and compared
oviposition rates on these leaves with non-tied leaf pairs on the same plant.
Because naturally occurring leaf ties typically contain leaf-tying caterpillars, frass, silk, and leaf damage, we also investigated the effects of these
occupancy-related cues on oviposition by comparing oviposition in initially

occupied vs. unoccupied leaf ties. Compared to non-tied controls, oviposition (across all taxa) was four times greater on unoccupied clipped leaves
and five times greater on previously occupied ties. These results indicate
that pre-existing shelters are actively sought out by adults, and that occupancy-related cues further enhance the level of oviposition. The magnitude
of the effect varied considerably among feeding guilds, with leaf-chewing
herbivores and scavengers showing the strongest response and sucking insects and predators showing less of response. Moreover, caterpillar species
differed in their response to occupancy-related cues. Our results highlight
the importance of herbivore-herbivore interactions in shaping natural arthropod communities. Our previously published experimental work demonstrated that plant architecture influences the level of leaf-tying in this
system. Therefore, the impact that these herbivore-herbivore interactions
have on the local arthropod communities depends in part on plant architecture.
LIN, GUANGHUI,1,2,* JOOST VAN HAREN,1 DANIELLE PIERCE,1
TIFFANY MORRIS,1 ALLEN WRIGHT1 and JOSEPH BERRY.3 1 Biosphere 2 Center, Columbia University, Oracle, Arizona, USA; 2 Lamont
Doherty Earth Observatory, Columbia University, Palisades, New York,
USA; 3 Department of Global Ecology, Carnegie Institution of Washington,
Stanford, California, USA. Responses of ecosystem gas exchanges to
simulated drought disturbance in a rainforest mesocosm.
Model and preliminary field studies indicate that drought disturbance associated with El-Nino events plays a significant role in controlling ecosystem exchanges of CO2, water and other trace gases. In this study, responses
of ecosystem gas exchanges to a series of simulated drought episodes in a
rainforest mesocosm were investigated using the facility of Biosphere 2.
Net ecosystem exchange of CO2, (NEE), soil respiration, evapotranspiration
(ET) and emission of N2O were measured before, during and after four
repeated month-long drought episodes in 2000-2002. Night NEE showed
a step-wise decrease to the drought, indicating differential sensitivities of
litter decomposition, plant respiration and oxidation of soil organic matter
to reduced soil moisture. Ecosystem respiration recovered quickly to predrought level after rewet treatment. Soil respiration showed significant reduction at the end of each drought episode. Canopy CO2 uptake also
showed a significant (40-60 %) reduction in response to the month-long
drought. In most cases, drought-induced reduction in ecosystem respiration
was balanced by the reduction in canopy CO2 uptake, resulting in little
change in carbon sink-source relationship at the whole system level. A
large decrease in carbon isotope ratio of ecosystem respired CO2 (about 5
per mil) right after rewet events suggested that (1) 13C-depleted carbons
from root exudates accumulated during the drought were quickly released
by soil microbes and (2) litter decomposition responded very quickly to
the rewet treatment. ET showed a significant decrease in response to
drought and ecosystem water use efficiency decreased as soil moisture decreased. The drought resulted in a 60-70 % decrease in N2O production
but a large pulse of N2O was observed within the first 36 hours immediate
after rewetting the mesocosm. Changes in the isotope ratios of N2O indicated a rapid shift between nitrification- and denitrification-dominated processes in response to drought and rewet episodes.
LIN, YICHING* and CAROL K. AUGSPURGER. University of Illinois,
Urbana, IL. Causes of a decline in species diversity in a deciduous
forest.
A theory of the maintenance of species diversity must include the causes
of both the maintenance and decline of species diversity. The objective of
this long-term study was to examine possible causes of the decline in species diversity over a 62-yr period (1939-2001) in Brownfield Woods, an
old-growth remnant in central Illinois. We hypothesized that the population
growth of Acer saccharum, the dominant species, arose from a lack of
density-dependent mortality, while the declining rare species were excluded
by competition from A. saccharum. Historical maps of trees (. 7.5 cm
DBH) from 1939, 1951, 1988, and 2001 were used to examine demographic patterns, population growth, and neighborhood effects of A. saccharum
on survival and recruitment of eight species. A. saccharum showed lower
mortality and higher recruitment than the majority of rare species. This
demographic advantage resulted in accelerated population growth. Population size of A. saccharum in 1988 and 2001 was significantly higher than

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207

the projected sizes estimated from transition matrices of 1939-51. Rare

species showed variable patterns of population growth. Aesculus glabra
expanded its population. Celtis occidentalis and two diseased species, Ulmus americana and Ulmus rubra, declined at an accelerated rate. Quercus
rubra declined at a constant rate, while Tilia americana and Fraxinus spp.
declined at a decelerated rate. Finally, survival and recruitment of all species, including A. saccharum,were higher in areas with significantly lower
basal area of A. saccharum. Thus, we detected both density-dependent mortality and interspecific competition in this study. The population of A. saccharum was regulated somewhat by its own density despite its fast population growth. Rare species were excluded by A. saccharum to a certain
extent, although their decline was slowed due to a moderate disturbance,
Dutch elm disease, in the 1950s. Historical datasets are invaluable in evaluating causes of species decline.
LINCOLN, DAVID E. University of South Carolina, Columbia, SC. Attraction of parasitoid enemies to a constitutively odorous plant.
Volatile emissions from leaves has been implicated in plant defense via the
attraction of herbivore enemies by induced emissions following foliar herbivory. Emissions from constitutively odorous plants, however, have received little attention. I examined the attraction of parasitoid wasps to an
odorous shrub, wax myrtle (Myrica cerifera), in two adjacent habitats. M.
cerifera is a common shrub in the zonal vegetation bordering coastal
marshes in the southeastern USA and as an understory shrub in inland
forests. The leaves constitutively produce a mixture of volatile compounds,
dominated by mono- and sesqui-terpenoids. Ten pairs of plants were selected along a marsh edge and one plant had 25 leaves mechanically
wounded while the other remained unwounded. Four yellow sticky traps
(7.5 X 10 cm) were attached to each plant. The same design was performed
approximately 100 m away in a mixed oak/pine forest adjoining the marsh.
The sticky traps were collected 24 h later and the abundance of parasitoid
wasps was determined. In both habitats, mechanically wounded plants attracted more parasitoid hymenopterans over the 24 h time period than did
unwounded plants (P.0.01). More parasitoids were captured in the forest
understory than at the marsh edge, despite greater volatile production
among plants in the high light prevalent at the marsh edge habitat. In a
separate experiment, groups (N55) of three plants were unwounded, 25
leaves wounded, or 50 leaves wounded. Increased wounding (50 vs 25
leaves) had a marginally significant attractive effect (P50.06). These results suggest that constitutively odorous plants may attract herbivore enemies via their emissions, but the effect may differ among habitats.
LINDER, ERIC T.* Do nested subsets occur in naturally fragmented
forests?
Examination of nestedness among communities has frequently been conducted upon communities in artificially fragmented landscapes or on true
island archipelagos. I examined the avian communities across a series of
naturally fragmented aspen stands in mountains of Utah. It is unclear how
sensitive species that evolve in a highly fragmented system such as this
should be to fragmentation. Furthermore, during times of environmental
stress it has been noted that rare species frequently become extirpated from
communities, which may or may not impact nestedness. Using the computer program NESTED I determined that avian communities formed a
nested subset pattern; implying nonrandom processes were governing the
distributional patterns of species in this landscape. This pattern held for all
three years examined. However, a drought during the second breeding season did not result in dramatic changes in nestedness compared to pre- and
post-drought seasons despite significant changes in avian community structure. Specific changes in community structure will be discussed.
LINDROTH, RICHARD L.,1 WILLIAM F.J. PARSONS2 and JAMES G.
BOCKHEIM.3 1 University of Wisconsin, lindroth@entomology.wisc.edu,
Madison, WI, USA; 2 Universite Laval, Ste. Foy, Quebec, Canada; 3 University of Wisconsin, Madison, WI, USA. Consequences of elevated carbon dioxide and ozone atmospheres for leaf litter decomposition at
Aspen-FACE.
The future capacity of forest ecosystems to sequester atmospheric carbon
is likely to be influenced by CO2-mediated shifts in nutrient cycling (via

208

Abstracts

changes in litter chemistry) as well as by interactions with other environmental pollutants, such as ozone (O3). We evaluated the independent and
interactive effects of enriched CO2 (560 ppm) and O3 (1.5 x ambient) environments on decomposition of trembling aspen (Populus tremuloides)
and paper birch (Betula papyrifera) leaf litter at Aspen FACE (Rhinelander,
Wisconsin, USA). Litter mass loss and chemical composition were monitored over two years. Reciprocal transplants of litterbags were employed
to separate the effects of substrate quality (chemistry) from those of environment (fumigation treatment). Calculation of mass loss rates (k-values)
revealed significant treatment effects that varied between tree species. For
aspen, both CO2 and O3 retarded mass loss, and their effects were additive.
Changes in loss rates were due primarily to shifts in litter chemical composition rather than to changes in the environment in which decomposition
occurred. For birch, enriched CO2 and O3 independently tended to accelerate decomposition, but in combination did not do so. Accelerated decomposition was due to changes in both chemical composition and treatment
environment. Differences in initial leaf litter quality were maintained between the species and sustained for at least one year among the fumigation
treatments. For both aspen and birch, starch and condensed tannins were
initially higher under elevated compared to ambient CO2, regardless of O3
level. Elevated C/N and lignin/N ratios were sustained under CO2 enrichment, and likely contributed to slower litter decay rates for aspen (+CO2
and +CO2+O3 treatments) and birch (+CO2+O3 treatment). These results
suggest that nutrient cycling dynamics of north temperate forest ecosystems
will change under atmospheric conditions predicted for the future.
LITCHMAN, ELENA,1,* CHRISTOPHER KLAUSMEIER,1 OSCAR
SCHOFIELD2 and PAUL G. FALKOWSKI.2 1 School of Biology, 310
Ferst Dr., Georgia Institute of Technology, Atlanta, GA, USA; 2 Institute
of Marine and Coastal Sciences, 71 Dudley Rd, Rutgers University, New
Brunswick, NJ. Resource-based niches of phytoplankton functional
groups.
Phytoplankton community composition profoundly affects patterns of carbon and other major element cycling as well as the structure of aquatic
food webs. Can we predict spatial and temporal occurrence and dominance
of major functional/taxonomic groups of phytoplankton? A significant step
would be to define the ecological niches of major taxa. To characterize
resource-based niches, we compiled a database of nutrient-dependent
growth and uptake parameters for different species of marine and freshwater phytoplankton belonging to major taxa: diatoms, dinoflagellates, coccolithophores, green algae and cyanobacteria. Parameters describing nutrient uptake and growth were significantly different among taxonomic
groups. Generally, diatoms are the best nutrient competitors, while dinoflagellates are the worst. Our database reveals important physiological constraints on the evolution of various traits that confer nutrient competitive
abilities: the maximum uptake rates of nitrate were positively correlated
with the minimum nitrogen quota and nitrogen half-saturation constant.
Parameters describing nutrient uptake and growth were significantly correlated with cell size: carbon-specific maximum uptake rates decreased
nonlinearly with cell size in a group-specific way.
LIU, HONG* and ERIC MENGES. Effects of wet- vs. dry-season fires
on vital rates of an endemic herb.
The season in which fire occurs may influence how plant populations respond to fire. In this paper, we report results from a four-year study on the
effects of replicated experimental burns, in wet vs. dry seasons, on demographic vital rates of Chamaecrista keyensis, a narrowly endemic understory herb of pine rockland forests of the Lower Florida Keys. Chamaecrista keyensis in the burned treatments had lower survival than unburned treatments, and plants in the wet-season burned treatments suffered
higher first-year post-burn mortality than those in the dry-season burned
treatments. However, survival 2-3 years post-fire was much greater in either
burned treatments than in the controls. The three-year cumulative survivals
in the wet-season burn treatments were significantly lower than either the
dry-season burn or unburned treatments. Dry-season burn treatments were
not different from the unburned treatments. Fires, regardless of the season,
did not stimulate growth of C. keyensis. In contrast, wet- and dry-season
burn treatments each stimulated C. keyensis reproduction comparing to unburned treatments. The dry-season burn treatments had greater seedling

recruitment than the wet-season burn or control treatments over the three
years post-burn. Overall, dry-season burn treatments seem to better stimulate population growth than wet-season burn treatments. This result is
unexpected because the wet season is the normal time for lightning ignited
fires in the Florida Keys. Nevertheless, slight differences in timing of burns
within the wet season may be important to the demography of C. keyensis.
LIU, JIANGUO,1 GRETCHEN C. DAILY,2 PAUL R. EHRLICH2 and
GARY W. LUCK.2 1 Department of Fisheries and Wildlife, Michigan State
University, East Lansing, MI, USA; 2 Center for Conservation Biology,
Stanford University, Stanford, CA, USA. Implications of household dynamics for resource consumption and biodiversity.
Human population size and growth rate are often considered important
drivers of biodiversity loss, while household dynamics are usually neglected. Aggregate demographic statistics may mask striking changes in the size
and number of households, and their effects on biodiversity. Household
dynamics influence per-capita consumption and thus biodiversity through,
for example, fuelwood consumption, habitat alteration for home-building
and associated activities, and greenhouse gas emissions. We found that
growth in household numbers globally, and particularly in biodiversity hotspots (areas rich in endemic species and threatened by human activities),
was more rapid than aggregate population growth between 1985 and 2000.
Even in places where population size declined, the number of households
increased substantially. Had average household size (i.e., the number of
occupants) remained static, there would have been 155 million fewer
households in hotspot countries in 2000. Reduction in average household
size alone will add a projected 233 million additional households to hotspot
countries from 2000-2015. Rapid increase in household numbers, often
manifested as urban sprawl, and resultant higher per-capita resource consumption in smaller households pose serious challenges to biodiversity conservation.
LIU, MINGLIANG,1,2,* HANQIN TIAN,1,2 SHUFEN PAN,1,2 JERRY
MELILLO,3 DAVID KICKLIGHTER3 and JOHN REILLY.4 1 Chinese Ecosystem Research Network, IGSNRR, Chinese Academy of Sciences, Beijing, 100101, China; 2 The University of Kansas, Lawrence, KS 66045;
3
Marine Biological Laboratory, Woods Hole, MA 02543; 4 MIT Joint Program on Science and Policy of Global Change, 77 Massachusetts Avenue,
Cambridge, MA 02139. Daily, seasonal and interannual variations in
carbon and water fluxes in terrestrial ecosystems: the Daily TEM and
its applications to Chinese Ecosystems.
The Terrestrial Ecosystem Model (TEM) has been widely used to investigate large-scale ecosystem processes in response to changing climates and
human impacts. The TEM model uses spatially referenced information on
climate, elevation, soils and vegetation to make monthly estimates of important carbon and nitrogen fluxes and pool size. This monthly model has
limited its applications to estimate daily exchanges of carbon and water
between terrestrial ecosystems and the atmosphere, and to simulate crop
yield in agricultural ecosystems which is sensitive to daily or weekly agronomic practices and climate variability. Therefore, there is an increasing
need to develop the model by expanding monthly time step into daily time
step to accurately simulate ecosystem processes. Here, we have developed
a daily version of the Terrestrial Ecosystem Model to simulate the terrestrial
ecosystem processes in which the mechanism of photosynthesis, nitrogen
cycling, nutrient redistribution, water cycle and soil organic pools are different from the current version of TEM. We use daily data from Chinese
Ecosystem Research Network (CERN) to calibrate the daily TEM. The
daily TEM has been used to investigate daily, seasonal and interannual
variations in carbon and water fluxes for three types of ecosystems: cropland, grassland and forests.
LLOYD, ANDREA H.,* ALEXIS WILSON and CHRISTOPHER L. FASTIE. Middlebury College, Middlebury, VT. Population dynamics of black
spruce at its northern limit in Alaska.
Black spruce (Picea mariana) forests dominate much of interior Alaska,
but black spruce is absent or extremely rare at arctic and alpine treeline
locations. In the Brooks Range in northern Alaska, the northern limit of
black spruce is several kilometers south of the northern range of white

spruce. Although white spruce have successfully advanced into tundra as

it has warmed in Alaska over the last several decades, the effects of climate
on the distribution of black spruce remains largely unknown. We investigated the dynamics of black spruce populations at the species northern limit
in the Brooks Range, Alaska, exploring the relative importance of fire and
climate as limits on black spruce population viability at its northern limit.
We reconstructed fire history and recruitment dynamics in three replicate
sites at the species northern limit, and developed a matrix population model
to explore black spruce population dynamics. All sites were affected by a
nearly stand-replacing fire slightly more than 100 years ago. Black spruce
experienced high recruitment rates in the 20 years following the fire, but
recruitment declined precipitously after that point. Black spruce currently
recruits primarily by layering, and although low densities of seedlings were
present in all sites, seed germination rates in greenhouse conditions were
extremely low (,1-8%). White spruce recruitment rates have been much
more consistent over time, and white spruce are now significantly overrepresented in the seedling size classes. Black spruce are likely to become
increasingly uncommon in these stands as time-since-fire increases, and the
persistence of black spruce on the landscape in the absence of fire will
depend almost entirely on asexual reproduction. The low success of black
spruce recruiting in the absence of fire suggests that the species limit is
unlikely to advance unless warmer conditions are accompanied by an increase in fire frequency.
LOCKABY, B. G.,1,* DAOWEI ZHANG1 and PHILIP CHANEY.2 1 School
of Forestry & Wildlife Sciences, Auburn University, Auburn, AL, USA;
2
Department of Geology & Geography, Auburn University, Auburn, AL,
USA. Ecological and socioeconomic influences of urban development
on forested landscapes in Georgia, USA: The West Georgia project.
An interdisciplinary investigation of urban development impacts was installed in 2001 on a forested landscape near Columbus, Georgia, USA. The
study area extends approximately 100 km to the northeast of Columbus
and reflects a gradient in human population density, rate of growth, and
land use / cover. Major land uses include urban, developing (residential),
forested (pine), forested (mixed), and rural (pasture). Field sampling is
watershed-based and encompasses approximately 22 catchments (1000 5000 ha). Feedback relationships between landscape development and economics, culture, biodiversity, water quality, vegetation patterns, habitat dynamics of large mammals, and aesthetics are being studied. Water quality
results indicate a strong influence of land use on fecal coliform, nitrate,
and sediment concentrations in surface water although no differentiation
was apparent between pine vs. mixed pine-hardwood watersheds. Econometric models have been developed to predict land use / cover changes in
5-20 years under different economic, policy, and demographic scenarios.
Regression relationships between land use descriptors and environmental
responses are being used to anticipate impacts of future land use changes
on water quality, biodiversity, and other response factors.
LOEWENSTEIN, NANCY J.,* MICHELE L. BURTON, LISA J. SAMUELSON and PHILIP L. CHANEY. Center for Forest Sustainability,
Auburn University, Auburn, AL, USA. Impact of urbanization and forest structure on non-native plant distribution: The West GA project.
Major sources of non-native plant propagules are lawns and gardens. As
urbanization expands into rural areas, an increase in non-native plants at
the urban-rural interface is expected. A study looking at the influence of
urbanization and forest structure on the distribution and invasiveness of
non-native plants across an urban-to-rural gradient north of Columbus, GA
was initiated in 2002. Plant sampling was conducted two times during the
growing season within 20 watersheds across the gradient. Quadrats were
randomly placed in each of three locations along a 500 m section of each
stream: within the channel, immediately outside the stream bank, or along
a short transect perpendicular to the stream. Also measured at each site
were: soil structure, pH and nutrient levels, slope, aspect, distance to nearest road, light availability, leaf area index, overstory structure and composition, size of forest fragment and land use-land cover within the watershed. Initial results indicate that: 1) at least one non-native plant species
was observed in every watershed, 2) the greatest number of non-native
plant species occurred at the urban sites, 3) several invasive species (Ligustrum sinense, Lonicera japonica, Microstegium vimineum and Murdan-

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209

nia keisak) were wide-spread and occurred in the majority of the watersheds. The potential exists for the spread of additional non-native plants
into the urban-rural interface, the success of which may impact or be impacted by forest structure.
LOFTIN, CYNTHIA S.,1 JON MCCLOSKEY,2 WILEY M. KITCHENS3
and MARSHA L. DUSEK.4 1 USGS-BRD Maine Cooperative Fish and
Wildlife Research Unit, University of Maine, Orono, Maine, USA; 2 University of Maine, Orono, Maine, USA; 3 USGS-BRD Florida Cooperative
Fish and Wildlife Research Unit, University of Florida, Gainesville, Florida, USA; 4 University of Florida, Gainesville, Florida, USA. Changes in
Savannah River tidal marsh vegetation distributions following removal
of a tidal flap gate.
We used remote sensing tools and field surveys to monitor changes in
vegetation distributions in the Savannah River tidal marsh following the
1991 removal of a tidal flap gate. Landsat TM and SPOT multispectral
imagery from October 1986, April 1987, May 1993, July 2000, October
2000, and April 2001 were compared using principle components analysis
to identify areas of changed brightness and greenness. Vegetation maps
were created from each image using classification procedures, and the maps
were compared with principle components images to identify from-to
change types. Predictions of a spatial vegetation succession model developed in earlier studies to estimate vegetation response to tide gate removal
were compared with the vegetation maps to determine if tidal freshwater
marsh had expanded as expected following tide gate removal. We identified
changes in greenness and brightness indicating changes in biomass, vegetation types, and water levels in the area during the study period. Increased
greenness was most extensive during 1987-1993. An increase in woody
and freshwater species occurred by 1993 in nearly 1/3 of the region predicted to be dominated by freshwater vegetation 2 years following tide gate
removal. Species found at sites with lowest salinity (,0.5ppt) before tide
gate removal have since expanded into previously brackish regions of the
marsh. By 2000 freshwater species had expanded to nearly 3/4 of the expected distribution of tidal freshwater marsh; Argyle Island and New Cut
approximate the current southward extent of vegetation associations intolerant of salinity .3.0 ppt. Shifts in dominance of Scirpus validus and
Zizaniopsis mileacea in mixtures with a variety of freshwater species have
occurred within and between years in this region since the gate was removed, and extent of Z. mileacea has increased in this region since tide
gate removal. Vegetation composition below Argyle Island has changed
little in response to tide gate removal, and remains primarily a mixture of
S. validus, S. robustus, and Spartina alterniflora.
LONG, JEREMY D.* and MARK E. HAY. Georgia Institute of Technology, Atlanta, GA. Learned aversion and retention: How marine fishes
avoid new aversive foods.
Numerous vertebrates and invertebrates learn to avoid foods that produce
physiological distress. However, the speed of learning, the duration of aversion without reinforcement, and the cues used are poorly investigated in
marine systems. In southeastern oyster reef communities, several co-occurring predators avoid the nudibranch, Doriopsilla pharpa, in the laboratory and the field. We examined the feeding behavior of two fish predators, Chasmodes bosquianus and Fundulus heteroclitus. Foods made with
tuna and lipophilic nudibranch extracts were initially consumed by C. bosquianus, but were regurgitated within minutes. Over the following week,
treatment fish rejected all tuna based foods, even those without nudibranch
extract, suggesting that they associate distress with tuna, not the extract.
Control fish unexposed to the extracts consumed tuna throughout this period. All fish were then switched to an alternate food. Treatment fish presented with tuna after 1 and 2 weeks without exposure to this food still
avoided tuna. When given daily presentations of tuna without compounds,
treatment fish avoided tuna for more than a week. In comparison, F. heteroclitus also initially accepted foods with lipophilic extracts and later
learned to avoid these foods, however, emesis was not evident nor did F.
heteroclitus learn to avoid palatable base foods. These results suggest that
F. heteroclitus rejected foods after detecting the nudibranch extract rather
than other cues associated with the food. Bioassay-guided fractionation of
nudibranch extracts identified a single bioactive chemical compound, the
sesquiterpene polygodial, responsible for extract unpalatability. Palatable

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Abstracts

foods with and without purified polygodial were initially consumed but
four days later, only half as many C. bosquianus ate foods with polygodial.
The observation that different predators use different cues to learn to avoid
the same food hints at the underlying complexity of learned aversions in
the marine environment.
LONG, ROBERT P.,1,* STEPHEN B. HORSLEY,2 PAUL R. LILJA,3
THOMAS J. HALL4 and SCOTT W. BAILEY.5 1 USDA Forest Service,
Northeastern Research Station, Delaware, OH; 2 USDA Forest Service,
Northeastern Research Station, Irvine, PA; 3 Pennsylvania Bureau of Forestry, Coudersport, PA; 4 Pennsylvania Bureau of Forestry, Middletown,
PA; 5 USDA Forest Service, Northeastern Research Station, Campton, NH.
Effects of forest liming on soil chemistry and sugar maple health in
Pennsylvania from 1986-2002.
A long-term liming study was initiated in 1985 to mitigate acidic soils and
has been intensively monitored at four forested sites on unglaciated soils
in north central PA. Dolomitic limestone was applied (22.4 Mg ha-1) in fall
1985. Soils were sampled and analyzed for exchangeable Ca, Mg, K, P,
Al, and acidity annually from 1986 to 1989 and in 1993, 1996, and 2001.
Soil pH in the upper 5 cm remained elevated at 6.4 on limed plots, compared with 3.8 on unlimed plots in 2001. Elevated Ca and Mg and decreased Al were measured at all depths to 60 cm in limed plots sampled
in 2001. Both P and K decreased in response to liming, but only in the
upper 5 cm. Changes in sugar maple (Acer saccharum Marsh.) crown vigor
have been evaluated yearly using a scale from 1 (dead) to 6 (healthy).
Crown vigor was consistently higher on limed plots, but has varied considerably due to stress events that include droughts (1988, 1991, 1995,
2001), and defoliations (1993, 1994). Mean sugar maple vigor in limed
plots peaked in 1998 at 5.5 while vigor in unlimed plots averaged 3.7.
Subsequent drought in 2001 decreased sugar maple vigor in all plots, but
by 2002 mean vigor in limed plots increased slightly from 3.9 in 2001 to
4.0 while trees in unlimed plots averaged 3.2 in both years. Sugar maple
basal area increment for trees on limed plots averaged 24.5 cm2 yr-1 and
trees in unlimed plots grew at a rate of 9.0 cm2 yr-1 from 1986 to 2002.
Cumulative sugar maple mortality from 1986 to 2002 was 30% in unlimed
plots and 13% in limed plots. These results indicate that a single lime
application has long-term beneficial effects on soil chemistry and sugar
maple vigor and growth.
LONSDORF, ERIC V.* University of Minnesota - Twin Cities, Saint Paul,
MN. Multiple disturbances in a metapopulation: The consequences of
inbreeding and nitrogen deposition to plant communities.
Nitrogen deposition, habitat destruction, and inbreeding can reduce the likelihood that a plant population may persist. While many studies explore the
effect of a single disturbance, native plant populations are likely to experience multiple and simultaneous disturbances. The efficiency with which
individuals acquire and use nitrogen (NUE) affects the response of populations to nitrogen deposition. This response is sensitive to inbreeding
caused by fragmentation that can reduce NUE. It is therefore important to
investigate these perturbations together. In this report, I use a metapopulation framework to study the potential consequences of nitrogen deposition
and habitat destruction to competing plant species with varying mating
systems. In this model, species coexist initially due to a competition-colonization tradeoff. Based on the mating systems of each species (selfing
rate) habitat destruction will cause differential changes to the initial competition and colonization abilities. Non-selfing species suffer from pollen
limitation, and thus a decline in colonization rate as destruction occurs,
while more selfing species are likely to suffer from inbreeding depression
in resource uptake, and thus a decline in competitive ability. Increasing
nutrient supply interacts with mating system, and pollen limitation drives
the likelihood of coexistence between species at high resource levels. Simultaneous consideration of genetic and environmental contributions to
population dynamics is essential as conservation biologists and other scientists make predictions about future persistence.
LORD, CYNTHIA C.* and JONATHAN F. DAY. University of Florida IFAS, Vero Beach, FL, USA. Modeling of arboviruses in Florida.
We are using simulation models to understand the ecology and transmission
of arboviruses, such as St. Louis encephalitis and West Nile viruses. These

viruses cycle between avian and mosquito species and human risk of disease is related to the levels of transmission in wild birds. Initial models
suggested an interaction between the timing of peak mosquito populations
and peak bird reproduction in the likelihood of outbreaks. A finer scaled
sensitivity analysis was performed with most parameters fixed at most
probable values and only those parameters controlling the timing of mosquito and avian populatons allowed to vary. There were more outbreaks in
this set of simulations than in previous sensitivty analyses, showing that
the most probable parameter values were very permissive for outbreaks.
This analysis showed that interactions between parameters are important in
the likelihood of outbreaks. We are pursuing this by considering fewer
parameters in combinations to look for interactions which affect the likelihood of outbreaks. None of the timing parameters affected the likelihood
of outbreaks directly, but mosquito mortality rates did influence the likelihood of outbreaks. Laboratory and field experiments have shown that
mortality is age dependent, which may change the influnce of mosquito
mortality on outbreaks. We are developing models incorporating age-dependent mortality to determine the effect on arbovirus transmission
LORTIE, CHRISTOPHER J., * ROB BROOKER, PHILLIPE CHOLER,
ZAAL KIKVIDZE,4 RICHARD MICHALET,5 FRANCISCO I. PUGNAIRE6 and RAGAN M. CALLAWAY.6 1 University of Montana, Missoula,
Montana, USA; 2 Centre for Ecology and Hydrology, Banchory, Banchory,
Scotland; 3 University Joseph Fourier, Grenoble, Grenoble, France; 4 Institute of Botany of the Georgian Academy of Sciences, Tbilisi, Republic of
Georgia; 5 Universite de Bordeaux, Talence, France; 6 Estacion Experimental de Zonas Aridas, Almeria, Spain. Rethinking individualistic theory
for communities: An integrative solution.
1,

Plant communities have been viewed as either a random collection of individuals or as organismal entities. For most ecologists, neither perspective
provides a modern view, yet formal positions continue to adopt this polarization. Exacerbating these extremes, a recent exchange in the literature
threatens polarization over the importance of negative versus positive interactions as predominant drivers of community organization. We propose
that current conceptual theory for communities is inadequate as a basic
foundation for the rapidly expanding empirical body of knowledge. It is
clear that species within natural plant communities can function individualistically and interdependently depending on synergism among: (i) stochastic processes, (ii) the abiotic tolerances of species, (iii) positive and
negative interactions among plants, and (iv) indirect interactions within and
between trophic levels. All of these processes are well accepted by plant
ecologists but not all fit comfortably within an individualistic paradigm of
plant communities. The position that communities simultaneously have
both individualistic and interdependent properties is analogous to the anthropic principle in quantum physics where light is both wave and particle.
Research conducted during the last 15 years strongly indicates that the
function of plant communities, like animal communities and ecosystems,
depends on both the properties of individuals and on the properties of
species aggregations. This dualistic nature creates environmentally and lifehistory driven variation from independence to interdependence among plant
species and is typically regarded as noise by many ecologists. However, as
in quantum physics, this noise should be viewed as a unique opportunity
for synthesis, not as a fundamental limitation of ecology.
LOTT, MICHAEL A.* and JOHN C. VOLIN. Florida Atlantic University,
Davie, FL. The growth and physiological ecology of two invasive nonindigenous fern species, Lygodium microphyllum and Lygodium japonicum.
Throughout the world, invasions by non-indigenous plant species threaten
the survival of many natural plant communities. This study examines the
growth and physiological ecology of two non-native species, Lygodium
microphyllum and L. japonicum, as well as two native vines, Vitis rotundifolia and Parthenocissus quinquefolia. The four species were grown in
shade houses under three light treatments. The relative growth rate (RGR)
of L. microphyllum was significantly greater in both the low- and mediumlight treatments as compared to the other three species. For example, at the
first harvest under low light conditions the RGR of L. microphyllum was
22.6 mg g-1 day-1 as compared to 18.8 mg g-1 day-1 for L. japonicum, 17.0
mg g-1 day-1 for V. rotundifolia, and 9.7 mg g-1 day-1 for P. quinquefolia.

Large differences in biomass allocation were observed between the two

non-native and native species. Under all three light treatments the nonnative ferns allocated significantly less resources into their climbing frames
(5-20%) as compared with the native vines (40-60%). Correspondingly, the
Lygodium species allocated greater resources to roots. However, it appears
that the differences in growth among the four species are largely explained
by differences in leaf physiology, where in both the low- and mediumlight treatments, whole-plant photosynthesis was significantly greater in the
two non-native climbing ferns, particularly so in L. microphyllum, compared to the two native vines.
LOUGHEED, VANESSA L.,1,* CHRISTIAN A. PARKER,1 MOLLIE D.
MCINTOSH,2 RICHARD W. MERRITT2 and R. JAN STEVENSON.1
1
Dept of Zoology, Michigan State University, East Lansing, MI, USA;
2
Dept of Entomology, Michigan State University, East Lansing, MI, USA.
Does size matter: a study of wetlands and biodiversity in an agricultural watershed.
Heterogeneity in plant species distribution produces much of the structural
variability throughout a wetland. This study was designed to look at the
loss of structural heterogeneity and its effects on the biota in both reference
wetlands and those impacted by agricultural land use. In July 2002, 8 small
(less than 1 ha) and 8 large (6 to 25 ha) isolated depressional marshes in
the Muskegon River watershed (Michigan, USA) were sampled for macrophyte, periphyton, zooplankton and macroinvertebrate communities using
a stratified random sampling design. Preliminary data indicate that diatom,
zooplankton and macrophyte communities in impacted wetlands show less
spatial variability and have fewer species than those in reference wetlands.
Initial analyses also indicate that, for the zooplankton community, both
reference and impacted small sites had greater beta diversity (ratio of total
species in landscape to local average) than large sites. Results from this
study suggest that land use practices negatively affect valuable wetland
functions by homogenizing habitats and reducing biodiversity. In addition,
smaller wetlands may represent valuable, unique habitats for wetland conservation.
LOVEYS, BETH R.,1,2 JOHN J. EGERTON,1,2,* WAYNE E. PIPPEN1,2 and
MARILYN C. BALL.1,2 1 Australian National University, jack.egerton@
anu.edu.au, Canberra, ACT, Australia; 2 CRC for Greenhouse Accounting,
Canberra, ACT, Australia. A mechanistic basis for lower freeze tolerance
under elevated [CO2].
Recent studies have shown that elevated [CO2] can adversely affect freeze
tolerance in evergreens such as snow gum (Eucalyptus pauciflora). The
mechanism, however, is unknown. In snow gum, lower stomatal conductance under elevated [CO2] results in an increase in leaf temperature of
approximately 3 8C. Here we test whether such higher leaf temperatures
during the photoperiod can influence the freeze tolerance of plants subject
to the same minimum night temperatures. A 3.5 8C increase in daytime
leaf temperature was simulated in the field using a Free Air Temperature
Increase (FATI) system. Freeze tolerance in these seedlings grown with or
without warming was compared with that of seedlings grown in adjacent
open-topped chambers (OTC) flushed with air containing either 350 or 700
ppm [CO2]. The experiment began at the start of autumn, prior to the onset
of freezing temperatures. Freeze tolerance was assayed by change in electrolyte leakage and photosynthetic efficiency (Fv/Fm) of leaf disks as a
function of minimum freezing temperature. Seven days after planting,
freeze tolerance was lower in leaves of plants grown with either warming
or elevated [CO2]. However, 27 days after planting, there were no differences in freeze tolerance between treatments. Thus, the elevated leaf temperature caused either by lower stomatal conductance in response to elevated [CO2] or by FATI warming increased the time required to achieve
freeze tolerance in snow gum.
LOYA, WENDY M.,1,* KURT S. PREGITZER,1 NOAH J. KARBERG,2
JOHN S. KING1 and CHRISTIAN P. GIARDINA.2 1 Michigan Technological University, Houghton, MI; 2 USDA Forest Service, North Central Researc Station, Houghton, MI. Tropospheric ozone reduces stable soil
carbon formation under elevated carbon dioxide.
Tropospheric O3 levels have increased by 35% over the last century across
the northern hemisphere, and are steadily increasing in association with

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211

development in tropical regions. Elevated tropospheric O3 inhibits forest

and agricultural productivity, including CO2-stimulated forest productivity,
but the impact of elevated O3 on carbon storage in soil is unknown. We
used O3 fumigation coupled with Free-Air CO2 Enrichment (FACE) technology in experimental aspen (Populus tremuloides) and birch (Betula papyrifera) forests to examine the impact of elevated O3 on formation of
decay-resistant soil carbon under elevated CO2. We found that increasing
O3 reduced formation rates of decay-resistant carbon in soil by 50%, relative to amounts entering this pool due to elevated CO2 alone. This change
was driven by reduced detrital inputs to soil and microbial utilization of
new carbon inputs. These results suggest that offsetting atmospheric increases in CO2 by increasing carbon sequestration in forest soils may be
limited by simultaneous increases in O3.
LUBCHENCO, JANE,1,* ANDREW ROSENBERG,2 RENEE DAVISBORN1 and LYDIA BERGEN.3 1 Oregon State University, Corvallis, OR;
2
University of New Hampshire, Durham, NH; 3 University of California,
Santa Cruz, Institute of Marine Science, Santa Cruz, CA. Connecting Science to Ecosystem-Based Management: Reflections on the Ocean Commissions.
In recent months, the country has heard from two national oceans commissions, both of which conducted comprehensive reviews of the Nations
ocean policies. The Commissions and the ensuing national dialogue are the
first broad scale assessment of the countrys attitudes, knowledge, management and policies regarding US waters in over 30 years. Both the independent Pew Oceans Commission (report released 9 April) and the congressionally mandated US Commission on Ocean Policy (in draft form as
abstract was written) relied heavily on scientific understanding in conducting their reviews. Both Commissions conclude there is a significant disconnect between current scientific knowledge about marine ecosystems and
the countrys policies and practices affecting these systems. Both Commissions highlight the extent to which the country should increase its investment in scientific research and monitoring necessary to understand the
dynamics of ocean ecosystems. The presentation will highlight relevant
findings of both commissions, then focus on two issues for further engagement by the scientific community. (1) Both Commissions emphasize
the need to take an ecosystem-based management approach to activities
affecting oceans, yet the guidance to do so is limited. (2) Scientific research
and monitoring programs that provide more comprehensive understanding
of the dynamics of marine ecosystems will be needed, but exactly what
they should look like needs to be defined. Many of the presentations in
this symposium provide models and information that relate to both of these
topics. The Partnership for Interdisciplinary Studies of Coastal Oceans,
PISCO, for example combines monitoring and research that focuses on the
near-shore portion of a Large Marine Ecosystem, the California Current
system along the west coast of the US. The new insights emerging from
PISCO suggest that it can serve as a model for other efforts to design
effective large-scale, long-term, interdisciplinary research and monitoring
programs that will inform policy and management.
LUCASH, MELISSA S.,1,* JOHANNES M. SCHOLBERG2 and RUTH D.
YANAI.1 1 SUNY-ESF, Syracuse, NY; 2 University of Florida, Gainesville,
FL. Diurnal changes in nitrogen uptake and root respiration in loblolly
pine seedlings.
Although energy derived from root respiration can be utilized for nitrogen
uptake, few studies have simultaneously measured root respiration and N
uptake. In this study, root respiration and N uptake were simultaneously
measured in loblolly pine seedlings grown in soil-uptake-monitoring
(SUM) columns filled with sand. To determine how the supply of photosynthate affects preference for NO3- or NH4+, N uptake was measured under
both light and dark conditions. N uptake was measured by filling columns
with a known volume and concentration of nutrient solution. After 2hrs,
the solution was removed by a vacuum pump and analyzed for NO3- and
NH4+. During the uptake measurements, root respiration was measured in
the column headspace using a LI-COR 6400. The measurements were repeated using a range of N concentrations.

212

Abstracts

LUNCH, CLAIRE K.* and DAVID R. BOWLING. Dept of Biology, University of Utah, Salt Lake City, Utah. Difficulties of measuring eddy
covariance fluxes of carbon dioxide at arid sites.
Arid and semi-arid ecosystems are dominant in most of the western United
States, but these regions have received comparatively little attention by the
CO2 flux monitoring community. Beginning in April 2001, we used the
eddy covariance technique to measure fluxes of energy, water, and CO2 in
a semi-arid grassland near Canyonlands National Park in southeast Utah.
The site was dominated by perennial native bunchgrasses and had low
vegetative cover and low productivity most of the year due to severe water
limitation. The majority of incoming energy (Rn) was converted into sensible heat flux (H). Little water was available to supply latent energy (LE)
fluxes except immediately after rain. The average midday Bowen ratio (H/
LE) was 16.8 during May-August and exceeded 50 during severe dry periods. These conditions presented a major challenge for accurate measurements of carbon dioxide flux due to temperature-induced density fluctuations in the CO2 analyzers used. Uncorrected CO2 fluxes were unreasonably
large, peaking at -25 to -30 mmol m-2 s-1 at midday (negative fluxes represent uptake of CO2), which is comparable to most temperate forests.
Density corrections based on the standard Webb, Pearman, Leuning approach resulted in unrealistic CO2 fluxes that were always positive regardless of time of day or season. Corrected fluxes did not follow soil temperature as expected for actual respiration fluxes but instead were correlated
with sensible heat flux. Reasons for these errors and potential solutions will
be addressed.
LUO, YIQI1,* and NCEAS N WORKING GROUP.2 1 University of
Oklahoma, Norman, OK; 2 University of California, Santa Barbara, CA.
Nitrogen limitation of ecosystem responses to rising atmospheric CO2:
Patterns and mechanisms.
One of the most controversial issues in global biogeochemistry is the regulation of terrestrial carbon (C) sequestration by nitrogen (N) availability.
Such a controversy is translated to a great deal of uncertainties in predicted
global terrestrial C sink in the next several decades. This talk will evaluate
the C/N interactions under a new conceptual framework that is progressive
N limitation (PNL). PNL takes place when available N in an ecosystem is
allocated to long-lived biomass pools and/or immobilized to increased soil
carbon stocks under elevated CO2. Synthesis of experimental results from
free-air CO2 enrichment (FACE) and open-top chamber (OTC) studies suggests three types of ecosystem responses: no PNL, delayed PNL, and immediate N limitation. The immediate N limitation happens in ecosystems
where elevated CO2 could not stimulate plant biomass growth and generate
no extra N demand. Delayed PNL occurs in ecosystems where CO2 stimulation in biomass growth and soil C storage is initially strong and progressively declines. The delayed PNL is usually associated with short-term
N stress alleviation mechanisms. Those mechanisms include N reallocation
among different plant and soil pools, changes in plant and soil C/N ratios,
and N mining via increased fine root growth. Those short-term mechanisms
usually do not significantly increase total N stocks in an ecosystem but
increase N use efficiency, supporting a positive ecosystem response to elevated CO2 even if labile N availability in soil declines. However, these
compensation mechanisms are not sustainable in the long term. The third
type of ecosystems is that CO2 stimulation of ecosystem productivity is
sustained while no PNL is observed within the lifetime of a CO2 experiment. The sustained CO2 stimulation is likely supported by long-term N
supply mechanisms. Such mechanisms balance extra N demand caused by
additional long-lived biomass growth and increased soil C stocks under
elevated CO2.
LUTSCHER, FRITHJOF* and MARK A. LEWIS. Centre for Mathematical
Biology, University of Alberta, Edmonton, Alberta, Canada. Dont get lost
in space.
Matrix models are a fundamental tool in population dynamics. Recently,
spatial structure has become recognized as a key factor in species survival
and persistence. Hence, spatial models are playing an increasing role in
management and conservation. However, spatially-explicit stage-structured
models can be very complex. Here, we show how such models can be
considerably simplified while still accurately capturing population growth

dynamics. The final model structure is the original matrix system modified
by average dispersal success. This technique provides a general but simple
framework for evaluating the effect of dispersal on population dynamics.
We will address questions such as minimal habitat size, reserve design,
habitat fragmentation and point-source pollution. As an example, we apply
this method to a simple model for marine invertebrates where larval dispersal differs significantly from dispersal at all other stages. We derive
conditions for reserve design and show how other influences such as habitat
deterioration can be incorporated.
LYNCH, ANN M.1,3 and THOMAS W. SWETNAM.2 1 USDA Forest Service, Rocky Mountain Research Station, alynch@fs.fed.us, Flagstaff, AZ,
U.S.A.; 2 The University of Arizona, Tucson, AZ, U.S.A.; 3 Colorado State
University, Fort Collins, CO, U.S.A. Dendrochronologic evidence of the
role of climate affecting insect outbreaks in western coniferous forests.
Tree-ring records were used to reconstruct spatial and temporal patterns of
western spruce budworm (Choristoneura occidentalis Freeman) outbreaks
in mixed-conifer forests in large regions of Colorado, New Mexico, and
Oregon, USA. We evaluated relationships between forest insect outbreaks
and climate by comparing these multi-century, multi-region reconstructions
to drought severity indices. These broad-scale comparisons show that wet
periods coincide with budworm outbreaks. Budworm population dynamics
appear to have a strong internal regulating component that may be a function of forest foliage production and budworm demographic response to
the timing, quantity, and quality of foliage. This system appears to be
entrained by regionally correlated cyclical patterns in summer moisture,
that lead budworm populations into oscillations varying in periodicity and
timing from one region and period to another. Budworm-forest modeling
and spectral analyses also suggest that strong, temporally variable cycles
of budworm populations are generated by internal controls, but that these
oscillations are consistently entrained across regions by precipitation cycles. Land-use history and fire exclusion may have altered the timing and
magnitude of contemporary outbreaks, but these changes are not consistent
across regions. Comparisons of the western spruce budworm reconstructions from the three regions indicate that disturbance regimes vary regionally, and that 20th century changes to these regimes have also varied.
Dendrochronologic reconstructions of pandora moth (Coloradia pandora
Blake) outbreaks indicate that there are complex interaction between precipitation, fire, and insect populations, with outbreaks initiating during dry
years. We found no evidence that either pandora moth or Douglas-fir tussock moth (Orgyia pseudotsugata (McDunnough)) outbreaks became more
or less extensive or severe, or of shorter or longer duration in the 20th
century.
LYNCH, ELIZABETH A.,1,* RANDY CALCOTE2 and SARA HOTCHKISS.3 1 Luther College, Decorah, IA; 2 University of Minnesota, Minneapolis, MN; 3 University of Wisconsin - Madison, Madison, WI. LateHolocene fire and vegetation history in the Wisconsin pine barrens.
Land managers seeking to restore "pine barrens" ecosystems in northwestern Wisconsin are interested in characterizing pre-European fire regimes.
Our objective is to use charcoal and fossil pollen in lake sediments to
reconstruct fire and vegetation patterns, and to investigate how fire and
vegetation interacted over the past 2000 years. At Ferry Lake we used
historical records to determine how known fires are represented in 210Pb
dated lake sediment, and then reconstructed forest fires for the past 2000
years. Charcoal accumulation rates (CHAR) of macroscopic (125-250 m)
fragments in contiguous core samples (each representing 5-10 years) were
calculated based on 210Pb and 14C dating. Graminoid charcoal fragments
were tallied separately to estimate the relative abundance of grass charcoal
over time. CHAR peaks greater than 1.25x background influx rates were
used to infer local forest fires. Pollen analysis shows a transition from oakdominated woodland to a relatively open pine forest or savanna at 1400
14
C yr BP. The fire regime also changed at this time from a 90-year fire
return interval to a regime with shorter return intervals, smaller CHAR
peaks, and less graminoid charcoal for 400 years after the vegetation
change. Beginning around 1000 14C cal yr BP the return interval lengthened
and both CHAR and graminoid charcoal influx began a decline that extended to the time of European settlement. Our results demonstrate that
even within the last 2000 years the vegetation and fire regime changed at

this site and that the vegetation patterns recorded in land survey records
from the mid-19th century may not represent typical conditions over the
last 2000 years. Restoring pine barrens ecosystem dynamics will require
an understanding of the temporal and spatial patterns of vegetation and fire
dynamics across the barrens region over thousands of years.
LYNCH, JASON A.* and FENG SHENG HU. University of Illinois, Urbana, Illinois. Does vegetation mediate the fire-climate relationships in
boreal regions?
Numerous paleoecological studies have provided information on Holocene
vegetational changes in Alaska. However, the relationship between climate
and vegetation composition on the boreal fire regime is poorly understood,
because few charcoal records exist from that region. We analyzed sediment
from Low Lake (62o 32.8 N, 153o 36.8 W) and Farewell Lake (62o 32.8
N, 153o 36.8 W) in the Alaska Range for lithology, ostracode trace-element
composition, pollen, and charcoal to reconstruct climate, vegetation, and
fire return intervals (MFI). Geochemical (Mg/Ca and Sr/Ca ratios) and
lithological data suggest cold and dry climatic conditions between 13000
and 9500 calibrated years before present (BP), highest growing-season temperatures from 8800 to 5000 BP, and a major increase in effective moisture
between 8500 and 6600 BP. Betula shrub tundra and Populus-Salixwoodland/forest dominated the early-postglacial landscape. Picea glauca arrived
in the region and formed a forest-tundra ecotone by 8500 BP, which was
replaced by closed Picea boreal forests by 8000 BP. Picea mariana replaced P. glauca as the dominant tree species by 6500 BP. The MFI was
.400 yrs before 6600 BP, 170 yrs between 6600 and 2000 BP, and 97 yrs
after 2000 BP. These data indicate that fires were much more common
under the relatively cold and wet climate of the middle and late Holocene.
We hypothesize that around 6600 BP, the establishment of P. mariana
dominated forests increased biomass flammability and fuel availability,
thereby promoting burning. Alternatively, increased frequencies of lightning and thus fire ignition may have come with the altered atmospheric
circulation that led to the colder/wetter climate after 6600 BP. The further
MFI reduction after 2000 BP in the absence of a vegetational change implies that fire-ignition variability was an important factor controlling borealfire response to climate change.
LYONS, KELLY G. Universidad Nacional Autonoma de Mexico, Instituto
de Ecologia, Hermosillo, Sonora, Mexico. Distribution, fitness and ecotypic variation ot the exotic annual grass, Aegilops triuncialis (barbed
goatgrass) on serpentine soil in California.
Colonization and spread of aggressive exotic species is often positively
correlated with anthropogenic disturbance; however, susceptibility of undisturbed habitats remains unclear. In addition, research is sorely needed
on post-establishment selection of exotic species encountering unusual
edaphic environments. Serpentine is a severe soil type that is relatively free
of anthropogenic disturbance and exotic species invasions in California. In
this study, I assessed the process of establishment and adaptation of an
aggressive non-native species, Aegilops triuncialis (Barbed goatgrass), in
serpentine grasslands in California. Reconnaissance of serpentine intrusions
and yearly monitoring suggest that A. triuncialis may grow preferentially
on serpentine soil but advances slowly on undisturbed serpentine sites. In
the greenhouse and field, reciprocal transplant studies were conducted to
assess edaphic adaptation of A. triuncialis. In the greenhouse, serpentine
populations performed better than non-serpentine populations when grown
on serpentine soil. This was particularly true for root to shoot ratios. These
results, however, were not supported in the field, where serpentine and nonserpentine populations did not differ in their performance on novel and
parent soil types. In the field, populations performed equally well on nonserpentine; alluvial, disturbed serpentine and shallow, undisturbed serpentine. Results of these studies suggest that A. triuncialis fitness is highest
on well drained, non-serpentine soils and disturbed serpentine soils but,
even in the absence of previous exposure to serpentine, is capable of establishment and spread onto more severe, intact sites.
VGREN, JOHAN,* MARKO REINIKAINEN and LENNART PERSLO
SON. University of Umea, Umea, Vasterbotten, Sweden. Effects of heterogeneity within trophic levels and resource dynamics on a model
aquatic food web.
The impact of food web complexity in an open and closed community was
explored experimentally under controlled laboratory conditions. We used

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213

an aquatic model community consisting of two forms of primary producers,

phytoplankton (Scenedesmus obtusiuscusculus) and periphyton (Nitzschia
perminuta) and two types of consumers, Daphnia pulex feeding on phytoplankton and Chydorus sphaericus feeding on both periphyton and phytoplankton. Three different food webs all having the phytoplankton (Scenedesmus obtusiuscusculus) and periphyton (Nitzschia perminuta), but having either one of the consumers or both were set up. These food webs were
studied with two different growth regimes for phytoplankton, semichemostatic growth and logistic growth in a factorial design. The experiment
was run for 90 days. D. pulex had a positive effect on the density of C.
sphaericus whereas C. sphaericus in turn had a negative effect on the
density of D. pulex. The mechanism behind the observed pattern was that
D. pulex regenerated nutrients while feeding on phytoplankton which were
utilized by periphyton. The periphyton was, in turn, grazed by C. Sphaericus which increased to very high population numbers and were able to
depress phytoplankton and thereby negatively affect D. pulex. The different
growth dynamics of phytoplankton was important for the strength of the
interaction between D. pulex and C. sphaericus in the sense that the indirect
interactions were more pronounced with semichemostate growth. The results imply that consumer identity, heterogeneity within trophic level and
the growth dynamics of organisms must be taken into account when evaluating food web interactions
MAAS, MARTHA M.* and NORMA L. FOWLER. University of TexasAustin, Austin, Texas, USA. Effects of fungal infection and competition
on a grass.
Individual plants of Nassella leucotricha, a perennial bunch grass common
in central Texas, are sometimes infected by the fungus Atkinsonella texensis. Infected N. leucotricha plants do not set seed; instead, their flowering
culms produce fungal fruiting bodies. However, previous studies have also
indicated that infected plants are on average larger than uninfected plants.
We hypothesized that infected plants are larger because they put resources
into vegetative growth that would have gone into culms, flowers and seeds.
To investigate the effects of infection on plant performance we conducted
a factorial experiment with two levels of infection status (infected or uninfected), two levels of simulated herbivory (unclipped or clipped), and
two levels of interspecific competition (alone or with 2 individuals of the
grass Bothriochloa ischaemum in the pot). Each of the eight treatment
combinations was replicated 20 times, for a total of 160 target plants. We
found that infection status did not affect final total above-ground plant dry
biomass. However, infection status did affect resource allocation: infected
plants allocated a greater proportion of their above-ground dry biomass to
non-flowering culms and associated leaves than did uninfected plants. As
hypothesized, infection by A. texensis evidently diverts resources normally
used by this grass for reproduction into vegetative growth. As expected,
the presence of two B. ischaemum plants in a pot reduced N. leucotricha
total above-ground dry biomass. Competition reduced average total biomass by 63%, and average vegetative biomass by 52%. In contrast, infection increased average vegetative biomass by 55%. Presumably, this infection-induced increase in vegetative size would improve a plants survival
in the future.
MACFARLANE, DAVID W.* and RICHARD KOBE. Department of Forestry, Michigan State University, 126 Natural Resources, East Lansing,
Michigan, USA. Relative growth rate is an ineffective response variable
for capturing size effects on tree growth.
Relative growth rate (RGR) has become the standard response metric for
tree growth analysis. It purportedly allows for size differences between
trees to be "scaled out" so that other factors that influence growth can be
evaluated independently of size. We suggest that RGR is a misleading and
ineffective response variable for capturing size effects on growth, and that
its mathematical convenience and corresponding ubiquity obscure further
progress toward understanding the effects of size, resources, and other environmental factors on growth. We use theory and empirical data sets on
tree growth to demonstrate undesirable qualities of RGR as a response
variable, including: 1) Rather than scaling out size, RGR scales in size
such that it makes size effects intractable and uninterpretable; 2) RGR
could lead to false conclusions regarding resource effects on growth because it confounds size and growth. For example, although foliar N is a

214

Abstracts

significant predictor of absolute growth rate in American beech saplings,

analyzing the growth response as RGR obscures nitrogen effect through
confounding growth and size. 3) In a similar vein, RGR-based models
generally resulted in poorer fits than absolute growth models because RGRbased models are size biased and thus decouple the growth response from
the predictor variables. To remedy these shortcomings of RGR, we advocate alternative models for growth analysis that relate absolute growth rates
to resource supply rates and tree size. These models use size explicitly as
a predictor variable, so that size effects on growth can be directly interpreted, rather than simply divided away.
MACK, RICHARD N. Washington State University, Pullman, WA. Biological invasions, stochastic events and global change: an uncertain
world becomes more uncertain.
Environmental stochasticity, random variation in the environment that affects an entire population (e.g. year-to-year changes in the abundance of a
populations predators and parasites), presents threats to the survival of all
populations; small populations are especially vulnerable. Most immigrant
populations of plants are small. Their extirpation is often caused by random
events that are catastrophic for populations that cannot withstand large
fractional reductions in size across consecutive generations. Predicting
whether any immigrant population will persist is daunting, given not only
the number of chance environmental events, but also the simultaneous influence of a populations size and the diversity and character of its genotypes. These predictions, however daunting, most usefully involve naturally
occurring populations for which the amplitude of expression of environmental factors is known. Climate change, which is only one component of
future global change, could directly and indirectly alter not only the direction but also the amplitude, periodicity, and duration of environmental stochasticity. These climatic changes could substantially increase the difficulty
of predicting the fate of an immigrant population. Computer simulations
of changes in weather that are beyond the range of a locales current regimen offer insight into how novel climatic dynamism could complicate
predictions on the fate of immigrant plant populations. As more species
become incorporated into international commerce, the role of environmental stochasticity in determining their persistence and spread in new ranges
will be shaped by ongoing permanent changes in the global environment.
MACKINNON, DANIEL,* RUTH A. HUFBAUER, ANDREW NORTON
and AARON JACKSON. Colorado State University, Fort Collins, CO.
Preference of an insect biological control agent for two invasive weeds,
yellow and Dalmatian toadflax.
Biological control of invasive plants is typically considered successful
when biological control agents (BCAs) substantially reduce population size
of these weeds. Slowing or halting the spread of invaders, although not as
dramatic or noticeable, can also be considered successful control. Exotic
plants tend to invade by radiation from numerous distinct patches or foci,
not as a single expanding front. The abundance of these satellite foci is
due to dispersal from larger populations, and these smaller patches can be
of greater importance than the larger populations to the spread of invasive
weeds. BCAs that reduce seed set and are good dispersers are valuable in
managing invasions because they can establish on undetected (and usually
smaller and less dense) weed populations that escape other types of control,
decreasing the establishment future infestations. Linaria genistifolia ssp.
dalmatica (Dalmatian toadflax) and Linaria vulgaris (yellow toadflax) are
invasive weeds that use seed for dispersal and establishment. Brachypterolus pulicarius (Coleoptera: Kateridae) is a BCA inadvertently introduced
from Eurasia to North America that reduces seed set in both toadflax species and disperses without assistance. In North America this beetle is abundant on yellow toadflax (and is thought to reduce its spread), however fewer
beetles occur on Dalmatian toadflax. To explore the causes underlying this
pattern, we examined whether populations of this beetle collected from both
host plants differ in their host-plant preference, measured by adult occurrence and oviposition. We found that beetles from both toadflax species
preferred yellow toadflax in laboratory and field experiments. Perhaps due
to this preference, Brachypterolus pulicarius is found on small, isolated
patches of yellow toadflax, suggesting that newly established foci of this
weed have a high probability of attracting this beetle. Studies to find populations of B. pulicarius that prefer Dalmatian toadflax may help stem the
spread of this weed as well.

MADDEN, KATHRYN,1,* REBECCA SHARITZ,1 STEVEN HARPER,1

DONALD IMM2 and BEVERLY COLLINS.1 1 Savannah River Ecology
Laboratory, Drawer E, Aiken, SC, US; 2 USDA-FS Savannah River, PO
Box 700, New Ellenton, SC, US. Tree composition and soil characteristics of Fall Line sandhill communities.
Southeastern Fall Line sandhill communities are often subject to management techniques similar to those used in adjacent or near-by pine communities. To evaluate differences between sandhill community and adjacent
forested community types, 15 sandhill woodlands were selected for survey
at the Savannah River Site in SC and at Fort Gordon and Fort Benning in
GA. Objectives were to identify distinguishing sandhill community level
patterns. Tree composition and soil characteristics were measured at 10
random points within each of the 15 sites. In most of the surveyed sites,
emerging characteristics include the composition of the tree canopy, which
is dominated (56% to 100% of total basal area per site) by a mixture of
four species: longleaf pine (Pinus palustris) and turkey oak (Quercus laevis), with associations of sandpost oak (Q. margaretta) and bluejack oak
(Q. incana). Canopy openness ranges from 20% to 39%. Emerging soil
characteristics include high percentage of sand (86% to 96%), low soil
moisture, and low soil nutrient content. An analysis of variance/Duncan
multirange test was used to determine if tree composition measurements
and soil characteristics differ among the 15 sites. The majority of the sites
demonstrate a range in composition of the four major tree species although,
a few sites were more heavily dominated by other hardwood and pine
species. Soil texture and soil nutrients were similar among sites. These data
are compared with data collected from near-by non-sandhill forests to aid
in distinguishing patterns characteristic of sandhill communities and to aid
in evaluating the effects of various management treatments.
MADRITCH, MICHAEL D.1,2,* and MARK D. HUNTER.1 1 Institute of
Ecology, University of Georgia, Athens, GA, USA; 2 Department of Entomology, University of Wisconsin, Madison, WI, USA. Phenotypic diversity affects ecosystem functioning in the presence of another species.
We have previously demonstrated that intraspecific phenotypic diversity of
leaf litter can influence ecosystem functioning during litter decomposition
in the field. It is unknown whether the effects of phenotypic diversity persist when litter from additional species is present. We used laboratory microcosms to determine whether the intraspecific diversity effects of turkey
oak leaf litter on nutrient dynamics are masked by the presence of naturally
co-occurring longleaf pine litter. We varied the phenotypic diversity of oak
litter (1, 3, and 6 phenotype combinations) in the presence and absence of
pine litter and measured fluxes of carbon and nitrogen over a 42 week
period. Soil C:N ratio peaked at intermediate levels of oak phenotypic
diversity whereas the total amount of DOC leached from microcosms declined as oak phenotypic diversity increased. In no case did the presence
or absence of pine litter influence the response of nutrient fluxes to changes
in oak litter diversity. Our results indicate that phenotypic diversity influences ecosystem function even in the presence of another species, and
suggest that conservation efforts should consider both inter- and intraspecific diversity.
MADSON, STEPHANIE L.* and DAVID C. COLEMAN. University of
Georgia, madsons@uga.edu, Athens, GA. Microclimate conditions within two-age regeneration stands in the southern Appalachians and the
effect on microarthropods.
Soil microclimatic conditions within forest stands have been shown to determine if successful establishment of seedlings will occur, the decomposition rate of organic matter, or the composition of soil fauna communities.
Changes in temperature and moisture have been shown to alter the abundance and distribution of microarthropods within the soil. We assessed the
soil microclimatic conditions within microhabitats established by two-age
regeneration treatments at Wine Spring Creek, North Carolina. We also
correlated microclimatic conditions to soil microarthropod abundance within microhabitats. The majority of the microarthropod taxa showed the greatest response to harvesting when examined at the scale of microhabitats.
The greatest fluctuation in temperature was in the skid rows. During the
warmest period of the day, soil temperature was on average warmer in the
skid rows and cooler at the base of coppices or shrubs. Skid rows had the

greatest bulk density and the least amount of moisture. Microarthropod

abundance was lowest in the skid rows and highest near the base of mature
trees, coppices, and shrubs. Microclimatic conditions within microhabitats
do have an effect on microarthropod community composition and abundance. Examining spatial patterns of microhabitat distribution and frequency across harvested stands may be a useful tool for elucidating treatment
effects within the high natural heterogeneity of microarthropod communities in forest stands.
MAERZ, JOHN C.,1,* BERND BLOSSEY,1 VICTORIA NUZZO2 and
JAME K. LIEBHERR.3 1 Ecology and Management of Invasive Plants
Prorgram, Department of Natural Resources, Cornell University, Ithaca,
NY, USA; 2 Natural Areas Consultants, Richford, NY, USA; 3 Department
of Entomology, Cornell University, Ithaca, NY, USA. Nonnative species
in forest food webs and their impacts on woodland salamander populations.
Using coverboard monitoring plots, we examined the prevalence of nonnative species in forests, and their impacts on woodland salamander populations at 10 sites in the northeastern United States. Numerous nonnative
invertebrates were found at all sites, and many were important prey for
woodland salamanders. Nonnative earthworms were particularly important
at all sites, as they influenced the abundance of several other nonnative
species and had a large effect on woodland salamander populations. Nonnative earthworms were important prey for adult salamanders; however,
high earthworm densities resulted in rapid loss of leaf litter. Decreased leaf
litter levels were associated with reduced abundance of small arthropods,
including some nonnative arthropods, and reduced abundance of salamanders that depend on small arthropods as prey. For two nonnative plant
species studied, garlic mustard (Alliaria petiolata) and Japanese barberry
(Berberis thunbergii), plant invasions were strongly associated with heavy
impacts from earthworm invasions. We could not detect any additional
effects of plant invasions on salamander abundance independent of earthworm impacts. Our research shows that multiple nonnative species from a
range of taxonomic and trophic groups are well established in forest food
webs and impacting a key native generalist predator. Further, this study
illustrates that invasions often occur in the context of other invasions,
which must be considered if we are to accurately assess the impact of a
particular invasive species.
MAHER, ELIZA L.* and MATTHEW J. GERMINO. Idaho State University, Pocatello, ID, USA. Variation in sky exposure and establishment
of conifer seedlings at the alpine-treeline ecotone across three ranges
in the Rocky Mountains.
Factors affecting the recruitment of trees into alpine and subalpine meadows are fundamental to understanding the dynamics of alpine-treeline ecotones (ATE), the transition from forest to alpine meadows. However, little
is known about the factors affecting establishment patterns in the ATE. The
purpose of this research was to determine if and how the range of sky
exposures (%SKY) for conifer seedlings in the ATE varies among three
mountain ranges. %SKY is a quantitative and integrated measure of the
complex canopy (surrounding plants and distance to trees) above seedlings,
and is determined using hemispherical photography. %SKY and surrounding vegetation was observed repeatedly for natural seedlings of Picea engelmannii, Abies lasiocarpa, and Pinus albicaulis found along transects
positioned throughout the ATEs of the Snowy Range, Teton, and Beartooth
mountains. The range of %SKY measured for all species combined differed
among mountain ranges, with the greatest number of seedlings occurring
at high %SKY in the Beartooths, and the least in the Tetons. Greater %SKY
for seedlings in the Beartooths appeared due to a greater relative abundance
of P. albicaulis, which occurred at higher % SKY than the other species.
The range of %SKY for each species also varied among mountain ranges.
Considering this, differences in climate, vegetation community, and unique
biological properties among conifer species may be collectively influencing
the variation seen in conifer seedling distribution in the ATE. Therefore,
these factors may be necessary for more complete explanations of treeline
dynamics that are transferable among these and other mountain ranges.

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215

MALLISON, SHANNON M.,1,* JERRY MILLER2 and PAUL LECHLER.3

1
289 Jennings Dr., Franklin, NC; 2 Department of Geosciences and Natural
Resource Management, Western Carolina University, Cullowhee, North
Carolina; 3 Nevada Bureau of Mines and Geology, University of Nevada,
Reno, Nevada. Assessment of variations in heavy metal accumulation
by Quercus alba: a dendrochemical-lead isotopic approach.
Spatial trends in both total lead (Pb) and Pb isotopes in tree-ring series
have been suggested as an effective means of assessing changes in both
source and abundance of atmospheric lead pollution through time. In this
investigation, total Pb, and selected Pb isotopic ratios were analyzed and
compared for tree-ring series of Quercus alba collected at two heights (0.3
and 4.0 m) within a given tree (5 trees total), and at similar heights between
trees (8 trees total) at a site in western North Carolina, USA. The Pb
isotopic signature of the soil corresponded with the signature of Pb in the
tree rings and, although not statistically significant, mean total Pb concentrations for the lower tree series consistently exceeded the upper series.
These trends indicate that Pb accumulation primarily occurs through the
roots for Quercus alba, and that some Pb is bound to woody materials
during its upward migration. However, the variability of isotopic ratios
within the tree rings was significantly greater than that of the soil, and
206
Pb/208Pb values for the upper series slightly exceeded the lower series of
the sampled trees. Assuming Pb fractionation does not occur, these data
suggest that Pb from an additional source is accumulated through leaves
or bark and bound in the upper portions of the tree. An increase in total
Pb concentrations for most tree ring series was evident beginning in 1930;
however, constant isotopic ratios indicate the source of the lead has not
substantially changed through the time period sampled, 1840-1999. Invariable isotopic ratios are inconsistent with the conclusion that additional
mid-20th century Pb found in these trees is from leaded gasoline. It may
be due, however, to an increase in the availability of Pb associated with a
shift in soil conditions, perhaps a decrease in pH related to acidic precipitation.
MALMSTROM, CAROLYN M.* Michigan State University, East Lansing,
MI. Plant pathogens and vegetation dynamics in natural systems: central questions.
Alterations in pathogen and vector dynamics can cause sudden and dramatic changes in plant community structure. For instance, some of the more
notable responses of vegetation to climate change are likely to be driven
by change in disease epidemiology. Predicting how disease epidemiology
in natural systems will respond to perturbation requires that we have a
baseline understanding of the ecological effects of pathogens. Often we do
not. This presentation introduces the session on plant pathogens in nature
by discussing three questions that are central to our understanding of plant
pathogens and natural vegetation dynamics: (1) How important are plant
pathogens in natural systems? Are they detail agents driving small adjustments in systems shaped by other forces, or more potent regulators? How
much of what has been attributed to plant competition might be better seen
as apparent competition, mediated by plant pathogens? How often do pathogens limit vegetation distributions? (2) Which natural plant pathogen systems do we understand the most, and which the least? Fungal, bacterial,
and viral pathogens differ substantially, yet our understanding of the distributions and roles of different pathogens remains quite limited. It was
only 15 years ago, for example, that researchers found that ocean waters
contain high concentrations of viruses, which may help regulate phytoplankton blooms. (3) When do pathogens drive quick change in plant communities and when do they act as slow sculptors? It is commonly assumed
that the greatest impacts of pathogens occur when new disease organisms
enter susceptible systems, and that over time pathogens and hosts tend to
co-evolve in ways that lead to moderation of disease severity. How often
is this true?
MALONEY, KELLY O.,1,* JACK W. FEMINELLA,1 PATRICK J. MULHOLLAND,2 RICHARD M. MITCHELL1 and LISA M. OLSEN.2 1 Department of Biological Sciences, Auburn University, Auburn, AL; 2 Environmental Sciences Division, Oak Ridge National Laboratory, Oak Ridge,
TN. The legacy of landscape disturbance on small southeastern coastal
plains streams.
We present results of a study designed to assess the relationships between
measures of contemporary stream conditions and historical versus current

216

Abstracts

measures of disturbance from land use. In 1999 we began quantifying instream physicochemical and biotic conditions (streambed organic matter
and coarse woody debris; streamwater NO3 and SRP; benthic macroinvertebrates) within 8 low-order catchments varying in contemporary disturbance intensity at Fort Benning, GA. Land use was quantified from 1974,
1983, 1990, and 1999 remotely sensed imagery using an ArcView extension (ATtILA). Catchment-specific disturbance was estimated as the sum
of the percentage of bare ground and early transitional forest on slopes
.3%, and the percentage of road cover. Contemporary sediment % organic
matter and Bray-Curtis macroinvertebrate community similarity with an
undisturbed reference stream were negatively and more strongly correlated
with historical disturbance. Contemporary measures of macroinvertebrate
functional feeding groups also correlated more strongly with 1974 disturbance conditions than more recent disturbance levels, with % of shredders
decreasing and % of collector-gatherers increasing with increasing catchment disturbance. Contemporary Shannon diversity also reflected historical
conditions, and was negatively correlated with disturbance in both the 1983
and 1990 classifications. In contrast, contemporary measures of streamwater SRP and NO3 concentrations, and streambed % coarse woody debris
abundance were more closely associated with contemporary (1999) than
historical disturbance conditions. Our results suggest that knowledge of
historical land use may be critical in interpreting present-day habitat conditions and macroinvertebrate communities in streams.
MANISH, CHINTAMANI S.* and WILLIAM T. PETERJOHN. West Virginia University, Morgantown, West Virginia. Interactive effects of UV
radiation and N availability on the growth of Quercus rubra seedlings.
A 4 month, 3-way factorial experiment was conducted in a greenhouse to
test whether the sensitivity of Quercus rubra seedlings to UV-B radiation
was affected by N availability and/or the level of UV irradiation. Seedlings
were exposed to 4 levels of UV radiation (UV-A or UV-A+B) in a split
plot design. The 4 levels of irradiation represented a predicted UV-BBE dose
for 2010 (7.02 KJ/m2/day), current levels 6.31 KJ/m2/day), and 2 sub-ambient levels (5.70 KJ/m2/day, and 0.63 KJ/m2/day). Plants were grown in
sand and watered daily with equal amounts of nutrient solutions. The N
concentrations of the nutrient solutions varied (400, 1600, and 4000 mM)
to provide three levels of N addition. UV-B stimulated total biomass production only at the highest level of N addition (p50.038), and this response
is attributable primarily to a stimulation of root biomass (p50.016). Growth
of above-ground tissues was not affected by UV-B. However, the effect of
UV dose, regardless of type (UV-A or UV-A+B), on stem growth depended
on N availability. Plant height (p50.013) and stem mass (p50.042) decreased with increasing irradiance levels but only for plants receiving the
highest level of N addition. This overall irradiance effect on stem growth
may be attributable to increased levels of UV-A since it is a component of
both light treatments. The production of UV-B absorbing compounds in
leaves was consistently stimulated by UV-B (p,0.001) but we found a
trend towards greater production at the lowest level of N addition
(p50.087). Since the sensitivity of red oak seedlings to UV-B (primarily
root growth) and the level of UV-A (stem growth) were apparent only at
the highest level of N availability, this suggests that the response of seedlings to altered light conditions in a regenerating forest may be significantly
altered by changes in N availability.
MARIANO, NESTOR A.1 and RODOLFO DIRZO.2 1 Dept. Ecologia,
Ceamish, Univ. Aut. de Morelos, Cuernavaca, MOR; 2 Instituto de Ecologia, Unam, DF. Herbivory, male and female fitness and compensation
in an annual plant.
Plants responses to herbivory range from undercompensation to overcompensation. Such flexibility of responses is possible by the plants modular
structure. However, few studies of herbivory have analyzed changes in the
demography of plant parts, which in turn determines the plants photosynthetic capacity to maintain growth and reproduction. Such quantitative dynamics may be reflected in qualitative changes in resource allocation to
male and female structures in order to maintain fitness in cosexual species.
We analyzed such effects using artificial herbivory treatments (control,
25%, 75% leaf area removed) on a monoecious annual plant, Cucurbita
sororia, under greenhouse conditions. We measured leaf natality, mortality
and longevity, number of male and female flowers, flower size, pollen

production and number of aborted male flowers. Our demographic analyses

showed that plants with moderate herbivory (25%) increased leaf natality
and longevity, yet, mortality was not affected. Under intense herbivory
(75%) leaf standing crop diminished due to increased mortality and reduced
longevity; unexpectedly, leaf natality was unaffected. Although herbivory
reduced overall flower production and effects were exacerbated in female
flowers, moderate damage did not affect male flower production. Increased
herbivory leads to an increase in the number of male aborted flowers and,
under heavy herbivory, those flowers that did not abort took longer to
develop. Pollen production/flower was affected only under intense herbivory, while corolla size was affected by both levels of herbivory. Considering effects on both growth and reproduction, we found that defoliation
leads to a negative relationship between leaf natality and male flower production. Defoliation had a greater impact on the more costly sexual function. In contrast, the greater allocation to male function seems to lead to a
relaxation of the tradeoffs between growth and reproduction. This may be
a mechanism to compensate for the negative effects of herbivory on plant
fitness.
MARKEWITZ, DANIEL,1,* RICARDO FIGUEIREDO,2 AZENETH
SCHULER,3 JORGE MORAES,3 THOMAS DUNNE4 and ERIC DAVIDSON.5 1 DB Warnell School of Forest Reources, The University of Georgia,
Athens, GA; 2 EMBRAPA Amazionia Oriental, Belem, Para, Brazil;
3
CENA, University of Sao Paulo, Piracicaba, Sao Paulo, Brazil; 4 Bern
School of Environmental Science and Management, University of California, Santa Barbara, Santa Barbara, CA, USA; 5 The Woods Hole Research
Center, Woods Hole, MA, USA. Pathways of dissolved nutrient flux in
a deforested Amazonian watershed.
Over the last four decades many watersheds in eastern Amazonia have been
largely converted from primary forests to degraded or managed pasture
lands. Some of our early work in streams draining one watershed in this
region demonstrated increased cation concentrations in solution during periods of high stream flow, a result contrary to patterns observed in other
tropical and temperate streams. We hypothesized that solution losses of
nutrients in this largely deforested watershed were derived from surface
runoff. To further investigate these pathways of nutrient loss we have instrumented two small watersheds, one in mature forest and another in managed pasture, with surface collectors, soil lysimeters, and groundwater
wells. Using a geochemical mixing model approach in which stream water
concentrations are approximated based on proportional mixing of surface
water, soil lateral flow waters, and groundwater, as well as ancillary stream
and well water data from the larger watershed, we infer important pathways
of nutrient flow. In the case of Ca, for example, volume weighted mean
surface solution concentrations are 45 mM while those of groundwater are
14 mM, two end points around a stream water concentration of 20 mM.
Based on measurements for solution cation concentrations we conclude that
surface and lateral flow pathways contribute important inputs to stream
waters in these deforested tropical watersheds.
MARKO, ADAM C.,* JOSH R. AULD, JASON T. HOVERMAN and
RICK A. RELYEA. University of Pittsburgh, Department of Biological
Sciences, Pittsburgh, PA, USA. Predator cues in aquatic systems: the
dynamics of cue concentration.
Many prey have evolved the ability to detect the presence of predators in
their environment. In aquatic systems, prey have been shown to adaptively
eavesdrop on recent predation events and subsequently adjust their phenotype to better match their environment. Understanding the concentration
dynamics (production and breakdown) of chemical cues can elucidate these
nonlethal aspects of predation and assist in understanding the mechanisms
of predator-induced phenotypic plasticity. We used a common pond snail,
Helisoma trivolvis, to analyze to the characteristics of the chemical cues
associated with predation. By crushing conspecifics we were able to induce
behaviors similar to those induced by crayfish (Orconectes rusticus). We
found that snails are sensitive to cue concentration in a continuous fashion
rather than using a threshold response. Peak responses occurred within 5
hrs and the cue was completely degraded within 14 hrs. Freezing the cue
had no negative impact on the activity of the chemical cue, suggesting that
freezing helped to retard degradation. These results demonstrate that chemical cues can be highly dynamic in aquatic systems and that prey can detect
and respond to a wide range of cue concentrations.

MARKO, MICHELLE D.,* FLORENCE K. GLEASON and RAYMOND

M. NEWMAN. University of Minnesota, Saint Paul, MN. Chemically
mediated host-plant selection by the milfoil weevil: A freshwater insectplant interaction.
To an insect, the air contains a myriad of chemical messages that signal
the location or presence of predators, mates and host plants. In aquatic
systems, the medium of water, rather than air, contains these chemical messages, which are readily broken down by microorganisms. The goal of this
research was to determine whether aquatic insects can use chemical cues
to locate their host plants. We have determined that the native weevil Euhrychiopsis lecontei has made a host-range expansion from the native northern watermilfoil, Myriophyllum sibiricum, to the non-indigenous Eurasian
watermilfoil, Myriophyllum spicatum. The weevil performs better on the
invasive Eurasian watermilfoil and behavioral bioassays indicate that the
weevil uses chemical cues released by Eurasian watermilfoil to select its
hostplant. Lyophilized water from jars containing field-collected milfoil
was used to determine the level of weevil attraction to milfoil exudates
with a dose-response curve (Regression; p,0.001, R250.97). Bioassaydriven fractionation (Y-tubes) of watermilfoil exudates was used to isolate
and identify the attractive fraction. Anion exchange, sizing, and reversephase high performance liquid chromatography were used in the fractionation. Chemical ionization-mass spectroscopy and 1H and 13C nuclear magnetic resonance were used to identify the components of the attractant,
which are common metabolites. Weevil response to each of the components
was determined. This is the first isolation of a water-borne insect attractant
in freshwater systems. Analysis of the watermilfoil-weevil interactions will
provide further understanding as to how insects locate their host-plants both
in aquatic and terrestrial systems and how host-range expansions develop.
MARQUARDT, PAULA E.,1,* CRAIG S. ECHT,2 BRYAN K. EPPERSON3 and DAN M. PUBANZ. 1 USFS-North Central Research Station,
Rhinelander, WI; 2 USFS-Southern Institute of Forest Genetics, Saucier,
MS; 3 Michigan State University, East Lansing, MI. Genetic and spatial
analysis of Eastern White Pine on the Menominee Nation Forest.
To ensure sustainability of forest resources, we must explore the effects of
silvicultural treatments on all aspects of resource management. While we
often focus on easily measured phenotypes, effects at the genetic level have
been largely ignored. Understanding the genetic implications of management practices may assist us in avoiding future bottlenecks. To supply basic
information on white pine genetic diversity for native, managed and locally
regenerated populations, six stands of eastern white pine were evaluated
across the 237,000-acre Menominee Nation Forest, Menominee County,
WI. Five of the sites are actively managed, while in the sixth no records
exist of live pine being cut (SP). Of the five managed stands, one was
regenerated as a white pine plantation (EL), three are under shelterwood
management (PP, C1, OL) and one a pine release management site (MC).
Fifty over-story white pine trees were sampled for genetic analysis at eight
microsatellite DNA loci for each stand except EL, where samples for DNA
analysis were taken from planted stock. In addition, 50 regenerated seedlings were sampled from C1, OL and MC. Genetic diversity was high and
maintained among native and managed sawtimber populations (mean He 5
0.58 + 0.22) and locally regenerated populations (mean He 5 0.56 + 0.23).
The plantation also contained high levels of genetic diversity (mean He 5
0.51 + 0.23). There was less than 2% genetic divergence among all populations (mean Fst 5 0.014). Inbreeding was significantly increased above
Hardy-Weinberg equilibrium for both the parents (mean Fis 5 0.20 + 0.14)
and seedlings (mean Fis 5 0.14 + 0.16). These results indicate some spatial
family structure, which is a major factor in bi-parental inbreeding. Spatial
structure among the sampled trees within each population was weak and
variable, which was not unexpected since the populations have been disturbed and sampled in various ways. The greatest positive spatial structure
was found in the first distance class for SP, the least disturbed population.
Based on the mean Morans I value of 0.008, Wrights neighborhood size
(Ne) for the SP population was approximately 300 trees.
MARR, DEBORAH L.* Indiana University South Bend, South Bend, IN,
USA. The role of fungal pathogens on seed mass variation and seed
production in 3 species of Hydrophyllum.
Many plant species exhibit little variation in seed size, however there are
some species in which seed size varies 10 fold or more. The persistence

Abstracts

217

of this variation is intriguing because both empirical and theoretical studies

predict that seed size should be less variable compared to other reproductive traits because larger seeds generally have higher fitness within species.
I studied seed mass variation in three species of Hydrophyllum, H. appendiculatum, H. canadense, and H. virginianum to better understand the range
of natural variation found in these closely related species. These species
coexist as understory herbs in deciduous forests throughout the midwestern
United States. The range in seed mass produced by individual maternal
plants varied widely among the three species. The greatest amount of seed
mass variation was observed in H. appendiculatum and H. canadense; over
30% of maternal plants in both species produced seeds that varied at least
8 fold in size. However, fewer than 7% of the H. virginianum maternal
plants produced seeds that varied 7 fold in size. Both H. appendiculatum
and H. canadense, were frequently infected by fungal pathogens (Fusarium
sp. and Rhizoctonia sp.) that caused stems and inflorescences to wilt. In
contrast, wilting was not observed in H. virginianum in the study sites. To
test whether seed production and seed mass were affected by the disease
status of the individual in H. canadense, seeds were collected from healthy
and diseased individuals that were hand-pollinated in the field with supplemental pollen. Diseased individuals produced significantly fewer fruits,
but no difference in seed mass variation between healthy and diseased
maternal plants was found. Further studies of all three species in which
maternal environment is controlled will help determine whether pathogens
that cause wilt contribute significantly to the unusual seed size variation in
Hydrophyllum.
MARRS, ROBIN A.,* RUTH A. HUFBAUER, HARSH PAL BAIS and
JORGE M. VIVANCO. Colorado State University, Fort Collins, CO, USA.
Effects of hybridization on allelochemical expression in invasive Centaurea.
Non-indigenous species become ecologically injurious when they displace
species native to the areas they invade and alter patterns of productivity in
an ecosystem. It is still fundamentally unclear which attributes enable benign pioneers to become aggressive invaders. In some cases hybridization
or allele frequency shifts can lead to increased invasiveness. Centaurea
maculosa and C. diffusa (Asteraceae, spotted and diffuse knapweed) are
native to Eurasia and have become highly invasive in North America, causing ecological and economic damage. These species have taken over millions of hectares of rangeland and wildland since their introduction and are
capable of invading undisturbed plant communities. Several different mechanisms may facilitate this displacement, including negative chemical interactions between species or allelopathy. These invaders each exude a phytotoxic compound into their rhizosphere; C. maculosa exudes (-)-catechin
while C. diffusa exudes a-hydroxyquinoline. Each species is immune to the
phytotoxic effects of its own exudate but is susceptible to the other species
toxin. Interestingly, the two species are capable of hybridization. Hybrid
individuals are intermediate in morphology between spotted and diffuse
knapweeds, but little is known of their ecology and physiology. Which
exudate is being produced by the hybrid individuals? They could be producing both compounds, just one, or neither. To address this question, seeds
were collected from a hybrid swarm population in Hood River, OR, where
morphological phenotypes ranged from "pure" C. maculosa to "pure" C.
diffusa, with all gradations in between. These plants were grouped into five
categories depending on how many characters they shared with either
"pure" species. Three offspring from five plants in each of the five categories are being assayed for production of (-)-catechin and hydroxyquinoline. This research will help elucidate the role of allelopathy and hybridization in the invasion of exotic plants.
MARSCHALL, ELIZABETH A.1,* and JAMES E. GARVEY.2 1 The Ohio
State University, Columbus, OH; 2 Southern Illinois University, Carbondale, IL. Understanding latitudinal trends in fish body size through
models of optimal seasonal energy allocation.
For fish at high latitudes, short growing seasons should constrain size-atage, although the converse often occurs. We used a dynamic-state variable
model to find energy allocation strategies to length, fat, and ovaries that
maximize expected egg production of largemouth bass Micropterus salmoides. We determined how latitudes and rations affect optimal allocation
and then simulated growth using optimal strategies. A theoretical reciprocal

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transplant explored how latitude-specific optimal strategies affected growth

at other latitudes. At low ration, allocation and growth were similar among
latitudes, with length selected in small individuals and reproductive tissue
and fat in large counterparts. At high rations, low-latitude fish invested
most energy to length and reproduction; high-latitude fish allocated to
length during summer and fat during fall, and developed ovaries earlier in
the year. Transplants revealed that smaller size-at-age occurs in the north
than the south, consistent with field patterns for largemouth bass. Although
northern strategies allowed fish to be successful in the south, southern
strategies were unsuccessful in the north. Latitude-specific energetic adaptations may compromise success of fish transplanted beyond their native
distribution.
MARSE, TODD J.* and JULIE L. WHITBECK. University of New Orleans, New Orleans, LA, USA. Assessment of aboveground litter decomposition rates across a hydrologic gradient in bottomland hardwood
forest.
We estimate aboveground litter decomposition rates across a 1.0-m elevation gradient in mature bottomland hardwood forest in Jean Lafitte National
Historical Park and Preserve, Jefferson Parish, Louisiana, USA. Hydrology
changes significantly along this gradient, such that the high elevation is
rarely flooded, while the low elevation experiences standing water for several months each year. We hypothesized that litter decomposition rate
would be slowest at the wet end of this gradient, and that dry summer
conditions at the high elevation site would also diminish the rate of decomposition. To assess aboveground decomposition rates, we measured litter mass change over time by placing known quantities of oven-dried litter
into fiberglass screen bags and deploying the bags on the forest floor for
varying exposure intervals. We placed groups of litter bags at five randomly
chosen replicate locations at each of three sites (ridge, intermediate,
swamp) along the hydrologic gradient. To estimate initial decomposition
rates, we collected litter bags each month for six months, then bi-monthly
to one year. We plan additional sampling at 15, 18, and 24 months after
installation to assess longer-term rates. The data for the first three months
of collection indicate a steady decrease in mass at the wettest site (swamp)
with a decomposition rate of 0.03 g dw per day. We observed an unexpected increase in mass during the first 60 days at the drier sites (ridge and
intermediate), which was then followed by a decrease similar to that observed at the swamp site. Close inspection of litter collected from the ridge
and intermediate sites suggests that one or more organisms may be using
the leaf litter at these sites as a substrate, adding mass to the litter.
MARSH-MATTHEWS, EDIE,* MELODY BROOKS and RAELYNN
DEATON. University of Oklahoma, Norman, OK. Matrotrophic provisioning: A bet-hedging mechanism in mosquitofish?
Livebearing fishes in the genus Gambusia provision offspring both by production of large, yolky eggs and by mother-to-embryo transfer of nutrients
(matrotrophy). We assayed nutrient transfer in Gambusia affinis and G.
geiseri using radiolabeled leucine. In G. geiseri, matrotrophic provisioning
was typically normally-distributed within broods. Provisioning to individual embryos was not a function of embryo size. In G. affinis, nutrient
transfer was typically positively skewed within a brood, with higher levels
of provisioning to larger embryos. In G. affinis, skewness was higher in
broods at later stages and in broods with smaller median size of embryos.
Differences between species in matrotrophic provisioning may reflect tactics evolved in environments that differ in predictability. Gambusia geiseri
is restricted to high-discharge, constant-temperature springs, so matrotrophic provisioning may represent a tactic to optimize offspring quality. Gambusia affinis occupies a variety of habitats over its extensive range, including those with extreme and unpredictable conditions. Highly-skewed matrotrophic provisioning in G. affinis is consistent with a bet-hedging tactic
to produce offspring of variable quality in unpredictable environments.
MARSHALL, JOHN D.,1 GREG GAFFORD-BARRON,2 KATIE
GRIEVE2 and RAMESH MURTHY.2 1 Dept. of Forest Resources, University of Idaho, Moscow, ID, USA; 2 Biosphere II, Columbia University, Tucson, Arizona, USA. Toward an isotopic mass-balance for carbon for
cottonwood (Populus deltoides) plantations grown under CO2-enrichment inside Biosphere II, Arizona.
Rates and stable carbon isotope composition (d13C) were measured for major fluxes to and from fast-growing cottonwood plantations in three sealed

bays at Biosphere II near Tucson, Arizona. Air and tissue measurements

were made at a range of CO2 concentrations 400, 800, and 1200 ppm. The
soil flux was measured from PVC rings with a Li-cor 6200 and d13C of
soil respiration was measured on air equilibrated with the soil atmosphere
in inverted 12-L dishes. We measured d13C of soluble carbohydrates in
leaves to estimate the isotopic composition of recent photosynthate. Leaftissue sampled in August had d13C averaging -31.4, -40.4, and -44.4 (SE
5 0.3) for the 400, 800, and 1200 ppm treatments, respectively. The
pure CO2 added to the bays, when sampled in October, had an average
d13C of -10.6 (SE 5 0.6) . The atmospheric d13C was surprisingly similar.
Soil respiration showed a small but consistent difference between the bays.
By combining these estimates of fluxes and d13C, we hope to construct a
closed isotopic mass balance for carbon inside each of the bays.
MARTIN, ANDREW,1,* LUCY RUBINO2 and J D. LEWIS.1 1 Calder Center, Fordham University, Armonk, NY; 2 Drew University, Madison, NJ.
Hemlock woolly adelgid density affects net photosynthetic rates but not
respiration rates or needle biochemistry in eastern hemlock.
Invasive, non-native insects and pathogens are a significant threat to native
forests. Currently, forests from North Carolina to Massachusetts are experiencing striking declines in eastern hemlock (Tsuga canadensis) associated with the invasion of the hemlock woolly adelgid (Adelges tsugae).
Although this invasion may dramatically alter carbon and nutrient cycling
in affected forests through effects on resource uptake and allocation within
infested trees, relatively little is known about the effects of the hemlock
woolly adelgid on hemlock physiology or biochemistry. This study examined the effects of differences in hemlock woolly adelgid density on
needle biochemistry and photosynthetic rates. Net photosynthetic rates declined by 30% as the average density of hemlock woolly adelgid increased
from zero to 0.5 adelgid per needle. However, net photosynthetic rates did
not significantly vary as adelgid density increased from 0.5 to two adelgid
per needle. Further, needle nitrogen and chlorophyll concentrations did not
significantly vary with adelgid density. Similarly, quantum yield, light compensation points, and daytime respiration rates did not significantly vary
with adelgid density. These results suggest that eastern hemlock may have
the capacity to partially compensate to low levels of hemlock woolly adelgid infestation, but not to high levels of infestation. In addition, effects of
adelgid density on net photosynthetic rates appear to be independent of
any effects on respiration rates, and do not appear to be driven by effects
on needle nitrogen or chlorophyll concentrations.
MARTIN, JENNIFER L.,1 STITH T. GOWER1 and JENNIFER PLAUT.2
University of Wisconsin-Madison, Madison, Wisconsin; 2 Brown University, Providence, Rhode Island. Measuring and modeling interspecies
competition of tree species in logged boreal mixed forests.
1

Succession in the boreal forest is challenging to predict because of the

frequent disturbances that alter successional trends. Some studies have
found that coniferous forests regenerate to deciduous forests following logging, yet many boreal forest stands are mixed, containing both evergreen
and deciduous species. Forest tree species strongly affect biogeochemical
cycles, yet most biogeochemical field studies and process models ignore
these mixed stands. The objective of this study was to determine, through
time, the successional trends and net primary production of dominant tree
species following logging in boreal forests of northern Manitoba. The experimental design consisted of eight replicate plots in four different-aged
stands that originated from clearcut harvests in 1935 (mature), 1971, 1983,
1990. Six trees of each major tree species were harvested and diameter
height growth relationships were determined by analyzing radial growth of
stem disks collected every 0.25 m up the stem. Trembling aspen (Populus
tremuloides Michx.) was the dominant tree species in the younger stands
(1983, 1990) while jack pine (Pinus banksiana Lamb.) and black spruce
(Picea mariana (Mill.) were the dominant tree species in the older stands
(1935, 1971). For each dominant species, diameter:height growth relationships were determined. Annual growth of jack pine and trembling aspen
decreased with age, whereas growth of black spruce increased. The relationship of annual radial growth with stand age for the three species is
discussed in relation to canopy architecture and biogeochemical cycles.

MARTINEZ, JEANNETTE C.* University of Minnesota, Dept. of Ecology, Evolution & Behavior, 100 Ecology, St. Paul, MN, USA. Habitat
specificity and population genetics of three dragonfly larvae in fragmented landscapes.
While anthropogenic land use change is a major component of environmental change and often results in fragmentation of habitat into isolated
remnant patches, habitat specialization is another important mechanism that
may lead to population patchiness. My research investigates the sensitivity
to extinction by fragmentation in three dragonfly species with differing
degrees of habitat specialization, a habitat generalist (Libellula quadrimaculata,), intermediate habitat specialist (Somatochlora williamsoni), and
habitat specialist (Somatochlora tenebrosa). This is based on principal
component analysis using multiple habitat variables. I combine population
genetics, ecology, and a modeling approach to gain a more complete understanding of species9 sensitivity to extinction by fragmentation. My study
species appear to exhibit metapopulation dynamics in the field. Preliminary
results of my metapopulation model reveal that when female adults disperse
into demes at low population densities, they provide a rescue effect because
female immigrants carry thousands of potential eggs with them. On the
other hand, male adult dispersal may generally contribute more to gene
flow between demes because of their ability to mate with multiple females
(pre-mating dispersal model). Preliminary population genetic results for the
intermediate habitat specialist revealed no population subdivision on a spatial scale of 300 miles when allozymes were used as molecular markers.
MARTINEZ, M. LUISA.* Departamento de Ecologia Vegetal, Instituto de
Ecolog
a A.C., Xalapa, Veracruz, Mexico. Climate change and community dynamics change on coastal dunes.
Studies on global climate change predict that sea level rise will lead to
reduced sediment supply to the beach-foredune interface and therefore,
alterations in substrate mobility in inland dunes. This will probably influence natural dynamics of dune vegetation since they are largely affected
by (and in many occasions dependent upon) substrate mobility. In a tenyear survey on primary succession occurring in a mobile dunes area (along
the coast of Central Gulf of Mexico) it was observed that sand movement
played a key-role in species replacement, especially during the early successional stages. The early colonizer (the shrubby legume Chamaecrista
chamaecristoides) was the species most tolerant to burial by sand, and
seemed to facilitate colonization by late colonizers (tall grasses such as
Trachypogon plumosus and Schizachyrium scoparium). Subsequent field
observations and experiments demonstrated the occurrence of facilitation:
species were spatially aggregated; environmental conditions (wind speed,
sand movement, and temperature on the sand surface) were significantly
ameliorated by the shrub, and seedling survival of the two grasses was
exclusive to the shade of the shrub. The above is evidence of the importance of sand movement for community dynamics. How will it thrive in a
scenario of global climate change and altered substrate ability? Markov
chains provide a useful tool to predict potential community dynamics and
species replacement in two contrasting scenarios. With reduced sand movement, it is predicted that early colonizers might disappear and become
locally extinct, which will be problematic for several reasons: the maintenance of the natural vegetation dynamics will lack its naturally restoring
species which is keystone after disturbance events; in addition, it is a conservation issue, since Chamaecrista is endemic to the Mexican mobile
dunes. With exacerbated sand movement, early successional stages will be
maintained and probably the system will remain unstable, negatively affecting late successional stages.
MARUCA, SUSAN,1,* ELI MEIR,1 DEREK STAL1 and CHRISTINE
REICH.2 1 SimBiotic Software, Inc., Ithaca, NY; 2 Museum of Science Boston, Boston, MA. Teaching ecology in a science museum using an
EcoBeaker simulation.
We developed a simulation model using the EcoBeaker software framework
to teach principles of disease dynamics and control to guests of the Museum of Science (MoS), Boston. The simulation, which allows users to
manage their own fish farm and implement various strategies for controlling disease outbreaks, is part of MoSs upcoming exhibit on using models
as tools for scientific exploration and discovery. We will present a synopsis

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219

of development, a demonstration of the simulation software, usage data and

assessment results from museum guest users, and a discussion of the future
of using simulation models for teaching ecology in outreach educational
settings.
MARUSSICH, WENDY A.* wendy.marussich@asu.edu, Arizona State
University, Tempe, AZ. The effects of five ant species on the germination of Datura wrightii.
Myrmecochory, or seed dispersal by ants, is often mediated by the presence
a lipid-rich appendage, or elaiosome, on a seed. Typically, ants gather
seeds, carry them to the nest, consume the elaiosome, and discard the seeds
unharmed either in the nest or refuse pile. The benefit to the ants is presumably the nutritional content of the elaiosome, whereas benefits to the
plant includes dispersal from the parent plant, protection from predators,
reduced seedling competition, protection from fire, transportation to nutrient-rich microsites, and enhanced germination due to seed scarification. I
tested whether seed scarification by ants enhances the germination of Datura wrightii by presenting D. wrightii seeds to field colonies of five ant
species (Pogonomyrmex rugosus, P. californicus, P. maricopa/occidentalis,
Aphaenogaster cockerelli, and Messor pergandei) and conducting a germination experiment with the seeds collected from the refuse piles of the
ant colonies. I also measured the percent of seeds destroyed by the different
ant species and the percent of missing seeds. Ant species did not affect
number of days to germination, plant height, or number of leaves per plant.
However, the total number of seeds germinated differed among ant species.
Pogonomyrmex maricopa/occidentalis enhanced germination, whereas
Messor pergandei most severely reduced germination rates. Benefits from
scarification by ants do not explain the presence of elaiosomes in Datura
wrightii. The reciprocal effects of ants and plants are explored with a mathematical model of seed fates.
MARX, LAURA* and MICHAEL WALTERS. Michigan State University,
East Lansing, MI. Decaying wood species composition and distribution
limit eastern hemlock and yellow birch seedling establishment.
Eastern hemlock (Tsuga canadensis) in Upper Michigan is usually found
in distinct patches of either pure hemlock or hemlock in association with
yellow birch (Betula allegheniensis). These patches often border areas of
northern hardwood forest. The distribution and species composition of
coarse woody debris (decaying logs, stumps, and large branches) help
maintain the eastern hemlock-northern hardwood patch structure and the
hemlock/yellow birch association. In four old growth hemlock-hardwood
forests, we found 93% of hemlock and yellow birch seedlings growing on
coarse woody debris, though this substrate covered only 3% of the forest
floor. Of the seedlings found on coarse wood, 62% were on conifer wood
(which made up 43% of total wood area), and average seedling density on
conifer wood was 3.1 6 0.67 s.e. seedlings/m2 while hardwood wood supported only 0.84 6 0.29 seedlings/m2. Conifer coarse wood was more abundant in hemlock patches than in mixed hardwood patches (paired t-test, N
5 7 paired forest patches, p 5 0.0492). The lack of suitable coarse wood
for establishment of hemlock seedlings in northern hardwood stands helps
to maintain the hemlock patch structure. Similarly, the restriction of hemlock and yellow birch seedlings to the same rare substrateconifer coarse
woodmaintains the strong association between adult hemlock and yellow
birch trees. Differences in wood chemistry may partially explain why conifer coarse wood is more suitable than hardwood coarse wood for seedling
establishment. Conifer coarse wood has modestly higher supply rates of
ammonium (paired t-test, n 5 13 field plots, p , 0.10) and lower pH (p
, 0.01) than hardwood coarse wood.
MASON, ERIN E.,* KATY M. KAMPF and ANDREW M. JAROSZ. Department of Plant Biology, Michigan State University, East Lansing, MI,
USA. Life history changes in Arisaema triphyllum due to pathogen
infection.
Habitat fragmentation is a known risk to plant populations, particularly
those with limited migration capability. The presence of a disease that
lessens individual fitness may compound the problem and lead to increased
local extinctions. Pathogen infections can influence plant life histories by
reducing growth and survivorship, and by altering the amount and pattern

220

Abstracts

of reproduction. Since 1999, we have monitored six populations of Arisaema triphyllum, a perennial forest herb, to investigate the impact of infections by Uromyces ari-triphylli, a systemic rust fungus. Effects on death
and dormancy are correlated with host size. Among smaller plants, diseased
individuals displayed higher rates of dormancy and mortality than nondiseased individuals. This trend diminishes as plant size increases. Infection
decreases growth rates across all plant sizes. Both infected and uninfected
individuals are able to reproduce asexually, yet asexual offspring of diseased plants have a lower survivorship. Disease directly limits a plants
sexual reproduction by reducing consecutive years of flowering and lowering seed output. Our data indicate that U. ari-triphylli infection consistently reduces host fitness by reversing growth patterns, limiting reproduction, and elevating rates of dormancy and mortality. Such effects on individual plants can influence entire populations. High disease incidence can
alter the size, structure, and persistence of local populations. Our demographic data indicates that the size of all populations is decreasing over
time, suggesting that fragmentation or some other environmental variable
is negatively influencing these populations. High incidence of disease exacerbates this trend and may increase localized extinctions or endanger the
species as a whole.
MATES-MUCHIN, JONATHAN,1 JAMES D. LEWIS,1,* GREG D.
TURNER,1 JEANNINE RICHARDS1 and WILLIAM SCHUSTER.2 1 Fordham University, Louis Calder Center, Armonk, NY; 2 Black Rock Forest
Consortium, Cornwall, NY. Variation in growth rate and above and
below-ground nitrogen of a hemlock-hardwood forest with the loss of
a dominant tree species.
The loss of dominant species due to invasive parasites may significantly
alter the relationship between above and below-ground carbon and nitrogen. Yet few studies have assessed how nitrogen is redistributed among
remaining species after the decline of a dominant species, and whether
changes in species composition and nitrogen may differentially alter short
term growth rates of coexisting species. Eastern hemlock (Tsuga canadensis) has declined over the past 15 years in eastern forests, associated with
the range expansion of the hemlock woolly adelgid (Adelges tsugae). We
hypothesized that eastern hemlock decline will cause short term increases
in total soil nitrogen availability and in total nitrogen in trees associated
with hemlock. Additionally, as the eastern hemlock declines due to mortality, we hypothesized that hemlock associates would show increased
growth rates linked with the competitive release and greater nitrogen availability. To test these hypotheses, we established nine 0.16 ha plots across
a gradient of hemlock dominance. Annual change in DBH between 1999
and 2002 was measured on all trees within each plot. Soil, live leaf and
leaf litter samples were collected in all plots, dried and ground, and analyzed for total carbon and nitrogen. The average of three tree cores from
red oak (Quercus rubra), chestnut oak (Quercus prinus) and hemlock were
used to determine long-term patterns in growth. Soil nitrogen concentration
was 10% higher on average in hemlock dominated compared to mixed
stands. N concentration was 4% greater in hemlock than in oak leaf litter.
Additionally, chestnut oak leaf litter N concentration was significantly
greater from hemlock compared to hardwood plots. Finally, hemlock
growth rate from 1997-2002 decreased by 65% relative to the average
growth rate while the average oak growth did not significantly change. With
the loss of eastern hemlock, there may be a shift in nutrients from decaying
trees to other healthy tree species, which may account for an increase in
the productivity of chestnut oak trees, which have shown greater amount
of leaf litter N.
MATHIAS, ANDREA* and PETER CHESSON. Univeristy of California,
Davis, Davis, CA, USA. Quantifying species coexistence from endogenous consumer resource cycles with relatively non-linear competition.
Theoretical research suggests that population density fluctuations may promote the coexistence of competing species. Such fluctuations can arise not
only as the consequence of environmental variability but also as the result
of endogenous population dynamical mechanisms. Despite the recent increase of interest in the role of endogenously generated population cycles,
the latest investigations remain descriptive and fail to disentangle and quantify the primary causes leading to coexistence. We analyze a model of two
consumers coexisting on a single resource with endogenous fluctuations.

The method uses a general framework that allows for separation of coexistence promoting mechanisms into well-defined groups and quantification
of their effects. The focal mechanism is relative non-linearity of competition, which is partitioned quantitatively into stabilizing and equalizing
mechanisms. The method uses a quadratic approximation to the long-term
low-density growth rate of a population, and works best for small fluctuations. However, an exact nonlinear analysis extends the qualitative conclusions of the quadratic approximation while correcting quantitative detail.
The above results can be extended for the case of multiple resources.
MATTHEWS, WILLIAM J.* and EDIE MARSH-MATTHEWS. University of Oklahoma, Norman, OK. Consistency of fish species associations
in stream pools: Implications for local coevolution.
Consistent patterns in contacts between species should promote coevolution
of species traits. Coevolutionary interactions have been assumed to require
long periods of intimate contact between species, but recent evidence shows
that coevolutionary adjustments can occur within decades, making interspecific contacts in ecological time important in understanding trait changes
in evolutionary time. The question of persistence of species contacts in
ecological time has been largely ignored in the literature. Many studies
have assessed changes in animal assemblages at fixed sites, but few have
explicitly tested persistence of species associations in complex communities. We visually surveyed fish species, and size-structured ecospecies, on
11 dates from 1995 to 2002 in 14 pools in a perennial reach of a small
stream, and calculated matrices of similarity in qualitative and quantitative
contacts among taxa on each date. We then used Mantel tests to assess
consistency of contact patterns between surveys at intervals of months and
years, and across four physical disturbances (floods and droughts). We used
data from similar surveys in 1982-1983 to test long-term consistency in
species associations. Associations of species and ecospecies associations in
direct contact were consistent across many, but not all, intervals between
surveys and were highly consistent between decades. Such consistency of
contacts among these taxa could promote coupled coevolution between
species pairs or diffuse coevolution within the community, and may be
applicable to a broader geographic mosaic model of coevolution that has
been proposed in recent literature.
MATTICE, JENNIFER A.,1,* DUANE R. DIEFENBACH1 and DANIEL
W. BRAUNING.2 1 PA Cooperative Fish and Wildlife Research Unit, Pennsylvania State University, 113 Merkle Laboratory, University Park, PA,
USA; 2 Pennsylvania Game Commission, 2001 Emerton Avenue, Harrisburg, PA, USA. Grassland songbird density and occupancy rates on
reclaimed surface mines in western Pennsylvania.
Widespread habitat loss, fragmentation, and consequent population declines
have drawn considerable conservation and research attention to grassland
bird communities. In order to assess the contribution of large tracts of
grassland habitat on reclaimed surface mines in western Pennsylvania to
global grassland bird conservation, we used distance sampling methods to
estimate the density of three obligate grassland species. Henslow9s (Ammodramus henslowii), savannah (Passerculus sandwichensis), and grasshopper sparrows (Ammodramus savannarum) had densities of 14, 7, and
28 singing males/100 ha, respectively. We also investigated patch-level and
landscape-level variables that could influence occupancy rates and abundance for these birds, including total patch area, patch core area, patch
vegetation, proximity to other reclaimed patches, and surrounding habitat
types. Our 102 patches ranged in size from 1 to 120 ha, and were between
3 and 25 years old. Patch area was a predictor of patch occupancy for all
three species. We found occupancy threshold values (the patch size at
which we found birds 80% of the time) to be 15 ha for grasshopper sparrows, 26 ha for Henslow9s sparrows, and 40 ha for savannah sparrows.
Succession proceeds slowly on reclaimed surface mines, and western Pennsylvania will likely provide suitable grassland bird habitat for decades.
Therefore, management of reclaimed strip mine areas as grasslands may
help mitigate overall declines in grassland songbird populations due to
habitat loss. We will also discuss other factors influencing grassland bird
abundance and occupancy on reclaimed mines.

MAUN.* University of Western Ontario, London, Ontario. The effect of

sea level rise on the vegetation adapted to burial along the coast.
During their evolution along coastlines, the beach and foredune species in
different continents have adapted to live under conditions of high wind
velocities, burial by sand, salt spray, sand blast and wave action. Such
shoreline habitats have existed throughout the Tertiary (65 million years
ago) or even since the Cambrian and Proterozoic periods. The changes in
global temperatures over time would have only forced the migration of
species northward or southward but the habitat has essentially remained
unchanged over these years. A rise in sea level may possibly have one or
more of three effects on burial tolerant vegetation. First, a complete submergence of a dune system will lead to the destruction of all vegetation.
However, this vegetation may not be completely lost because propagules
(fruits, seeds, rhizome or stolon fragments) of many of these species have
the ability to disperse rather long distances to other shorelines where they
may find conditions suitable for re-establishment. Second, the rise in sea
level may only cause shoreface erosion and sand may start to move inland.
In such a scenario, species intolerant of burial will be killed; tolerant species, if any, will start to increase in coverage; and pioneer species will start
to re-establish. Examples of this case can be found in areas where blowouts
are advancing inland. Third, if sea level rise creates conditions under which
no sand is being deposited on the shoreline, the burial tolerant species on
higher ground will eventually be replaced by non tolerant species during
succession.
MAY, DAVE Z.,1,* KIM J. BROWN2 and STEPHANIE A. BOHLMAN.3
Ohio University, Department of Geography, dave.may@ohio.edu, Athens,
OH, USA; 2 Ohio University, Department of Environmental and Plant Biology, kim.brown@ohio.edu, Athens, OH, USA; 3 University of Washington, College of Forest Resources, bohlman@u.washington.edu, Seattle,
WA, USA. Evaluation of high-resolution, multi-band imagery for determining proportions of oak and maple LAI in Black Rock Forest,
NY.
1

Spatial estimates of leaf area index (LAI) are difficult to obtain but are
essential in estimating ecosystem productivity. Remote sensing technology
can be an effective tool for measuring LAI in simple systems, but complications arise if LAI .;5 or if canopy structure is complex. Highly
productive deciduous forests possess species and structural heterogeneities
that make LAI difficult to quantify through remote sensing methods. Ideally, remote sensing technologies could be used to determine tree speciesspecific differences in productivity. Tree- or stand-based relationships for
predicting LAI are often used, but are site-specific, labor intensive, and
highly variable. At the end of the growing season in deciduous systems,
leaf litterfall traps allow for the accurate assessment of species specific leaf
area (LAI(spp)), but the labor, temporal, and spatial sampling issues make
remotely sensed techniques attractive. The goal of this research is to utilize
multi-band high-resolution remote sensing imagery to determine key species groups (Acer rubrum, Quercus rubra, and Quercus prinus) during the
2000 growing season in the forest canopy at Black Rock Forest, NY. Spectral reflectance data were obtained at the level of the leaf, branch, and forest
canopy at two sites exhibiting differences in water availability and topography (Turnbull et al. 2002). These multi-spatial spectral reflectances were
then integrated into a classification to quantify the percentage of canopy
cover that the species of interest occupy within the forest. Correlation statistics were performed using LAI(spp) and percentages of canopy cover at
both sites. Comparisons of remote methodology versus ground-based
LAI(spp) will be presented, to determine if the technique will be useful for
more rapid and accurate estimations of LAI(spp) for eastern deciduous forests.
MAYER, AUDREY L.,1,* CHRISTOPHER W. PAWLOWSKI2 and HERIBERTO CABEZAS.1 1 U.S. Environmental Protection Agency, Cincinnati,
OH; 2 U.S. Environmental Protection Agency, ORISE Research Fellow,
Cincinnati, OH. Using Fisher Information to detect gradual and rapid
ecosystem regime shifts.
As ecosystems experience perturbations of varying regularity and intensity,
they may either remain within the state space neighborhood of the current
regime, or "flip" into the neighborhood of a regime with different charac-

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221

teristics. Although the possibility of such regime shifts needs to be incorporated into informed environmental monitoring and management efforts,
the identification of regime shifts typically requires an intimate knowledge
of the ecosystem dynamics as well as extensive supporting data. We demonstrate the utility of a dynamic information index that lowers the threshold
of knowledge required to detect shifts in ecosystem regime. Information
theory can be used to quantify the order inherent in systems in spite of
imperfect observations or signals from the source system. We use the
Fisher Information index to develop a measure of the variation of a dynamic steady state based on a generated probability density function. Periodic systems that spend more time in a few segments of their trajectory
have a high Fisher Information, whereas periodic systems spending an
equal amount of time in all the segments of their trajectory have zero
information. This form of the Fisher Information is a measure of system
order. A transition from one periodic regime to another appears as a change
in Fisher Information. We calculate changes in Fisher Information from
time series data for ecosystems at several scales (Lake Erie, the Pacific
Ocean, and the global climate system), and demonstrate that the differences
in Fisher Information behavior are correlated with regime shifts. Fisher
Information changes more gradually for systems undergoing a gradual shift,
whereas the index changes rapidly (either positively or negatively) for rapid
shifts. Fisher Information can be calculated for an unlimited number of
variables, although including spurious variables that are not associated with
ecosystems dynamics decreases the sensitivity of the index to regime shifts.
MAYFIELD, MARGARET M.,* DAVID ACKERLY and GRETCHEN C.
DAILY. Stanford University, mmayfield@stanford.edu, Stanford, CA,
USA. Functional diversity of herbaceous and shrubby plants in countryside landscapes of southern Costa Rica.
In recent years, increasing attention has been directed toward the study of
human mediated biodiversity loss in the tropics. However, most of these
studies focus on species richness and composition without considering other dimensions of diversity. One dimension of growing interest to the scientific and conservation communities is functional diversity. Unlike species
richness, studies of community functional diversity can provide valuable
information about ecosystem processes and resource pools available to other organisms in the community. For example, the presence of fleshy fruits
may be of major importance to the survival of certain bird and mammal
species in deforested areas regardless of the species producing the fruits.
Knowledge of functional diversity is extremely valuable for understanding
the ecological importance of individual species in vulnerable communities
and will improve our understanding of how ecosystems are changing in
human-altered landscapes. In this study we examine the functional diversity
of herbaceous and shrubby plant communities in 10 common habitats in
human-dominated landscapes of southern Costa Rica. These habitats include pasture, roadside vegetation and primary forest. We focus on simple
physiological traits such as leaf area, growth form and SLA, as well as
plant traits associated with animal activities including pollination, seed dispersal and herbivory. We ask two basic questions: 1) has functional diversity been lost or gained in deforested habitats of southern Costa Rica? And
2) does the level of redundancy in the functional traits found in forested
and deforested habitats differ? We found that most deforested habitats had
fewer functional traits present than forested habitats. However, the abundance of species with specific functional traits was generally higher in
deforested than in forested habitats.
MAZIA, NOEMI C.,1 ENRIQUE J. CHANETON1,* and THOMAS KITZa, Universidad de Buenos Aires, chaBERGER.2 1 Facultad de Agronom
neton@ifeva.edu.ar, Buenos Aires, Buenos Aires, Argentina; 2 Universidad
Nacional del Comahue, Bariloche, Rio Negro, Argentina. Facilitation vs.
apparent competition and the establishment of tree seedlings beneath
nurse shrubs in northern Patagonian.
Facilitative interactions play a major role in plant recruitment in stressful
environments. Yet, negative indirect effects mediated by herbivorous consumers can potentially mask the effects of facilitation between neighbouring plants. Our objective was to evaluate how seedling predation by tenebrionid beetles affects the likelihood of tree establishment in a xeric
forest-steppe ecotone in northern Patagonia, Argentina. We hypothesized
that apparent competition arising from incidental predation of newly

222

Abstracts

emerged seedlings by omnivorous beetles would decrease Austrocedrus

chilensis recruitment under "nurse" shrubs, a process otherwise facilitated
by shrub-canopy shading. Seedlings of A. chilensis were transplanted into
each of three microsites (beneath shrub, off-shrub sheltered and off-shrub
open), both within enclosures having natural or elevated beetle densities
and in unenclosed areas. Desiccation was the main source of seedling mortality (90%) in open microsites. Beetle addition increased seedling predation from 15 to 60% beneath shrubs, and from 3 to 20% in off-shrub
sheltered microsites. Pitfall trapping confirmed that beetles numerically aggregated in shrub patches, while litter removal/addition indicated that
shrubs provided both food and microhabitat to tenebrionids. Laboratory
trials showed that tree seedlings were most vulnerable to beetle attack within one month from emergence. Thus, early seedling mortality due to beetle
predation reduced the strength of, but did not completely eliminate, tree
facilitation by shrubs. Our results suggest that the balance between facilitation and apparent competition may influence the long-term structure of
this ecotonal plant community.
MCAULIFFE, JOSEPH R.,1,* ERIK P. HAMERLYNCK2 and MARTHA
C. EPPES.3 1 Desert Botanical Garden, Phoenix, AZ; 2 Rutgers University,
Newark, NJ; 3 University of North Carolina, Charlotte, NC. Repeated geological perturbations and long-lived creosotebush in the Mojave Desert.
Individual genetic clones of creosotebush (Larrea tridentata) have persisted
for thousands of years at a site in the Mojave Desert, California. Paradoxically, these clones are rare or absent on the older, more stable geomorphic
surfaces in the area, but instead are found where fluvial and eolian processes have repeatedly disturbed the soil surface throughout the Holocene.
These disturbances prevent the formation of a fine-textured soil horizon
(vesicular A or Av horizon) directly beneath the surface. When present,
this horizon strongly inhibits infiltration of precipitation and negatively
impacts plant water status. The geologically dynamic nature of the Mojave
Desert environment, rather than geological stability per se, has allowed the
persistence of clones through lengthy episodes of extreme climatic drought
during the last several thousand years.
MCCAIN, CHRISTY M. The University of Kansas, Lawrence, Kansas.
Species richness patterns of small mammals along an elevational gradient in Costa Rica.
Evaluations of elevational diversity theories show that elevational gradients
constitute one of the least understood patterns in biogeography. Recently,
null models, termed Mid-Domain models, have been proposed that suggest
that elevational patterns of species richness may be due solely to geographic constraints. These null models predict mid-domain peaks in species richness as a consequence of geometric patterns resulting from overlapping
species ranges between two geographic boundaries. Species richness and
abundances of small mammals were surveyed along an elevational gradient
at Monteverde Cloud Forest Reserve in Costa Rica during 2000-2002. The
four data sets constituted three temporal replicates: late wet, early wet, and
dry season, and one spatial replicate. In 16,900 trap-nights, 1561 individuals from 17 species were captured. Species richness peaked at mid-elevation between 1000-1300 m, with richness declining both at higher and
lower elevations. I developed a new program, Mid-Domain Null, to run
50,000 Monte Carlo simulations using empirical and randomized range
sizes and range midpoints to produce mid-domain prediction curves. Most
of the empirical curve of species richness occurs within 95% prediction
curves of the Mid-Domain model, although deviations from the model
exist. Other diversity theories: productivity hypotheses, habitat diversity,
resource diversity, competition, community overlap, and historical hypotheses are considered; only the mid-domain and climatic factors were supported moderately, although not one factor, but many factors appear to
underlie the pattern of species richness.
MCCALL, ANDREW C.* and ROBIN D. WAUGAMAN. University of
California-Davis, Davis, CA. Induced floral resistance in a native California annual, Nemophila menziesii.
Induced responses to foliar herbivory have been relatively well-studied, but
such responses in floral tissues have received considerably less attention.
If there is a cost to resistance traits or if resistance traits in floral tissue

negatively affect pollinator service, then plastic resistance responses in

flowers may be effective strategies for plant populations. Thus far, there
has been little evidence that induced floral responses to either prior leaf or
floral herbivory occur in natural populations. Here we demonstrate induced
floral resistance in a population of Nemophila menziesii (Hydrophyllaceae)
in California over two years. In the first experiment, flowers damaged with
larvae of Platyprepia virginalis (Lepidoptera: Arctiidae) produced subsequent flowers that were less palatable to P. virginalis than control plants
without damage. In a second experiment, plants with artificially-damaged
flowers produced both second and third flowers that received roughly 10
times less natural floral herbivory than flowers on control plants. There
were, however, no significant differences in total seed set, seeds per fruit,
or average weight of seeds between treatments. Induced resistance in flowers seems to occur in N. menziesii, although more experiments are needed
to demonstrate if this trait is adaptive in the face of natural floral herbivory.
MCCALLUM, HAMISH I.,1,* LEAH GERBER,2 KEVIN D. LAFFERTY3
and ANDREW P. DOBSON.4 1 University of Queensland, hmccallum@zen.uq.edu.au, Brisbane, Queensland, Australia; 2 Arizona State University, Tempe, Arizona; 3 USGS, University of California, Santa Barbara,
California; 4 Princeton University, Princeton, New Jersey. Exposing population viability analysis to disease.
Despite the importance of pathogens in natural populations, little attention
has been given to host-pathogen dynamics in population viability analysis.
The usual advice is to treat epidemics as "catastrophes", which occur with
a low probability and kill some random proportion of the current population. We present a modeling framework for testing hypotheses about the
role of disease in extinction risk analyses. We ask: (1) how disease affects
variability in abundance and thus, population viability and (2) whether
viability estimates of real populations suffering from disease require explicit modeling. As would be expected, we found that the presence of a
lethal disease decreased the median survival time of simulated populations.
Treating disease as a catastrophe in PVA leads to several erroneous conclusions. A key feature of epidemics is the existence of a host population
threshold for disease introduction. This threshold means that the effect of
epidemics is both density-dependent and overcompensatory. Modeling epidemics as increased stochastic mortality (catastrophe) grossly overestimated the probability of extinction, because of the density dependent effects of the threshold. Another effect of disease was to change the predicted
relationship between population growth rate and median time to extinction.
Populations with high growth rates that would normally be in little danger
of extinction were more likely to exceed the host-density threshold and
suffer virulent epidemics. These results suggest that, when confronted with
disease, populations with high rates of increase could be more susceptible
to extinction than populations with lower rates of increase. If potential
management actions involve manipulating pathogens, then it will be important to model disease explicitly.
MCCAMENT, CORINNE L.* and BRIAN C. MCCARTHY. Ohio University, Athens, Ohio. Survival and growth of American chestnut (Castanea dentata) seedlings under various silvicultural regimes in a mixed
oak forest ecosystem.
A blight resistant variety of American chestnut Castanea dentata is scheduled to be available for large-scale restoration efforts within the next decade. However, the regeneration ecology of this species is poorly understood relative to other hardwood species. The purpose of our study was to
evaluate the forest microenvironments best suited to C. dentata seed germination, seedling survival, biomass resource allocation, and leaf area to
target the best environments for future plantings of blight resistant seeds.
Using blight susceptible chestnut seeds, we planted 100 (n 5 10 reps of
10) in each of four silvicultural units (control, burn, thin, and thin/burn) in
three mixed oak forests (total of 1200 seeds) in southeastern Ohio. Germination and survival percentages were observed monthly (June-September, 2002). Seedlings were harvested in September and patterns of biomass
allocation determined. Germination or survival did not vary (P . 0.05)
among forests or treatments. However, patterns of biomass allocation varied
considerably among silvicultural treatments. Seedlings in any stand that
received thinning (increased light) were larger in basal diameter, taproot
length, taproot mass, fine root mass, stem mass, stem length, and leaf area

(all P , 0.01). Our data suggest that American chestnut is very responsive
to patterns of available light relative to growth and biomass allocation.
Thus, future restoration efforts should target gaps or recently thinned stands
for chestnut reintroduction.
MCCARRON, JAMES K.,1,* KAREN A. GARRETT,1 PHIL A. FAY,1
SHAUNA P. DENDY,1 HELEN M. ALEXANDER2 and B M. BROECKELMAN.1 1 Kansas State University, Manhattan, KS; 2 University of Kansas, Lawrence, KS. The effect of altered rainfall patterns on leaf rust
severity in Solidago canadensis.
Altered precipitation patterns predicted by climate change models may affect the incidence and severity of plant disease in natural systems. We
studied the effects of precipitation patterns on incidence of a leaf rust
caused by Puccinia dioicae on goldenrod, Solidago canadensis, a common
native forb in tallgrass prairie. Growing season rainfall patterns were manipulated at Konza Prairie Biological Station by reducing rain event volume
by 30%, increasing the length of the dry interval between rain events by
50%, or both, using rainout shelters fitted with sprinkler systems. An unsheltered plot and a shelter irrigated at ambient quantities and intervals
were the controls. We rated goldenrod plants for incidence of leaf rust for
four growing seasons, and used Poisson regression in a log linear model
to examine how precipitation treatments affected incidence. Goldenrod in
plots receiving reduced rainfall quantity, rainfall at increased intervals, or
both, had a significantly lower incidence of leaf rust in the wetter year. In
other years, proximity of a primary host species was a more important
factor. Both quantity and pattern of rainfall inputs may affect disease incidence in native grasslands. In addition, we are modeling the effects of
climate change, in terms of predicted changes in leaf wetness duration, on
the severity of leaf rust and similar diseases.
MCCARTHY, BRIAN C.,1,* CAROLYN H. KEIFFER2 and STEVEN H.
ROGSTAD.3 1 Ohio University, (mccarthy@ohio.edu), Athens, OH, USA;
2
Miami University - Middletown, Middletown, OH, USA; 3 University of
Cincinnati, Cincinnati, OH, USA. Composition, structure, and diversity
of a chestnut-dominated hardwood forest in southwestern Wisconsin,
USA.
American chestnut (Castanea dentata (Marshall) Borkh.; Fagaceae) was
virtually extirpated as a dominant tree species throughout the Appalachian
forest region by the 1930s due to the chestnut blight (Cryphonectria parasitica; Ascomycota). Fortunately, a small number of disjunct chestnut populations escaped infection throughout the Midwest. One such stand, believed to be the largest, exists near La Crosse, Wisconsin. A settler reportedly planted 12 chestnut trees in the early 1900s, from which 11 exist
today (the largest being 127 cm in diameter). Since this initial introduction,
American chestnut has naturalizedmore than 6,000 individuals now dominate this mixed oak forest. The purpose of our project was to establish a
long-term study of forest dynamics, conduct a detailed study of the current
composition and structure of the vegetation, and assess the regeneration
ecology of chestnut, which was largely extirpated in Appalachian forests
before it could be studied. In the summer of 2001, we established 30 permanent plots, 500 m2 throughout the stand. Mature trees, saplings, seedlings, shrubs, herbs and groundcover were sampled in each plot. In addition, hemispherical canopy photos and soil samples were taken to describe
the microenvironment and regeneration conditions. A total of 27 species
were found in the overstory. The total basal area of the stand was 35 m2ha1
, of which chestnut accounted for 43%. Chestnut has acted almost as an
invasive species in colonizing this oak-hickory forest. This occurrence provides considerable hope for restoration ecologists considering the re-introduction of chestnut into oak forest ecosystems, as disease resistant varieties
become available.
MCCARTHY, HEATHER,* BONNIE MILLAR and RAM OREN. Duke
University, Durham, NC, USA. Drought sensitivity of growth in a pine
forest under CO2 enrichment and differing nutrient supplies.
Basal area increment and shoot and needle elongation were monitored in
a free air CO2 enrichment experiment in a 20-year-old loblolly pine forest,
during a growing season with severe drought (;50% lower than average
April through September precipitation). Combining these measurements

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223

with previous measurements during non-drought years allowed for a comparison of drought sensitivity across woody (stems and shoots) and nonwoody (needles) components, exposed to differing levels of atmospheric
CO2 and soil fertility. Although the stem growth period during the dry
growing season lasted longer than in growing seasons with average precipitation, annual basal area increment was significantly reduced by
drought. Under drought conditions, trees at ambient CO2 with native soil
fertility had 56% lower stem growth and 51% lower shoot growth (both
P,0.001) than under non-drought conditions. At the same time, needle
growth was reduced by 22% (P50.002) in the upper canopy. Under elevated CO2 with native soil fertility there was a lesser decrease of 47% in
stem growth. However, in contrast to ambient CO2, drought caused no
reduction in shoot or needle length. This indicates that elevated CO2 may
provide a buffer for drought-induced growth reductions, but that this buffer
is greater within the canopy. With fertilization, drought-induced reductions
in stem growth under ambient and elevated CO2 were 55% (P,0.001) and
46%, respectively. Thus, stem growth is similarly buffered against drought
by elevated CO2 whether nutrients are added or not.
MCCARTHY, MEGAN C.,1,* BRIAN J. ENQUIST1 and KARL J. NIKLAS.2 1 University of Arizona, Department of Ecology and Evolutionary
Biology, Tucson, AZ, US; 2 Deparment of Plant Biology, Cornell University, Ithaca, NY, US. Global allocation rules for plant biomass partitioning: Evolutionary and functional trends across land plants.

ed after radicle emergence under low light (3% full sun) in a shadehouse.
Hampea appendiculata and Prestoea decurrens showed the strongest mortality response with 0% and 83% survivorship in the pathogen treatment
vs. 67% and 100% in the control. Vochysia ferruginea and Welfia regia
showed modest decreases in survivorship (10%) as well as decreases in
biomass (20%) in the pathogen treatment. Tapiria myriantha exhibited no
difference in survivorship but a 12% decrease in final height. Individuals
of Vitex cooperi although exhibiting a mortality response (60% survivorship in treatment vs. 73% in control) had a positive growth response (20%)
to pathogens. Neea psychotrioides was the only species that showed an
increase (10%) in survivorship whereas Apeiba membranacea, Virola koschnyi, Pentaclethra macroloba, Iriartea deltoidea and Socratea exorrhiza
showed no response to the pathogen treatment. Contrary to expectations
derived from the Janzen-Connell Hypothesis, there was no relationship between disease sensitivity and adult abundance. In addition, disease sensitivity was not related to independent shade tolerance rankings. Overall,
proximity to adult conspecifics showed weak effects and patterns contrary
to expectations, suggesting possible independent effects of conspecific
seedling density.
MCCAULEY, LISA1,* and DAVID G. JENKINS.2 1 University of Illinois
at Urbana-Champaign / IL DNR, Springfield, IL; 2 University of Illinois at
Springfield, Springfield, IL. GIS-based estimates of former isolated depressional wetlands in an agricultural landscape.

Understanding how plant biomass is partitioned between roots, stems,

leaves, and reproductive tissues is central for many questions in life history
evolution, ecology, and ecosystem studies. Recently, a biomass partitioning
model based on the constraints on resource transport and photosynthetic
harvesting capacity was developed to explain how plants should divide
their energy between leaf, stem and root function. The biomass partitioning
model predicts that leaf mass, and total, stem, root and leaf production
should scale to the 3/4th power of body size and stem and root mass should
scale isometrically to body size. However, while empirical data fit these
predictions, there was additional residual variation not explained by the
model. Here we assess the relative importance of evolutionary history,
functional trade-offs, and large scale environmental variation on explaining
additional variation not explained by the predicted allometric constraints.
Phylogeny and taxonomy did not influence the predicted biomass partitioning for any of the biomass partitioning relationships examined, suggesting that the constraints acting on the organ mass and production scaling
relationships are similar across evolutionary groups. However, taxa differed
significantly in the relative proportions of allocation to stems, roots, leaves,
and fruits. Angiosperms and gymnosperms only differed in the amount of
leaf mass and the amount of production for a given leaf mass. Nested
Analysis of Variance across taxonomic hierarchies indicated that most variation in vegetative partitioning exists at the species and genus levels indicating a strong environmental signal. The differences in leaf mass and
production for a given leaf mass appear to be due to differences in leaf
phenology between angiosperms and gymnosperms. However, variation in
allocation to reproduction is deeply rooted within the land plant phylogeny.
Thus, it is shown how a general allometric model provides a framework
for quantifying the relative influences of environment and functional differences on influencing plant biomass partitioning.

Before European settlement, 23% of Illinois (8 million acres) was covered

by wetlands. Currently, only 3.5% of original wetlands (1.25 million acres)
exist in Illinois, this is a loss of an estimated 90% of the states wetlands.
Champaign County was one of the most extensively drained counties in
the State: 40-61% of its pre-settlement area was covered in wetlands: today
only 0.9% of its area is covered by wetlands (i.e., 39-60% of original
county area was drained), and 98% of the wetland number. We used GIS
to map the spatial extent, density, pattern, and sizes of drained and extant
depressional wetlands in Champaign County, Illinois. We derived several
models of former wetlands; models used digital soil maps to identify hydric
soils, but varied by using 10 m and 30 m Digital Elevation Models (DEM),
Digital Raster Graphics (DRG), or Digital Orthophoto Quadrangles (DOQ).
We also conducted visual field surveys for saturated or ponded conditions
to test the models. Our most reliable model to date is based on DRGs and
hydric soils, and indicates that circa 900 depressional wetlands (75% of
number, 69% of area) were drained, accounting for 868 ha of lost wetland
habitat in Champaign County alone. Thus, depressional wetlands accounted
for the majority of historical wetland loss. Nearest-neighbor distances have
dramatically changed, so that an organism adapted to disperse in the former
landscape could reach 72% of all nearest neighbors within 500 m. Today,
that probability never exceeds 23%, even at 3000 m distance. The modern,
drained landscape of Champaign County poses potential risk for remaining
wetland populations, and GIS models of precise former wetlands locations
can be valuable for wetland conservation and restoration efforts.

MCCARTHY, SARAH E.* and RICHARD K. KOBE. Michigan State University, East Lansing, MI. Survivorship and growth of tropical tree seedlings to fungal pathogens extracted from soil cultured under conspecific
adults.

Phenotypic plasticity can change community dynamics by altering the nature of species interactions. Therefore, it is important to quantify phenotypic change and incorporate its effect into studies of community ecology.
To compare magnitudes of plasticity across taxa and to understand the
adaptive significance of these responses, we require unbiased estimates of
changes in morphology induced by environmental factors. Because morphology changes with overall size, analytic procedures must control for
allometric scaling of specific traits with body size. Many studies of phenotypic plasticity use a technique called shearing to correct for size. Shearing, like all methods of size correction, is based on the assumption that
groups being compared share a common allometric pattern. In shearing,
this common allometry is usually characterized by the first principal component (PC1) of the pooled data (across all treatment groups). Residuals
from ordinary least squares regression of each trait against PC1 are then
compared among groups. Shearing assumes that (1) PC1 obtained from
pooled data correctly describes the within-group scaling relationships; and

Soil fungal pathogens are a likely but understudied mechanism of distanceand density- dependent mortality patterns that could contribute to the maintenance of tree species diversity. Using seedlings of 11 tree species that
varied in abundance and shade tolerance in a wet tropical forest (La Selva
Biological Station, Costa Rica), we compared species-specific survivorship
and growth responses in soils with and without inoculations of pathogenic
spores extracted from soils that had been cultured by conspecific adults.
Pathogen treatments and controls were inoculated with arbuscular mycorrhizal fungi (AMF). To minimize potential artifacts associated with soil
sterilization, we used a wet sieving method to extract pathogenic and AMF
spore fractions, which were added to sterile medium. Seedlings were plant-

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Abstracts

MCCOY, MICHAEL W.,* CRAIG OSENBERG, BENJAMIN M. BOLKER, BENJAMIN MINER and JAMES VONESH. University of Florida,
Department of Zoology, PO Box 118525, Gainesville, Florida, USA.
Quantifying phenotypic plasticity in ecological experiments.

(2) the allometric relationships (covariance matrices) of the groups are identical. Morphological data may violate both of these assumptions; in particular, inducing phenotypic changes in experimental organisms often changes
allometric patterns. Nonetheless, most studies of phenotypic plasticity in
amphibians and other vertebrates have applied shearing to compare phenotypes without acknowledging or testing these important assumptions. In
this study, we discuss potential biases imposed by violations of these assumptions and discuss Common Principal Components (CPC) and Burnaby9s back-projection method as an alternative for the study of phenotypic
plasticity. We use simulated data and results from an experiment testing
for predator-induced plasticity in pinewoods treefrog (Hyla femoralis) to
compare the two approaches.
MCCULLEY, REBECCA L.,1,* ESTEBAN JOBBAGY,2 WILL POCKMAN3 and ROBERT B. JACKSON.1 1 Duke University, Durham, NC,
USA; 2 Universidad de Buenos Aires, Buenos Aires, Argentina, Argentina;
3
University of New Mexico, Albuquerque, NM. Understanding patterns
of root uptake using strontium isotopes, root and soil nutrient distributions, and modeling.
Broad geographic trends in root biomass distributions across climatic gradients and for plant functional types do not indicate patterns of root activity
or function. In an attempt to assess root uptake patterns, we sampled native
semi-arid and sub-humid grasslands (to a depth of 7 m) for root biomass,
soil nutrient concentrations (P, C, N, K, Ca, Mg, Cl and Na), and strontium
isotopic signatures (from both plant material and soil). The strontium isotopic data indicate that the integrated depth of cation uptake by the grassland species decreases with increasing mean annual precipitation from an
average integrated depth of 2.8 m at the most arid location to 0.1 m at the
most humid. Grassland root biomass was highly concentrated in the top 1
m of soil (accounting for 94.6 - 99.7% of total root biomass across sites),
but roots were encountered 8 m deep at the driest site. Soil chloride profiles,
environmental indicators of long-term water movement, indicate that majority of the soil water entering these ecosystems as precipitation does not
percolate below 2 m of depth. Biologically-cycled nutrients were highly
concentrated in the top 0.25 m of soil, with a concentration factor ranking
of C . N & P . K . Ca & Mg. Consistent with previous work, our data
suggest that through biological cycling limiting nutrients are removed from
the soil at depth and concentrated at the soil surface. Our data reveal that
nutrient uptake occurs surprisingly deeply in aridland soils (. 2 m), perhaps as a result of deep roots for water uptake.
MCCULLOH, KATHERINE A.* and JOHN S. SPERRY. Department of
Biology, University of Utah, Salt Lake City, Utah, USA. The application
of Murrays law to Psilotum nudum, an analogue of an ancestral vascular plant.
Murrays law (ML) predicts that the optimal design for a transport system
that maximizes hydraulic conductance for a given investment in vascular
tissue occurs when the sum of the conduit radii cubed (Sr3) is constant
across ranks. This law has been shown to apply to the xylem conduits of
plants when the conduits are only functioning in a water transport role and
not providing the plant with structural support. Until now, work on ML in
plants has concerned relatively derived angiosperms and gymnosperms.
Determining when plants evolved this optimal system would provide valuable insight to plant vascular systems. The fern Psilotum nudum closely
resembles plants known only from the fossil record such as Cooksonia and
Rhynia. In these fossils and the extant P. nudum, the xylem is centrally
located in the stem, suggesting it does not significantly contribute to the
structural support of the plant and that ML should apply. However, ML
assumes that the volume flow rate (Q) in the transport system remains
constant, which is not the case in the stem-photosynthesizer, P. nudum. The
derivation of ML was extended to the case of variable Q, and it was shown
that in the optimal system Sr3 remains proportional to the decrease in Q.
By measuring the transpiration along the stem with a porometer, to estimate
Q, and measuring the Sr3, we were able to determine if the Sr3 did remain
proportional to Q. Data show a strong correlation between Sr3 and Q,
suggesting that ML does apply to these simple plants and the optimal
design may have developed early in vascular evolution. Futhermore, P.
nudum deviated from the pipe model, exhibiting increasing numbers of
conduits from the base to the tip of the shoot. This deviation results in a
higher ML optimum over the less efficient pipe model.

MCDANIEL, JOSH M.,* KELLY ALLEY, ELIZABETH TURES and

PHILLIP CHANEY. Auburn University, Auburn, AL. Linking environmental knowledge, land use, and policy: A survey of rural, urban, and
suburban watersheds in west Georgia.
Knowledge of the environment can be affected by a variety of variables
including education, cultural traditions, geographic residence, occupation,
economic pursuits, recreational activities, differential access to information,
and unequal power relations. In this study, we explore variation in environmental knowledge by groups that are discrete in relation to demographic
and socio-economic criteria. We also examine the connections between
knowledge and human impacts on the environment. The changes in land
use in an area of economic development and urbanization provide the setting against which we sort out these dynamics. Our objective is to identify
the variables that increase or lower knowledge of environmental impacts
of urbanization, and then relate our findings to actual household land use
practices and participation in public policy debates. In this poster we present results regarding a survey done in rural, suburban, and urban watersheds in three rapidly urbanizing counties in West Georgia. We test hypotheses regarding the factors that influence different levels of environmental knowledge, land use practices, and perceptions regarding environmental policy. We also examine how variations in environmental
knowledge influence land use practices that may impact water quality, biodiversity, and forest ecosystems at the household level. Explanatory variables for individual households include presence of different waste disposal
systems and practices, recreational activities, pesticide/herbicide use on
lawns, gardens, and timber lands, construction of streamside management
zones (SMZs), and use of best management practices (BMPs) on timberlands. Finally, we assess how varying levels of knowledge affect individual
participation in public policy platforms concerning regulatory mechanisms
designed to promote environmental conservation and/or economic development. Variables of policy participation include membership in civic organizations and other non-governmenal organizations concerned with development/conservation issues, voting records, and attendance at public
meetings concerning land use regulation and economic development.
MCDONALD, MICHAEL E.1,* and BARBARA LEVINSON.2 1 Environmental Monitoring and Assessment Program, Office of Research and Development, US EPA, RTP, NC, USA; 2 STAR Grants Program, Office of
Research and Development, US EPA, Washington, DC, USA. Why do we
need environmental indicators.
Coastal ecosystems are complex and dynamic. Establishing the condition
of all the organisms and their relevant abiotic environment in any given
estuary is not feasible. Environmental indicators are used as surrogates in
these systems with the tacit assumption that the condition of the indicator
represents the condition of the ecosystem. Benchmarks for these indicators
to be measured against are also required in order to determine if there is
an evironmental problem, the magnitude of the problem, where it is occurring, and what is causing it. Thus, these indicators must be generally
applicable over a relatively wide regions of the country. Eventually, these
same indicators may be required to establish the efficacy of management
programs and policies being used to reduce the environmental problems.
Developing environmental indicators to meet these needs has been a major
research emphasis for EPA, intramurally through its Environmental Monitoring and Assessment Program and extramurally with the academic community through its STAR grants program. As new environmental problems
emerge, the next generation of environmental indicators will be required
to establish coastal ecosystem condition. The results of these combined
research efforts will provide better scientific assessment of the state of our
coastal ecosystems, and better informed national, regional and local decisions on priority coastal issues.
MCDONALD, ROBERT I.* and DEAN L. URBAN. Duke University,
Durham, NC. The effect of forest edges on tree growth rates.
Forest fragmentation is a common process occurring in forests worldwide
that has been shown to have important implications for tree species composition and abundance. In particular, the effects of forest edges on microclimate are often profound, usually resulting in increased light available to
plants along the forest edge. Using dendrochronological techniques, we

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225

assessed the effect of forest edges on the growth rate of 4 species that are
common in the North Carolina Piedmont: Acer rubrum, Liriodendron tulipifera, Liquidambar stryacifula, and Pinus taeda. Transects from the forest edge into the forest interior were established on more than 60 forest
edges in the Duke Forest (Durham, NC), incorporating edges of varying
aspect and ages. Within a transect, all stems greater than 10cm were cored,
and their spatial position and species recorded. For each transect, a set of
environmental and edaphic variables were measured, including soil texture
and nutrient values, and percent canopy closure. All transects were located
in a GIS system, and land cover data classified from a time-series of Thematic Mapper images were used to assess the age of each edge. All four
species had significant increases in growth rates within 5 meters of an edge
(P , 0.05 in all cases), but the effect did not appear to deeply penetrate
into the forest interior. However, the increase in growth rate near edges
explains a substantially smaller portion of the variance than edaphic or
other types of explanatory variables. There is an indication of possible
interactions between soil texture and the effect of forest edges on tree
growth rate. Overall, this increased growth rate may prove to be quite an
important effect for some applications, since analyses of land-cover maps
for the region suggest a substantial percentage of trees are within 10 meters
of an edge.
MCELRONE, ANDREW J.,1,* WILL T. POCKMAN,2 ROBERT N. ADDINGTON1 and ROBERT B. JACKSON.1 1 Department of Biology, Duke
University, Durham, NC, USA; 2 Department of Biology, University of
New Mexico, Albuquerque, NM, USA. 20,000 mm under the surface:
Structure and function of deep tree roots in central Texas.
Deep roots often contribute substantially to whole tree water use through
uptake of reliable deep water resources, particularly in water-limited environments. Despite their importance, most studies of deep roots have favored indirect methods over direct physiological measurements because of
limited access. We used caves in the Edwards Plateau region of central
Texas to access deep tree roots utilizing a perennial underground stream at
approximately 20 meters depth. Daily and seasonal measurements of sap
flow, water potential, and transpiration on Bumelia lanuginosa and Quercus
fusiformis were combined with xylem anatomy and hydraulic conductivity
(Kh) measurements to characterize the response of these species to changes
in environmental conditions along the soil-plant-atmosphere continuum.
Sap flow and water potential data in deep roots closely tracked similar
changes in stem sap flow for both tree species. Changes in sap flow and
water potential in all plant components were most strongly correlated with
the changes in vapor pressure deficit and photosynthetically active radiation. Xylem pressure changes in stems propagated to deep roots in as little
as 25-45 minutes, suggesting low water storage for the trees in this system.
During whole-tree shade treatments repeated throughout the season on B.
lanuginosa, stem sap flow was reduced 50-75% as a result of decreased
transpiration. The concurrent reduction in deep root sap flow during these
periods varied in magnitude depending on the availability of shallow soil
water from recent precipitation events. Changes in xylem structure and
function were detected for both species with mean vessel diameter and Kh
largest in deep roots, intermediate in shallow roots, and smallest in branches. Such adjustments would reduce flow resistance in deep roots and maintain deep water use as environmental conditions change throughout the
season. These data show that deep roots are tightly integrated with aboveground plant components, which may contribute to the persistence of
woody species in water-limited habitats.
MCEUEN, AMY B.1,2 and LISA M. CURRAN.3 1 University of Michigan,
Ann Arbor, MI; 2 Present address: Wilkes University, mceuen@wilkes.edu,
Wilkes-Barre, PA; 3 Yale University, New Haven, CT. Seed dispersal and
recruitment limitation at large spatial scales in northern hardwood
forest fragments.
Despite increasing evidence of seed limitation in forest ecosystems, data
on spatial patterns of seed availability at large (.1ha) spatial scales remain
sparse. We monitored seed rain in five northern hardwood fragments in
Michigan for two years (40m spacing, n 5 114). Of the 37 woody plants
sampled (n 5 98,032 mature seeds), only three had widespread seed availability within all sites containing resident sources (seed in .70% of traps
in each fragment). Seed colonization, measured as the percentage of traps

226

Abstracts

within a fragment receiving seed, differed based on dispersal vector, with

animal-dispersed species hitting fewer traps than wind-dispersed, even after
adjusting for lower seed production. High colonization percentages (.70%)
were found for light-seeded, high-fecundity species (Betula papyrifera, Betula alleghaniensis) at low source densities and distributions (,0.15m2 basal
area*ha-1 and ,4% plots containing mature individuals), but only occurred
for heavy-seeded, lower-fecundity species (Acer rubrum, Fraxinus pennsylvanica, Tilia americana) when source densities and distributions were
high (.0.7m2 basal area*ha-1 and .19% plots containing mature individuals). Heavy-seeded wind-dispersed species and animal-dispersed species
also showed evidence of recruitment limitation, with seedling presence associated with seed presence for Carpinus caroliniana, Fagus grandifolia,
Prunus avium, and Tilia americana. No seed dispersal into fragments was
detected for Acer rubrum, Betula alleghaniensis, or Tsuga canadensis, despite high seed colonization levels (.50%) in fragments with resident
sources, suggesting these species can be seed-limited at larger spatial
scales. Our results confirm the importance of seed dispersal to forest dynamics and suggest seed availability plays a significant role in determining
variation in plant community composition across scale.
MCEWAN, RYAN W.,1,* ROBERT N. MULLER2 and BRIAN C. MCCARTHY.1 1 Department of Environmental and Plant Biology, Ohio University, Athens, OH, USA; 2 Santa Barbara Botanical Garden, Santa Barbara, CA, USA. Vegetation-environment relationships among four canopy-layers in an old-growth mixed mesophytic forest.
Previous work in an old-growth mixed mesophytic forest has indicated that
Acer rubrum and Acer saccharum are dominant in the understory strata,
while xerophytic oak species (e.g. Quercus montana) are becoming increasingly restricted to the overstory canopy. This apparent shift in dominance
is similar to that seen in xeric oak forests throughout the eastern U.S. and
yet the species composition and environmental conditions in those oak
forests represent only one extreme of the gradient present in mixed mesophytic forest systems. We examined the oak-to-maple transition phenomenon within our mixed mesophytic study site in order to examine vegetation-environment relationships among woody species in four canopy-strata.
We hypothesized that 1) the most important environmental variables determining vegetation structure would differ among canopy-layers and 2)
relative abundance of Acer and Quercus would be maximized under very
different environmental conditions. Overstory vegetation was arrayed in
Canonical Correspondence Analysis space along a gradient of soil fertility
(Ca, r 5 -0.72) and elevation (r5 0.76). In the midstory, soil fertility (Ca,
r 5 -0.78), elevation (r 5 0.66) and light availability (r5 0.57) were important factors. Vegetation patterns of the shrub-layer and the ground-layer
were driven by soil fertility (Ca, r 5 -0.84, r 5 -0.78, respectively) and
soil pH (r 5 0.63, r 5 0.75, respectively). Across strata, Quercus sp. were
consistently located in ordination space in upper elevation areas with high
light availability and low soil pH. Acer rubrum was located near the center
of each diagram, suggesting plasticity, and Acer saccharumappeared to be
restricted to more nutrient rich areas. Though a dominance shift seems to
be underway, our data indicate that the environmental criteria necessary
for maximum relative dominance of Acer and Quercusare similar between
strata.
MCFARLIN, CAROLINE R.,1,* TRACY L. BUCK,2 STEVEN Y. NEWELL2 and STEVEN C. PENNINGS.3 1 The University of Georgia, Athens,
GA; 2 The University of Georgia Marine Institute, Sapelo Island, GA; 3 The
University of Houston, Houston, TX. Impact of fertilization on the abundance of two detritivorous snails.
Because of anthropogenic alterations to the global nitrogen cycle, there is
increasing interest in how nutrient supply affects community structure and
processes. Most fertilization studies have focused on changes in plant biomass or community composition, and only a few studies have examined
how nutrient effects propagate through the food web. Salt marshes are
particularly vulnerable to eutrophication because they receive nutrient inputs from adjacent watersheds. Here we examine the impact of fertilization
on two detritivorous snails that feed on litter from two abundant salt marsh
plants, Spartina alterniflora and Juncus roemerianus, at 19 sites within the
Georgia Coastal Ecosystems LTER study area. In control and fertilized
plots, densities of both snail species were positively correlated with nitro-

gen content of live Juncus, but were not correlated with nitrogen content
of live Spartina, nitrogen content of litter from either plant species, or plant
biomass (live or litter). The two snail species were negatively correlated
with each other in each treatment, supporting previous studies suggesting
that they compete. Fertilization significantly increased nitrogen content of
live plants of both species, but did not affect the nitrogen content of either
litter type, perhaps because plants resorbed nutrients from senescing leaves
or because litter was in variable states of decay. Fertilization increased the
biomass of Spartina litter and decreased the biomass of Juncus litter, but
did not alter densities of either snail species, nor did it alter the negative
correlation between species. Sites differed strongly in snail numbers, irrespective of experimental treatments. Our results suggest that impacts of
eutrophication may not propagate through the food web to affect detritivore
densities, possibly because eutrophication may not strongly affect litter
quality, or because other factors such as competition and predation may
more strongly mediate detritivore densities.
MCGHEE, JAY D.1,* and JIM BERKSON.2 1 Virginia Polytechnic Institute
and State University, Blacksburg, VA; 2 Virginia Polytechnic Institute and
State University, Blacksburg, VA. A meta-analysis of wild turkey reproductive parameters.
Understanding the reproductive parameters of wild turkey populations at
both the local and landscape levels is important for their effective management. We attempted to assess reproductive parameters for the eastern
wild turkey (Meleagris gallopavo silvestris) at the landscape level. To accomplish this, we used a meta-analysis approach to combine information
from multiple radio-telemetry studies. We then examined estimates of nesting and renesting rates and success, hen success, clutch size, hatching success, and poult survival at 2 weeks. We conducted x2 tests to determine
homogeneity across studies and determine overall differences in reproductive vital rates between adult and subadult hens. We used weighted linear
regression on heterogenous parameters to assess potential geographic structure (latitude, longitude). Analysis indicated that adult hens exceeded subadult hens in nest rates (x2 5 73.953, df 51, p , 0.0001), renest rates (x2
5 10.831, df 51, p 5 0.0010), and hen success (x2 5 15.092, df 5 1, p
5 0.0001) only. Only clutch size qualified as homogenous across studies
(x2 5 2.83, df 5 14, p 5 0.9993). Regression analysis indicated that adult
nest rate (F2,10 5 50.84, p , 0.0001) and adult renest rate (F1,7 5 29.76,
p 5 0.0010) varied according to latitude. We conclude that reproductive
vital rates in wild turkey hens vary primarily according to extrinsic factors
with some underlying latitudinal structure. For the majority of reproductive
parameters, landscape level estimates cannot adequately be applied to local
populations.
MCGINNIS, THOMAS W.,1,* JON E. KEELEY,1 MATTHEW L.
BROOKS,2 ROBERT L. SANFORD3 and JAYNE BELNAP.4 1 U.S. Geological Survey-Biological Resources Discipline, Three Rivers, CA; 2 U.S.
Geological Survey-Biological Resources Discipline, Henderson, NV; 3 University of Denver, Denver, CO; 4 U.S. Geological Survey-Biological Resources Discipline, Moab, UT. Above and belowground fire-related temperatures in a cheatgrass (Bromus tectorum L.) infested Sierra Nevada
yellow pine forest: Effects on cheatgrass seeds and native plants.
The unforeseen influx of cheatgrass following prescription burning in the
old growth yellow pine forest of Kings Canyon National Park, California,
emphasizes the need for more research on fire ecology. In 420 5x5 m
treatment plots we have measured how fire may alter cheatgrass and native
plant populations. This paper focuses on the range of temperatures that
occur above and belowground in plots with different fuel loads and burning
seasons, and how these relate to plant survival and recruitment. Late fall
burns in 2001, with light fuel loads and high relative humidity, did not
cause noticeable mortality in established perennial natives or in cheatgrass
seeds. Soil temperatures at 5 cm belowground in the late fall burn reached
a mean peak temperature of only 188C, rising 58C due to fire (+58C). To
simulate a longer fire return interval, additional fine fuels (5 cm-deep pine
needles) were added in summer and fall 2002 burn-plots resulting in significantly higher above and belowground heating. At 2 cm belowground,
needle-addition plots reached a mean peak temperature of 2428C (+988C)
in the summer and 1558C (+1388C) in the fall, while untreated plots reached
758C (+408C) and 698C (+468C), respectively. Emulating conditions in the

field, we subjected cheatgrass seeds to several heat, cold and moisture

treatments, then incubated them to determine mortality. Some dry seeds
survived brief 1208C heating while wet seeds survived 1008C. From our
lab results we predict that most, but not all of the seeds in needle-addition
field plots should have been destroyed, while a significant number of seeds
would have survived in the unaltered plots. Preliminary field observations
in February 2003 agree with the predictions. We will discuss our final
results from the spring 2003 growing-season.
MCGRAW, JAMES B.* and MARY ANN FUREDI. West Virginia University, Morgantown, WV. Demography of central populations of American ginseng.
American ginseng is being harvested from natural populations throughout
the central part of its range. Previous demographic studies at the northern
range margin suggested that levels of harvest typically seen were unsustainable there. Over a two year period, we censused 27 natural populations
containing over 3000 individuals spread over four states in the central part
of the distribution. Using population projection matrix models, we determined individual, regional, and overall population growth rates. Overall, in
2001-2, populations were found to be expanding slightly (l5 1.0427). In
the two states with the largest harvest rates, contrasting population growth
rates were found: Kentucky populations were found to be expanding, on
average (l 5 1.0846), while West Virginia populations were declining
slightly (l 5 0.9781). The cause of these differences is not clear, however
Kentucky populations appeared to have higher rates of seed production
(albeit lower germination rates) than West Virginia populations. Another
difference is that West Virginia populations were browsed by deer at high
rates. Further demographic simulations showed that the fate of seeds is
particularly important for ginseng population growth rates, and this represents two very important management opportunities. First, the ripeness of
seeds at the time of harvest was shown to significantly affect subsequent
germination; harvest prior to full seed ripening can drive population growth
rates below 1 (declining populations). Thus, it is exceedingly important
that the onset of the harvest season occur after completion of seed ripening.
Presently, there is very little data to suggest current harvest seasons are
optimally timed. Second, the disposition of seeds by the harvester is critical
to population growth. Harvesters can dramatically influence population
growth and viability if they are encouraged to (a) harvest only seed-producing plants, (b) leave seeds in the population, and (c) plant the seeds 2
cm deep to enhance subsequent germination.
MCGRAW, JODI M. University of California, Berkeley, CA, USA.
Mounds enhance endangered plant persistence and native community
structure by reducing exotic plants and litter.
Mounds created by burrowing animals can enhance diversity by creating
spatial and temporal refugia that allow persistence of fugitive native plant
species. However, many exotic plants can also colonize mounds and compete with native plant populations, rendering it difficult to predict the effects of these disturbances in invaded plant communities. Understanding
the mechanisms by which mounds influence plant populations can enhance
our understanding of these disturbances as well as facilitate conservation
of rare, disturbance-dependent plant species. I examined the effects of gopher mounds on plant community structure and the persistence of two
federally endangered plant species (Chorizanthe pungens var. hartwegiana
and Erysimum teretifolium) in a unique edaphic community in central
coastal California known as the sandhills. Using a factorial experiment, I
examined the roles of exotic plant species and litter in determining the
effects of mounds created in two seasons (spring vs. fall) on the demographic performance of the two endangered species and plant community
composition. I found that mounds removed accumulated leaf litter on the
soil surface and reduced the cover of exotic plant species by over 75%. In
addition, there was a trend towards higher native plant cover and native
plant richness on mounds. Mounds also facilitated the performance of the
two endangered plants. They enhanced germination of Erysimum by removing accumulated leaf litter and increased seedling growth and survival
to reproduction by reducing exotic plant competition. Mounds had additional beneficial effects beyond simply removing leaf litter and reducing
exotic plant competition, however, as both Chorizanthe and Erysimum grew
larger on mounds than on undisturbed plots with litter and exotic plants

Abstracts

227

removed. This effect was most pronounced for mounds created in the fall,
perhaps due to increased soil nutrient availability. Overall, these natural
disturbances enhance native plant performance and community structure.
MCGREGOR, RACHELLE L.* and LENORE FAHRIG. Carleton University, Ottawa, ON, Canada. Do small mammals exhibit road aversion?
We hypothesize that small mammals exhibit road aversion and do not attempt to cross roads. We predicted animals that avoid roads should demonstrate two characteristics: they will not include roads in their home range
and will turn back from roads (i.e., make U turns) when traveling through
forested habitat. To test whether small mammals include roads within their
home ranges, we established a mark-recapture protocol using three trapping
grids: two adjacent to but on opposite sides of the road, and one farther
into the same forest as one of the other grids. We found no difference in
small mammal densities between grids adjacent to roads and those more
interior (p 5 0.56). However, we found animals were less likely to cross
roads than expected (p ,0.001), although suitable habitat across roads
could be included in their home range. These results suggest that animals
will use all habitat up to but excluding roads. To test for behavioural responses to roads, we translocated fluorescent-powdered small mammals
across roads and within continuous forest. We recorded paths created by
sloughed-off powder and found that animals translocated across roads had
straighter paths, but there was no significant difference in their return rates
(p 5 0.18). We conclude that small mammals may prefer to exclude roads
from their home ranges but will cross if necessary (i.e., to disperse).
MCGRODDY, MEGAN E.,* LARS O. HEDIN and TANGUY DAUFRESNE. Department of Ecology and Evolutionary Biology, Princeton University, Princeton, NJ, USA. Scaling of C:N:P ratios in forest ecosystems
world-wide.
The relative constancy of carbon:nitrogen:phosphorus (C:N:P) ratios has
been key to understanding the role of limitation and biotic-abiotic interactions in shaping nutrient cycles and ecosystem function in marine ecosystems for decades. In contrast, research on stoichiometry in terrestrial
ecosystems has primarily focused on variations at the individual and stand
level and fluxes and stocks within individual sites, although there has been
interesting work at larger scales looking at ratios as indices of nutrient
limitation. Using the marine ecologists approach we looked for Redfieldtype ratios in terrestrial forests on a global scale. While we were more
interested in the relative constancy of the C:N:P ratios than their numerical
value we did note that the atomic ratios calculated for foliage (1100:29:1)
and litter (3400:47:1) reflect the increased proportion of C-rich structural
material characteristic of terrestrial vegetation. We found that the global
forest C:N:P ratios in both foliage and litter were more variable than those
of marine particulate matter, biome level (temperate broadleaf, temperate
coniferous and tropical) ratios were as constrained as marine ratios and
statistically distinct from one another. These results led us to ask if the
biome scale patterns are generated by physiological constraints of foliar
tissue or are they the result of constraints imposed by community or ecosystem structure. Nitrogen:P ratios held constant across the range of biomass stocks and productivity in all forests suggesting that physiological
constraints are effective at all scales. Coniferous forests showed a shift to
lower C:N and C:P ratios at higher foliar biomass. Litter C:N and C:P
ratios in tropical forests and C:N ratios in temperate broadleaf forests also
decreased at higher productivity. We suggest that these diagnostic differences between biomes with respect to ratios and their scaling patterns may
lead to further understanding of strategies of forest nutrient cycles on global
scales.
MCGUIRE, MARY ANNE* and ROBERT O. TESKEY. University of
Georgia, Athens, Georgia, USA. Release of CO2 from xylem sap is responsible for apparent wound respiration in tree stems.
A rapid increase in CO2 efflux from plant tissues after they have been
disturbed or damaged has usually been considered a respiratory wound
response. However, the CO2 concentration ([CO2]) in tree stems is normally
very high (3-10%, or 30,000 to 100,000 mmol mol-1) relative to atmospheric concentration (370 mmol mol-1). The high CO2 gradient from xylem to
atmosphere is maintained by significant barriers to diffusion in the cam-

228

Abstracts

bium and bark layers. This suggests that any disturbance to the stem that
affects these barriers, for example, cutting through bark, will increase the
rate of efflux of CO2 from the xylem to the atmosphere. We examined the
role of [CO2] in the xylem of trees on apparent wound respiration by experimentally manipulating xylem CO2 concentration over a range of 1 to
17% [CO2] in saplings of two tree species, sycamore (Platanus occidentalis) and sweetgum (Liquidambar styraciflua) and measuring CO2 efflux
from the stems before and after wounding. In all cases (n518), pre-wound
and post-wound stem CO2 efflux was linearly related to stem [CO2] (R2 5
0.83 and 0.87, respectively). Wounding caused a rapid increase in CO2
efflux that was approximately four-fold greater than pre-wound rates of
efflux, and could be attributed almost entirely to the release of CO2 from
the xylem. These results indicate that the rapid increase in CO2 efflux from
tree stems observed after bark disturbance has been misinterpreted as a
wound response, when actually much of the effect can be explained simply
as CO2 degassing from the xylem. This finding has important implications
for studies of stem respiration if the bark was disturbed prior to measurement, for example, to facilitate attachment of a cuvette.
MCINTYRE, AIMEE P.,1,* RICHARD A. SCHMITZ1 and CHARLIE M.
CRISAFULLI.2 1 Oregon State University, Corvallis, OR; 2 Forest Service,
Pacific Northwest Research Station, Olympia, WA. Associations of the
Van Dykes salamander with abiotic riparian features.
The Van Dykes salamander (Plethodon vandykei, or PLVA) is a rare species endemic to Washington State. PLVA is commonly associated with
habitat features that exhibit specific hydrological, geomorphologic, and disturbance characters. It is often encountered in cool moist microhabitats
along streams, splash zones of waterfalls, and headwater seeps. However,
no study has investigated the specific habitat associations of PLVA. We
conducted salamander surveys, measuring biotic and abiotic habitat variables at stream sites in the Cascade Mountains. Our study design allowed
us to compare sites where PLVA was detected, to those where PLVA was
not detected, between stream sites. Using logistic regression, with presence
and absence as the response, we were able to develop models to predict
PLVA detection between sites. We developed a priori models and ranked
them with Akaikes Information Criterion (AIC). The top model for the
between stream analysis included abiotic riparian features: the proportion
of boulder on the valley wall, the number of additional stream channels
entering the main channel being surveyed, and the proportion of valley
wall with vertical morphology. Sites that exhibited an increase in any one
(or combination) of these features had an increased probability of PLVA
detection. These results suggested that the likelihood of PLVA detection at
a site is largely driven by abiotic factors. Future analyses will allow us to
explore how the abiotic variables suggested by the between stream model
relate to the probability of PLVA detection at a micro scale (within site
modeling) and a landscape scale (modeling using a Geographical Information System).
MCINTYRE, BLODWYN M.* University of Redlands, Department of Environmental Studies, 1200 East Colton Avenue, Redlands, CA, USA. The
common raven as a threat to esert tortoise, west Mojave Desert, California.
The common raven (Corvus corax,) is a subsidized predator of the desert
tortoise (Gopherus agassizii). Raven predation on juvenile tortoises is believed to be one of the most important threats to desert tortoise. Although
many threats to the desert tortoise are known, few, including raven predation, have been quantified, which is essential to conservation management. An area in the West Mojave Desert encompassing five Breeding Bird
Survey (BBS) routes was chosen for study. Raven population increases
since the 1970s were calculated for this area. Using GIS, features in this
landscape identified as "raven attractants" were quantified. Spatial and statistical analyses were used to estimate and rank the importance of various
"raven attractants" (landfills, water sources, urban areas, etc.) in their potential to increase raven populations locally. The results of this study can
be used in adaptive management of desert tortoise through management of
"raven attractants".

MCINTYRE, NANCY E.,1,* RICHARD A. NISBETT,1 ALISA ABUZEINEH,1 YONG-KYU CHU,2 CARL W. DICK,1 TYLA HOLSOMBACK,1
MARILYN A. HOUCK1 and COLLEEN JONSSON.2 1 Texas Tech University, Lubbock, TX, USA; 2 New Mexico State University, Las Cruces,
NM, USA. A hierarchical habitat selection analysis of claims that Hantavirus host rodents are asymptomatic.
Although host rodents infected with Hantavirus have been shown to be
indistinguishable from uninfected animals in terms of longevity, growth
rates, fitness, and health, natural selection (from a viral standpoint) should
favor differences in the behavioral ecology of infected and uninfected animals in such a way to facilitate viral transmission. However, the behavioral
consequences of infection with Hantavirus have not previously been examined. We therefore conducted a study in which habitat use by the marsh
rice rat (Oryzomys palustris, primary host for the Bayou strain of Hantavirus) was compared at multiple spatial scales between Hantavirus-positive
and uninfected animals, and these patterns were further compared to habitat
availability. Results to date show that there is significant habitat selection
by marsh rice rats in terms of both macro- and micro-habitats (i.e., habitat
types are not used simply in proportion to their availability). Although rates
of seroprevalence differed significantly between macrohabitat types, these
differences were a function of host density. Furthermore, microhabitat selection did not differ between seropositive and seronegative animals. Our
study thus provides corroboration for claims that, with regard to habitat
requirements, Hantavirus-infected rodents are apparently indistinguishable
from uninfected individuals.
MCINTYRE, PETER B.* and ALEXANDER S. FLECKER. Department
of Ecology and Evolutionary Biology, Corson Hall, Cornell University,
Ithaca, NY, USA. Trophic ecology in a Neotropical river: Insights from
stable isotopes.
We report the results of a stable isotope analysis (C, N) of the food web
in Rio Las Marias, a small Andean piedmont river in Venezuela. Basal
resources were represented by samples of epilithic algae, sediments, and
riparian vegetation, and consumer taxa included 55 fish species as well as
gastropods, crustaceans, amphibians, reptiles, insects, and birds. Carbon
isotopes suggest that allochthonous food sources are important in this food
web, contrary to expectations based on measurements of benthic productivity and results from other Neotropical rivers. Nitrogen isotopes indicate
that fishes occupy approximately three trophic levels, and most species held
intermediate trophic positions. There was no relationship between body size
and trophic position of fishes, in contrast to patterns observed among temperate fishes. Species traditionally categorized as benthic algivores differed
significantly in carbon and nitrogen isotope ratios, suggesting trophic segregation based on consumption of algae versus sediment. Spatial patterns
in carbon isotopes were also evident from comparisons of algae and consumers in forested tributary streams and the main channel. Carbon signatures in tributaries were relatively depleted, offering the potential to track
movement of organisms between the main channel and tributaries. These
results offer insight into trophic ecology in Rio Las Marias, and suggest
interesting contrasts with patterns in other aquatic ecosystems.
MCKANE, ROBERT,1,* PAUL RYGIEWICZ,1 PETER BEEDLOW,1
CHRISTIAN ANDERSEN,1 RENEE BROOKS,1 WILLIAM HOGSETT,1
MEAGAN HYNES2 and JOHN LAURENCE.3 1 U.S. Environmental Protection Agency, Western Ecology Division, Corvallis, OR, USA; 2 Dept. of
Land, Air and Water Resources, University of California, Davis, CA, USA;
3
U.S. Forest Service, PNW Research Station, Corvallis, OR, USA. Lateral root distribution of trees in an old-growth Douglas-fir forest inferred from uptake of tracer 15N.
Belowground competition for nutrients and water is considered a key factor
affecting spatial organization and productivity of individual stems within
forest stands, yet there are few data describing the lateral extent and overlap
of competing root systems. We quantified the functional root distribution
of different tree size classes in an old-growth forest in the Oregon Cascades
by measuring aboveground uptake of 15N (99 atom % as NH4Cl) injected
at different treatment radii around target Douglas-fir trees. Tree size classes
included sapling, intermediate and dominant trees. Each treatment included
50 evenly-spaced soil injections of 15N at 0.5, 1 or 1.5 times the mean

crown radius of each tree-size class. Preliminary analyses of the 15N content of aboveground tissues 4 months after injection show a monotonic
decrease in root function with distance, with uptake beyond 1 crown radius
accounting for over 1/4 of total uptake. These results suggest significant
extension of root systems beyond the dripline of trees and direct belowground competition among neighboring stems. We illustrate applying the
tracer data in combination with spatial data on stem location and size to
calculate the functional belowground overlap among neighbors. We also
examine how the intensity of belowground overlap is distributed throughout this stand and how it is correlated with growth rates of individual stems.
Our approach establishes a means for improving the representation of belowground competition in stand models of forest growth, a feature that
existing models either lack or treat theoretically.
MCKELVEY, STEPHEN R., PABLO PARYSOW* and ROBERT L. MATHIASEN. Northern Arizona University, Flagstaff, AZ. Modeling white
fir site productivity in southwestern USA: Parameter prediction and
guide curve methods.
The objective of this study was to model site productivity for Rocky Mountain white fir in Arizona and New Mexico. For that purpose, we developed
site index models and curves using both the parameter prediction and guide
curve methods. Site index is commonly used in growth and yield models
to project forest stands under alternative management scenarios. We collected stem analysis data from 91 white fir trees to carry out the parameter
prediction method. We also used the height and age of 1,054 white fir trees
to apply the guide curve method. We compared the white fir site index
curves produced by the parameter prediction and guide curve methods. In
addition, we compared the site index curves for white fir and Douglas fir
growing in Arizona and New Mexico, using the guide curve method for
both species.
MCKENNA, KERRY C.,1,* DARYL L. MOORHEAD,1 JOHANNA LAYBOURN-PARRY2 and EMILY ROBERTS.2 1 University of Toledo, Toledo,
OH; 2 University of Nottingham, Nottingham, United Kingdom, United
Kingdom. Structural changes in the pelagic food web of Lake Fryxell,
Antarctica.
Total biomass of the pelagic food web of Lake Fryxell, Antarctica showed
a significant increase from 1996 to 1998 (105 to 436 ug C/L, respectively).
Mass also varied with depth, ranging from 161 ug C/L under surface ice,
to a maximum 475 ug C/L at 9m, which corresponds to the deep chlorophyll maximum (DCM), then dropping to 275 ug C/L at 13m. Significant
differences also existed among months with maximum values in December
and January. Bacteria comprised the largest pool, and increased rapidly
from ,40 ug C/L at 8m to .300ug C/L at 10m. Stepwise linear regression
showed that variation in bacterial biomass (R25.61 p ,0.001) was related
to predator biomass and month. Photosynthetic nanoflagellates (PNAN)
comprised the second largest pool, and showed greatest seasonal values in
December, greatest annual values in 1998, and greatest values at 9m depth.
Regression analysis revealed significant relationships (R2 50.48 p , 0.001)
to predators, month and year. Relative contributions of bacteria and PNAN
changed with depth; bacteria increased below the chemocline whereas
PNAN declined below this depth. Rotifer biomass decreased with depth
and maximum values occurred in December and January. There was a
significant relationship (p50.0011) between rotifers, the top predator in
this system and the biomass of the other plankton groups. Heterotrophic
nanoflagellates (HNAN), herbaceous - and bacterial-feeding ciliates all increased significantly with depth (p50.0001), with maximum values at the
chemo-cline, with highest values in December and January. Variation in
herbaceous- and bacterial-feeding ciliates were related to the biomass of
their prey (p , 0.0001) and month.
MCKERNAN, PAMELA,1,* JOSEPH D. WHITE,1 KEVIN J. GUTZWILLER,1 WYLIE C. BARROW2 and LORI RANDALL.2 1 Baylor University, Waco, TX; 2 U.S. Geological Survey, National Wetlands Research
Center, Lafayette, LA. The effect of fire on a mixed Chihuahuan Desert
plant community.
The effect of fire on plant community composition and successional response was investigated in Big Bend National Park, Texas (1) locally by

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229

observing the short-term response of vegetation within a 500ha prescribed

burn and (2) regionally through land cover change analysis using Landsat5 Thematic Mapper (TM) data. We predicted that fire intensity and severity
within this patchy fuel environment would reduce overall shrub densities
and promote grass establishment. Predictions of long-term community
changes were based on trends observed between Landsat TM images of
1986 and 1999 and on seed bank analyses. Within the burn sites, the prefire characterizations of fuel distributions and vegetation cover indicated a
shrub-dominated community with disconnected fuels and vegetation canopies. Seed pools also indicated higher proportions of shrub species. Surface fire temperatures, rates of spread, and post-fire severity ratings were
related to fuel patchiness and pre-fire species composition. Post-fire seed
banks did not change appreciably due to the low intensity of the burn.
Shrubs were most reduced by the prescribed burn, while site patchiness
was significant in determining mortality rates. Analysis of landcover
change from Landsat TM derived vegetation maps between 1986 and 1999
indicated a region-wide proliferation of shrub communities. Comparison of
burned and unburned areas in the park showed no difference in the rate of
landcover change suggesting a negligible impact of fire in patchy shrub
dominated communities. This supports the idea that the occurrence of a
single fire in an extremely patchy fuel and vegetative environment may not
be sufficient in changing community composition.
MCLACHLAN, JASON S.,* JAMES S. CLARK and PAUL S. MANOS.
Biology, Duke University, Durham, NC. Conservation lessons from the
Last Glacial Maximum.
Some places, like Brazils Atlantic Forests, are hotspots of local species
endemism. In other places, like Amazonia, many rare species have geographically extensive overlapping ranges. Global conservation efforts increasingly focus on regions with the former kind of rarity, because species
there are thought to be more prone to extinction. The end of the last ice
age provides an interesting contrast to the modern extinction crisis because
most species did not go extinct, despite rapid environmental change. Many
currently common temperate tree taxa were rare at the last glacial maximum (LGM) and it would be useful to know if their survival was aided
by extensive diffuse populations. The consensus of the paleoecological
community, though not unanimous, is that rapid expansion from isolated
southern refugia (ice age hotspots) was common in Europe and in North
America. We show that reconstructions of LGM refugia using the fossil
record have suffered from systematic bias against the detection of widespread diffuse populations. Molecular data and a reanalysis of fossil pollen
data from eastern North America show that temperate hardwood species
thought to be restricted to isolated refugia during the late glacial, were, in
fact, widely distributed, though rare. Large ranges may have helped rare
species survive rapid environmental change in the past, as conservation
biologists hope they will today.
MCLAUCHLAN, KENDRA K.* and SARAH E. HOBBIE. University of
Minnesota, St. Paul, MN. Role of texture and vegetation in determining
soil organic matter dynamics of former agricultural fields.
Soil organic matter (SOM) formation over annual to millennial timescales
is an important component of ecosystem and soil development. Yet little
is known about how vegetation and soil texture influence SOM accumulation at decadal time scales on lands converted from agriculture to permanent vegetation cover. In particular, as SOM levels recover from depletion by agriculture, it is unclear how the dynamics of labile organic matter
fractions compare to those of more recalcitrant fractions, and how factors
such as soil texture and vegetation composition affect the dynamics of
different SOM fractions. We sampled a chronosequence of former agricultural fields in western Minnesota to determine how soil texture and vegetation characteristics, individually and interactively, control the rate and
type of SOM formation over 35 years. Total SOM contents depleted by
agriculture are increasing after establishment of perennial grassland. We
hypothesized that three characteristics of plant species - biomass production, tissue N concentration, and C chemistry - would be important determinants of labile SOM quantities. We also hypothesized that soils with
high clay content would accumulate recalcitrant SOM at a faster rate than
soils with low clay content. In contrast to our expectations, variation in
tissue chemistry between warm-season and cool-season grasses did not

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translate into significant variation in the type or quantity of SOM formed.

Also in contrast to our expectations, differences in soil parent material,
between clay-rich glacial till or sandy glacial outwash, did not predict the
rate or magnitude of SOM accumulation. These results suggest that much
of the variation in SOM dynamics during recovery from agriculture arises
from factors other than the composition of the established vegetation or
the texture of soil parent material.
MCMILLAN, BRETT A.,1 A. SCOTT BELLOWS1 and THOMAS R. ALLEN.2 1 Biological Sciences, Old Dominion University, bmcmilla@odu.
edu, asbell@vcu.org, Norfolk, VA; 2 Political Science and Geography, Old
Dominion University, tallen@odu.edu, Norfolk, VA. Application of a
Pseudo-NDVI image enhancement to coastal landscape ecology.
The Normalized Difference Vegetation Index (NDVI) is a long and widelyused vegetation index applied to combinations of spectral bands from digital remotely-sensed images and used to accentuate and measure vegetation
characteristics. NDVI uses red and infrared wavelengths, such as the multispectral sensors on Landsat, to detect chlorophyll and leaf structure variations in vegetation. The coarse spatial resolution of most satellite observations (e.g., 30 x 30 m pixels from Landsat Thematic Mapper) makes
them most appropriate for landscape-level investigations on a regional
scale. Although multi-spectral satellites with finer spatial resolutions are
becoming available, they often lack sensing abilities in the infrared wavelengths that are relevant to plant ecology. The primary historical source of
multi-spectral imagery with pixel size , 5 x 5 m is color-infrared (CIR)
aerial photographs, which lack visible red wavelengths. Having a need to
detect changes in vegetation over short distances (, 2 m) on a small island,
we developed a modified or pseudo-NDVI (P-NDVI) for use with CIR
aerial photographs. Our P-NDVI substitutes blue band values for red band
values in the same function used for original Landsat NDVI, with the
rationale that chlorophyll absorbs blue and red wavelengths nearly equally.
Furthermore, blue light scattering, a common problem with satellite images,
is reduced in aerial photography because of lower observational altitude.
Classification of vegetation at test locations using NDVI on Landsat images
and P-NDVI on aerial photographs produced comparable results. This protocol has an advantage over multivariate statistical classification procedures
such as principal components analysis (PCA), in that detected variation can
be attributed to true differences in vegetation with greater confidence. This
technique should be useful for conducting coastal vegetation and plant
ecology studies at a finer spatial scale and over a longer time range than
is available with satellite imagery.
MCNEELY, CAMILLE* and MARY E. POWER. University of California,
Berkeley, Berkeley, California, USA. Role of the armored grazer Glossosoma penitum in a headwater stream food web.
We investigated the effect of grazing by an invertebrate herbivore on the
energy sources used by other invertebrate consumers in a headwater stream
food web. We hypothesized that Glossosoma penitum might act as a sink
for energy produced by local algae in a small, forested stream in Northern
California. Glossosoma larvae are very abundant in the stream (average
midsummer densities were 10126194/m2 in 2000 and 10646430/m2 in
2001), and in 2000 they made up 48% 6 5% of the invertebrate biomass.
Stable isotope studies have shown that Glossosoma larvae in this stream
derive most of their carbon from local productivity, despite low algal standing crops. We hypothesized that the caddisfly might compete with other
herbivores for algal food, and increase their reliance on inputs of terrestrial
detritus. Because Glossosoma are defended from predation by a heavy
stone case, they may also reduce transfer of algal energy up the food chain.
We removed Glossosoma by hand from five 3m long sections of the stream
during the summer of 2000, and measured the response of algae, invertebrate herbivores (Heptageniid mayfly nymphs and Psphenid beetle larvae),
and invertebrate predators (Calineuria stonefly nymphs) in removal sections and paired controls. We examined the diets of consumers using stable
carbon isotope ratios. We also examined gut contents of one herbivore
group (Heptageniid mayflies). The treatment reduced Glossosoma biomass
by 80-90% compared to control plots, and chlorophyll-a density doubled
in removal sections (paired t-test of log-transformed values at 32 days: t
5 -2.92, p 5 0.043). However, there was no significant difference in the
d13C of invertebrate herbivores or predators, suggesting that increased algal

availability did not result in greater assimilation of algal carbon by these

consumers.
MCPHEARSON, P. TIMON,* KYLE F. BENNETT, ROBERT M. COX,
MATHEW I. PALMER and PETER J. MORIN. Rutgers University, New
Brunswick, NJ. Mutualism or slavery? Classical mutualists positively
affect associated communities albeit by symbiont host control.
With recent exceptions, ecological studies have not examined mutualism
beyond within-trophic level study, though theoretical implications for community-level effects have arisen from many studies. We used a model positive association between protists, Paramecium bursaria and its endosymbiont Chlorella, to test theory that posits mutualism should be more common and therefore a more important determinant of community dynamics
in ecologically stressful systems. We constructed tri-trophic level communities in aquatic protist microcosms along a temperature stress gradient in
light and dark incubators. We compared communities with mutualistic interaction to identical structured communities without functional mutualistic
interaction. Results demonstrate that communities with mutualists have
higher species abundances when under stress than communities without.
However, our mutualists may not exhibit simple reciprocally positive interactions as classically described. Instead, by assessing the effects on freeliving and endosymbiotic Chlorella, we reveal an apparent case of host
slavery which determines major treatment effects in our communities.
Analyses suggest that P. bursaria controls free-living Chlorella densities
which increases abundance for other species in the community when under
significant temperature stress. We reveal an often overlooked conditionality
inherent in many species interactions by showing the community context
that results in mutualism acting as slavery and further confirm that under
significant environmental stress communities may fair better when they
have mutualist species in them.
MCQUATTIE, CAROLYN J.* and JOANNE REBBECK. USDA Forest
Service, Northeastern Research Station, Delaware, OH, USA. Mycorrhizal
colonization of maple and oak seedling roots exposed to prescribed fire
and/or overstory thinning.
Disturbance from prescribed burning and/or thinning may change light and
moisture conditions to favor oak reproduction and limit competitors such
as red maple. However, the effects of these treatments on mycorrhizal colonization of oak and maple seedlings have not been evaluated. Thinning
and prescribed burning, alone and in combination, are being used to study
seedling responses to these disturbances in Vinton County, OH with four
treatment combinations: undisturbed control (U); overstory thinned by 29%
(T); prescribed burning (B); burning and thinning (BT). In June and August
2001 (first growing season following treatments), four seedlings (2 black
oak, Quercus velutina, ectomycorrhzial species, and 2 red maple, Acer
rubrum, endomycorrhizal species) from six plots per treatment were excavated from three subsites with different moisture levels (2 mesic, 2 intermediate, 2 xeric). Seedling root systems (48 per collection) were chemically preserved. Maple roots were cleared and stained, and percent endomycorrhizal roots per root system was quantified by point counting. In
oaks, all root tips were inspected microscopically for a fungal mantle or
Hartig net. Beaded oak roots displaying no obvious fungal mantle were
stained to determine if endomycorrhizal structures were present. At both
collection times, mycorrhizal colonization of maple roots ranged from 55%
to 76%. Colonization was lowest at xeric sites. Colonization increased between June and August in each treatment; the greatest percent increase
(15%) occurred in BT, possibly due to increased light and less competing
vegetation. Oak roots were predominantly ectomycorrhizal. Colonization
was highest in June for the U treatment (69%) and in August for the T
treatment (61%). Beaded oak roots with endomycorrhizal structures (hyphal
coils, arbuscules, vesicles) were observed at both collection times in the B
and BT treatments. On disturbed sites, oak roots may form both ectomycorrhizal and endomycorrhizal associations that contribute to increased nutrient uptake and seedling survival.
MEADOR, ANDREW J., PABLO PARYSOW* and MARGARET M.
MOORE. Northern Arizona University, Flagstaff, AZ. Reconstructing
ponderosa pine presettlement forest structure in northern Arizona using permanent plots.
We present preliminary results of an on-going study to reconstruct presettlement ponderosa pine forest structure in northern Arizona. We base our

reconstruction on post-harvest tree data collected between 1909 and 1913

from 7 permanent plots on the Coconino National Forest. Plots were established at the time of data collection. Data contain information on live
trees with diameter at breast height (dbh) of at least 4 inches, stumps, snags,
and downed dead trees. Plot sizes range from 3 to 10 acres. Using the
GENGYM (Generalized Growth and Yield Model) system as well as dendrochronological techniques, we modeled the forest structure that would
have existed at the time of plot establishment accounting for disturbances
caused by European settlement. We present dbh frequency distributions, as
well as average number of trees per acre, basal area per acre, and quadratic
mean diameter.
MEGONIGAL, PATRICK,1,* JOHANNA WEISS,2 SCOTT NEUBAUER1
and DAVID EMERSON.3 1 Smithsonian Environmental Research Center,
Edgewater, MD, USA; 2 George Mason University, Fairfax, VA, USA;
3
American Type Culture Collection, Manssas, VA, USA. Mechanisms
promoting a wetland rhizosphere ferrous wheel.
Competition between anaerobic microorganisms for electron donors suppresses methane emissions from wetlands. Fe(III) reducing bacteria outcompete methanogens for H2 and acetate, but relatively little is known
about the mechanisms by which Fe(III) oxides are deposited and reduced
in wetlands. We investigated Fe(III) oxide deposition in the rhizosphere of
wetland plants and determined that the process does not proceed strictly
by chemical oxidation as once believed. Chemolithotrophic Fe(II)-oxidizing bacteria were ubiquitous in wetlands and mediated up to 60% of Fe(III)
deposition in laboratory studies. Furthermore, the mineralogy of rhizosphere Fe(III) oxides favored rapid reduction by Fe(III)-reducing bacteria.
Labile, poorly-crystalline Fe(III) oxides were 66% of the rhizosphere Fe
pool, compared to 23% of the bulk soil Fe pool. As a result, the rhizosphere
Fe(III) oxides were consumed in respiration at twice the rate of those in
the bulk soil during anaerobic incubations. This difference persisted when
the incubations were spiked with an Fe(III)-reducing bacterium (Geobacter
metallireducens) and an electron-shuttling compound (AQDS), and the
same pattern was observed during a year-long field experiment in which
90% of rhizosphere Fe(III) oxides were reduced compared with 40% in the
bulk soil. We concluded that the wetland plant rhizosphere is a "hot spot"
of Fe cycling because roots are ideal sources of all the substrates required
by both Fe(II)-oxidizing and Fe(III)-reducing bacteria, including labile
Fe(III) oxides, labile organic C, O2, and Fe(II) (from Fe oxide reduction
or diffusion from wetland porewater). This work helps explain why the
presence of plants reduces the proportion of organic carbon that is metabolized by methanogens.
MEIER, OUIDA W.,* ALBERT J. MEIER and SCOTT GRUBBS. Western
Kentucky University, Bowling Green, KY, USA. Relationship between
landuse and stream conditions in the karsted upper Green River watershed of Kentucky.
The Green River watershed in Kentucky, with 151 fish species and 73
mussel species (nine of which are endangered), has an aquatic faunal community that is among the highest in biodiversity in the U.S. The most
diverse and least impacted sub-basin of this watershed, the upper Green
River watershed, has been targeted for a USDA Conservation Reserve Enhancement Program (CREP) to convert pasture and crop land to riparian
buffers along the Green River and its tributaries. We initiated a sampling
program to develop a comprehensive picture of pre-enhancement conditions in the region. Within our sampling program, measurements of water
quality, fish and macroinvertebrate species, and habitat parameters were
taken on 43 sites in the upper Green River basin during 2002. Using a
geographical information system (GIS) approach, we have quantified landuse for the watershed and at smaller scales within the watershed. Some
of the water quality parameters related most strongly to current landuse
patterns are fecal coliform bacteria, total suspended solids, and turbidity.
High fecal coliform values in this system are related to both animal input
(from animal feeding operations) and human input (from failing septic systems and straight pipes). The presence of exceptionally well-developed
karst in parts of this watershed make inputs of both human and animal
waste much more difficult to keep out of the aquatic ecosystem. As the
CREP plan is implemented, it will be important to separate human from
animal inputs of fecal coliform in order to assess the programs effective-

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231

ness in protecting the watershed and the high biodiversity of the upper
Green River.
MEINERS, SCOTT J.,1,* MARY L. CADENASSO2 and STEWARD TA.
PICKETT.2 1 Eastern Illinois University, Charleston, IL; 2 Institute of Ecosystem Studies, Millbrook, NY. Predictability of within-site invasion dynamics for native and exotic plant species.
The rate of within-site spread is a key diagnostic for setting management
priorities as slower invaders tend to be lower management priorities than
faster invaders. However invasion dynamics are rarely quantified. Here we
present data from the Buell-Small Succession Study, a long-term study of
post-agricultural vegetation dynamics, to evaluate the dynamics of plant
species invasions over a 44-year period. The analyses focus on a range of
species of varying successional status, life history and form. We have determined the rates of invasion for the 20 most abundant native and exotic
invaders at the site (40 species in total). Based of rates of increase in
frequency of occurrence in the sites, native and exotic species do not differ
in invasion rate. However, plant life forms vary predictably in rate of invasion. Short lived and herbaceous plants invade at much greater rates than
more long-lived woody species. These life forms follow the following ranking from fastest to slowest invaders: annuals . biennials . perennials .
woody spp. The length of population lag times (the time between when an
invader first appears in a site and when its population begins to grow)
follows the same ranking, with annuals having the shortest lag times. These
results suggest that short-lived invaders may pose more immediate management concerns than longer-lived species. This information provides land
managers with a simple way to assess management priorities without extensive population monitoring. Furthermore, exotic and native plant invasions appear not to have fundamentally different dynamics.
MEIR, ELI,* ELEANOR STEINBERG and SUSAN MARUCA. SimBiotic
Software for Teaching and Research, Inc., Ithaca, NY, 14850. The next
EcoBeaker: Introducing EcoBeaker 2.5 for introductory biology and
beyond.
EcoBeaker is the worlds leading software for teaching ecology, conservation biology, and evolution, used in 300+ universities and almost 100
high schools. The software teaches by constructing simulated communities
of species in the computer and then asking students to do experiments on
those communities, as a practicing ecologist would, to discover important
concepts. We are releasing a major upgrade of EcoBeaker in spring 2003,
and pre-release users have been extremely positive about the direction
weve taken. This talk will highlight some of those changes. Among the
new features are: several new labs, including labs on life tables, on harvesting natural resources, and on optimal foraging; major improvements to
some of our older labs; a new workbook format for the labs that is especially useful in larger and introductory courses; a better looking and simpler
to use interface; and more. At the end well also discuss some of our plans
for future versions of EcoBeaker and related teaching software we are
developing.
MELBOURNE, BRETT.* University of California, Davis, CA, bamelbourne@ucdavis.edu. Fitting nonlinear population models to data with
both process and measurement error.
Most models fitted to data are of the general form N(t+dt) 5 F(N(t)), where
abundance or density N is a function of N at a previous time. We want to
estimate parameters for the function F, given data N(t+dt) and N(t). Dependent variables other than N, such as abiotic variables, might also be
included in F. All of these models are fitted to data most commonly assuming that N(t) and other dependent variables are measured without error,
or less often assuming that N(t+dt) is free of process error. These assumptions are rarely met for real data. Most data contain both process error,
which arises because the true process deviates from the model, and measurement error, which arises because the measured data deviate from the
true values. It is well known that ignoring either source of error results in
biased parameter estimates. Cook and Stefanski (1994) proposed a simulation based method of model fitting (SIMEX) related to jackknife estimation, with wide application to nonlinear models when the measurement
error variance is known or can be estimated. I studied the performance of

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Abstracts

SIMEX for the logistic model of population growth with both process and
measurement error. For simulated data, SIMEX provided unbiased estimates of growth rate r, carrying capacity K, and process error variance. In
contrast, naive estimates under the assumption of no measurement error
overestimated r, underestimated K, and overestimated process error. I used
SIMEX to fit the logistic model to data from a density perturbation experiment, the aim of which was to measure density-dependence in the growth
of benthic stream algae. The data were initial and final biomass. The variance component for the measurement error was obtained separately from
replicate samples in a nested sampling design and a bootstrap approach
was used to obtain confidence intervals for r and K.
MELILLO, JERRY.* The Ecosystems Center, Marine Biological Laboratory, Woods Hole, MA, USA. Science, policy and the grand element
cycles.
Cycles of several of the grand elements, including carbon and nitrogen,
have been disrupted by human activity. Since the beginning of the Industrial Revolution humans have caused the carbon dioxide content of the
atmosphere to increase by more than thirty percent. Over this same period,
we have more than doubled the amount of reactive nitrogen cycling through
the biosphere annually. In recent years, interdisciplinary teams of scientists
have conducted assessments of the disruption of the carbon and nitrogen
cycles and reported their findings to policy makers and the public. Examples include reports from the Intergovernmental Panel on Climate Change
and the First and Second International Nitrogen Conferences. Scientists
have made it clear to policy makers that these disruptions have, or will
soon have, many negative consequences for people and other life on earth.
In response, policy makers, relying on sound science, have begun to frame
protocols for managing the grand cycles of carbon and nitrogen in ways
that minimize negative environmental effects while fostering economic
growth. As they continue their work, policy makers will rely even more
heavily on natural and social scientists, economists and engineers partnering to provide management options for a host of environmental issues
associated with element-cycle disruptions. Prime candidates for immediate
attention are the stabilization of the atmospheres carbon dioxide concentration and the reduction of eutrophication of the worlds estuaries and
coastal waters.
MELLARD, JARAD P.* and BRYAN L. FOSTER. University of Kansas,
Lawrence, KS. Effects of initial conditions and resource supply on species assemblage dyamics: A microcosm study.
Community assembly theory predicts that initial conditions, invasion sequences, and priority effects are important in regulating the development
of communities. An important goal in ecology is to isolate the ways in
which historical processes and deterministic mechanisms (competitive
dominance, species traits etc.) may influence communities of interacting
species. We present results from the first year of a plant microcosm experiment designed to isolate the role of initial conditions (initial species
abundances) on multispecies interactions and development of community
structure. Initial relative abundances of eight grassland species were manipulated at two levels of nutrient supply to assess: (1) the degree to which
assemblage dynamics are affected by initial conditions versus deterministic
effects of particular species; and (2) how nutrient availability may alter the
effects of initial conditions on assemblage dynamics. Abundance distributions at the end of the first growing season strongly reflected initial conditions, but also exhibited the dominant effects of two highly competitive
species. Seedling growth rate was a good predictor of a species relative
abundance in some treatments at the end of the first growing season. Resource levels interacted with initial conditions and species-specific effects
to influence community states by altering the abundance hierarchy. Preliminary results indicate that: (1) both initial conditions and species-specific
traits can interact in complex ways to regulate community development;
and (2) the nature of these complex interactions may be modified substantially by resource availability.
MENDELSSOHN, IRVING A. and MATTHEW G. SLOCUM. Louisiana
State University, Baton Rouge, LA, USA. Effects of nutrient enrichment,
disturbance, and herbivory on vegetation and soils in an oligohaline
marsh.
Ecosystems are maintained in a relatively stable state by background levels
of disturbance and stress, and multiple stressors may interact in important

ways. Alteration in these stress and disturbance levels may therefore lead
to loss of ecosystem integrity and diversity, especially if multiple stress or
disturbance factors are affected. We studied how multiple stressors/disturbances affected soils and vegetation in an oligohaline marsh on the north
shore of Lake Pontchartrain, Louisiana. Stressors/disturbances included
herbivory, whose background levels have been raised with the introduction
of nutria (Myocaster coypus), and a nutrient addition treatment, which simulated increased inputs due to agricultural runoff. We also applied a lethal
disturbance (Roundup application), which allowed assessment of recovery
rates of vegetation and soils while stressed by herbivory and nutrient addition. All stressors affected vegetation and soils, and multiple stressors
sometimes reinforced effects. Lethal disturbance and herbivory interacted
to reduce biomass recovery; without fences biomass only reached 43% of
controls over four years, as herbivores preferred fresh shoots in lethally
disturbed plots. In contrast, within fences biomass gradually reached control levels. Fertilizer increased biomass up to 70%, but the effect was highly
variable and of moderate significance, and did not offset the effects of
herbivory. However, fertilizer consistently reduced species richness from
five species to four species per plot, increased decomposition rates by 20%
over control levels, and elevated exchangeable NH4-N and P in the soil.
Exchangeable NH4-N was also increased by lethal-disturbance and herbivory, such that when combined with fertilization it reached 4.5X over control
levels. This research shows that multiple stressors can have additive, negative effects on the diversity and resilience of this oligohaline marsh. In
addition, increased soil NH4-N and the accompanying increase in decomposition may lead to a lowering of elevation, resulting in higher water
levels and consequences for ecosystem health.
MENGE, BRUCE A.,1 JANE LUBCHENCO,1 STEVEN D. GAINES,2
FRANCIS CHAN,1 MELISSA FOLEY,1 GREG HUDSON,1 KARINA J.
NIELSEN1 and JACQUE PAMPLIN.1 1 Department of Zoology, Oregon
State University, Corvallis, OR, USA; 2 Department of Ecology, Evolution
and Marine Biology, University of California, Santa Barbara, CA, USA.
Dramatic changes in rocky intertidal communities associated with
oceanographic regime shifts.
Marine community theory predicts that community structure and dynamics
should vary both with rates of propagule input (larval transport model or
LTM) and food (phytoplankton and detritus) subsidies (bottom-up model
or BUM). Both models predict that with increases in subsidies, rates of
recruitment, predator abundance, and predator effect should all increase.
They make opposite predictions for prey abundance, which increases with
subsidies in the BUM but decreases with more recruitment in the LTM.
Because larval transport rates and food subsidies in coastal habitats vary
over tens to thousands of km, tests of such models have lagged. Recent
studies in New Zealand are generally consistent with both models, except
that contrary to the LTM prey abundance increases rather than decreases
with recruitment. Studies on the US west coast, in concert with a recent
oceanographic regime shift, provide an opportunity to examine these predictions across appropriate scales of space and time. The Pacific Decadal
Oscillation (PDO) is a large scale (Pacific Ocean Basin) long-term (25 year)
cycle of change in surface temperature. In about 1998, the Northeast Pacific
shifted from a warm phase to a cold phase. Long-term studies of intertidal
community dynamics underway since 1989 (Oregon) and 1995 (California)
have revealed major changes in several factors. In Oregon, but not California, upwelling intensity increased, average water temperature decreased,
and concentration of phytoplankton, recruitment of mussels, and abundance
of mussels on the shore all increased. Prior research suggests that the increase in phytoplankton results from higher nutrient inputs due to more
intense upwelling, and that this higher level of food supply enhances mussel larval growth and survival. Observed changes to date are consistent
with all predictions of the bottom-up model and, again excepting prey
abundance, the larval transport model. Ongoing investigation through the
PISCO project will permit a detailed mechanistic understanding of the ecological consequences of these changes.
MENGE, DUNCAN L.1,* and CHRISTOPHER B. FIELD.1,2 1 Stanford
University, Stanford, CA; 2 Carnegie Institution of Washington, Stanford,
CA. Phosphorus dynamics under manipulated global change conditions in a California annual grassland.
Global environmental factors changing with anthropogenic influence include increased CO2, NO3- deposition, warming, and precipitation. The Jas-

per Ridge Global Change Experiment (JRGCE) at Stanford Universitys

Jasper Ridge Biological Preserve manipulates all four of these factors in
an annual grassland ecosystem, maintaining all combinations of ambient
and elevated treatments for each factor. Phosphorus limitation may influence the extent to which these manipulations affect ecosystem processes
such as NPP. This study investigates this biogeochemical feedback in the
JRGCE. Extracellular phosphatases, enzymes secreted by plants and microbes, have been shown to indicate phosphorus demand in this system
and others. Soil phosphatase activities were measured to assess ecosystem
level P demand under these manipulations. Elevated NO3- significantly increased phosphatase activity, indicating higher P demand. Elevated precipitation significantly decreased phosphatase activity, indicating lower P demand. CO2, warming, and all interaction terms were not significant.
MENNINGER, HOLLY L.1,* and MARGARET A. PALMER.2 1 Behavior,
Ecology, Evolution & Systematics Program, University of Maryland, College Park, MD, USA; 2 Department of Entomology, University of Maryland, College Park, MD, USA. Terrestrial-aquatic linkages: Herbaceous
vegetation and headwater streams.
Open-canopy streams with dense edges of herbaceous vegetation are common throughout agricultural landscapes in the Mid-Atlantic region, yet the
interaction between vegetation and invertebrate dynamics of such streams
is rarely studied. Along headwater streams, edge vegetation bridges the
terrestrial2aquatic interface and may play important roles in channel shading, creating habitat structure, and providing a detrital resource. We removed the edge vegetation from three open-canopy headwater streams and
measured the removal effects on 1.) rates of key ecosystem processes (primary production, organic matter decomposition) and 2.) diversity of aquatic
invertebrates as compared to reference streams and pre-manipulation observations. We predicted that vegetation removal would result in a shift
from an apparent detritus-based system to a primarily algal-driven system.
With changes in food resource base, we anticipated a decrease in the abundance of detritivorous invertebrates as well as an increase in algivorous
invertebrates. We also predicted that removing edge habitat structure would
result in a decline of sedentary invertebrate predators. We found increases
in primary production, the result of greater insolation to the stream channel,
although algae did not grow homogeneously throughout the stream reaches.
Rates of leaf decomposition of a common grass did not change significantly
as a result of vegetation removal. Shredding invertebrates (Isopoda, Lepidostomatidae) declined across reference and experimental streams, suggesting no effect of manipulation on this feeding guild. Other invertebrate
feeding groups, however, did respond to the removal treatment. Algivorous
midge larvae (Chironomidae) increased nearly four-fold following vegetation removal and the subsequent increase in algal production. The number
of predaceous odonate larvae (Calopterygidae) declined significantly following vegetation removal, a consequence of the elimination of important
habitat structure. These results suggest that herbaceous vegetation growing
along the edges of open-canopy headwater streams may play key roles in
regulating in-stream food resources and structuring the aquatic invertebrate
community.
MENTZER, JESSICA L.,1,* ROBERT M. GOODMAN2,3 and TERI C.
BALSER.1 1 Soil Science Department, University of Wisconsin-Madison,
Madison, WI; 2 Gaylord Nelson Institute for Environmental Studies, University of Wisconsin-Madison, Madison, WI; 3 Plant Pathology Department, University of Wisconsin-Madison, Madison, WI. Seasonal changes
in wetland microflora under varied nutrient and hydrologic regimes.
Wetlands are a unique ecosystem type, especially in terms of nutrient cycling under fluctuating hydrology. This makes them important to studies
of ecosystem functioning. Many processes of interest in wetlands, such as
denitrification and decomposition, are mediated by microbes; therefore,
characterization of microflora composition and function may lead to insights about the dynamics of wetland ecosystem functioning and nutrient
cycling. In this experiment, we measured several microbial parameters over
the growing season in a simulated wetland undergoing varied nutrient and
hydrologic regimes. Mesocosms were grown for two years with native
Wisconsin wetland species and in the summer of 2002 were subject to
three nutrient (none, low and high) and three hydrologic (early season,
constant, and intermittent floods) regimes in a full factorial experiment with

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233

five replications of each treatment. We collected soil cores at four times:

1) in late May, before treatments started, 2) at two different peaks in the
vegetation growth (August and September), and 3) after senescence of the
aboveground biomass in early December. We assayed each sample for nitrification potential, denitrification potential, and various enzyme activities
(phosphatase, beta-glucosidase, alpha-glucosidase, chitinase, phenol oxidase, cellobiohydrolase, and hemicellulase ). We also performed lipid analysis to determine microbial community structure and composition. Results
show that nitrification potentials decreased significantly with increased nutrient levels and that the intermittent flood regime had higher nitrification
potentials than early season or constant flood regimes. These results indicate that the nutrient and flood regimes did in fact impact microfloral functions that are important to the wetland system.
MERCHANT, AMETHYST G.* and STEPHEN S. MULKEY. University
of Florida, Gainesville, Florida. Effects of nitrogen and phosphorus
availability on Dicerandra (Lamiaceae) congeners with different lifehistories.
Endemic to the southeastern United States, the genus Dicerandra consists
of nine species displaying different growth habits in specific ecosystems.
Annuals range throughout recently disturbed sandhill habitats of Georgia
and northern Florida. Perennials are limited to small, geographically isolated scrub habitats along relict beach ridges and dunes within Floridas
peninsula. Scrub has been described as nutrient deficient, notably in phosphorus, relative to other habitats. We applied a factorial experiment crossing three levels of nitrogen and phosphorus within two blocks to individuals of D. densiflora (herbaceous annual) and D. cornutissima (semi-woody
evergreen perennial) to determine how variation in nutrient availability affects growth, biomass allocation, gas exchange, and nutrient concentration
of tissues. Annuals were expected to increase growth and fruit production
with changes in biomass allocation while perennials exhibit greater nutrient
accumulation and little change in growth and biomass allocation as nutrients increase. Total dry weight increased with nutrient availability for both.
Biomass allocation to floral structures (including seeds) in D. densiflora
was relatively consistent throughout treatments. Stem, leaf, and root weight
ratios were also similar throughout treatments for this annual. Root : shoot
ratio decreased significantly for D. cornutissima as nitrogen availability
increased. Root : shoot ratio was significantly higher in D. cornutissima.
Significant differences in maximum photosynthetic rates (area-based) were
not apparent due to great variability within treatments although larger rates
were expected as phosphorus availability increased. Specific leaf area,
branch production, and tissue nutrient concentrations will be discussed in
reference to interactions between nitrogen and phosphorus availability.
METZGER, KRISTINE L.,1,* MONICA G. TURNER1 and WILLIAM H.
ROMME.2 1 University of Wisconsin-Madison, Madison, WI; 2 Colorado
State University, Fort Collins, CO. Foliar nitrogen in early postfire vegetation in the Greater Yellowstone Ecosystem.
Foliar nitrogen (N) and carbon (C) concentrations were studied at two sites
that differed in substrate and pre-fire species composition in the Greater
Yellowstone Ecosystem following stand-replacing fires in 2000. In this paper we addressed three questions: (1) Do foliar N concentrations and foliar
C:N ratio differ among dominant herbaceous species and/or functional
types after fire? (2) Do foliar N concentrations and foliar C:N ratios in
dominant herbaceous species differ between burned and unburned forests
at the same site? (3) Does total foliar N and foliar N contributions by
species and functional types differ by site, soil nitrogen availability, or fire
severity? We analyzed dominant herbaceous vegetation for carbon and nitrogen content. Vegetation was sampled in areas that burned in 2000 and
adjacent areas that escaped the burn. Total biomass, plot level C:N ratio,
and foliar N were determined by sampling ten 0.25-ha plots for percent
cover of herbaceous vegetation and then applying species-specific allometric equations to convert cover to biomass. Species foliar N concentrations ranged from 0.70 to 3.57% and C:N ratios ranged from 12 to 66;
both differed significantly among species and functional types. Nitrogen
fixers (Lupinus argenteus, Ceonothus velutina) contained the highest N
concentration and a grass (Calamagrostis Canadensis) the least. Five species had significantly higher N in burned plots than in unburned plots (10
to 41%), and three species showed no differences. At the plot level, the

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Abstracts

contribution of N fixers was negatively correlated to inorganic N availability, whereas plot-level C:N ratio was positively correlated. Total foliar
N and total aboveground biomass did not vary by site, inorganic nitrogen
availability, or fire severity. Our results suggest remarkable resilience to
the effects of severe fire at the ecosystem level; despite substantial variation
in foliar N among species, different species assemblages produced similar
values in aboveground biomass and total foliar N.
MEYER, CLIFTON W.,1 DAVID D. BRESHEARS,1 FAIRLEY J.
BARNES1 and ORRIN B. MYERS.2 1 EES-2, Earth and Environmental
Sciences Division, Los Alamos National Laboratory, Los Alamos, NM,
USA; 2 Center for Population Health, University of New Mexico, Albuquerque, NM, USA. Drought-induced mortality following dampened
soil moisture pulses: Results from a semiarid woodland.
Drought can produce landscape-scale mortality of both woody and herbaceous plants, as has been documented in response to the 1950s drought in
southwestern USA. Recent years have produced the most intensive drought
in parts of the southwestern USA since the 1950s drought, with extensive
observed tree mortality. Here we quantify mortality at a semiarid pinyonjuniper (Pinus edulis and Juniperus monosperma) site, Mesita del Buey,
located in northern New Mexico. We evaluated drought impacts using a
stand map obtained in the early 1990s and updated in 2003, and continuous
soil moisture measurements prior to and during the drought. We document
substantial mortality of pinyon but not juniper in response to the drought.
We also document the magnitude to which soil moisture pulses were attenuated during the drought. Our results have implications for improving
our understanding of drought-induced tree mortality, which is likely to
become more important at landscape scales as climate change progresses.
MEYER, EUGENE.* Loyola College, Baltimore, Maryland, Baltimore,
MD, USA. For camouflaged and elusive species, the concept of Detection Space. Example: midsize predators.
It is easy to miss the largest predators in an animal community if the
predators are well camouflaged and elusive. Such is the case with snapping
turtles in my field searches, and I suggest for other midsize predators in
the eastern US. The evidence: I recorded parallel searches for living amphibians and reptiles by experienced observers. I compared the results to
my field experiments for non-living non-moving animal replicas. I found
that a few species are so close to the threshold of detection that they are
entirely missed by some observers and by generalized search methods. The
most-missed taxa range from the smallest or most specialized salamander
and frog species, to the largest reptile present. I introduce the concept of
Detection Space to clarify detection effort and results. Detection Space is
shown simply on transparencies upon conventional graphs of community
structure. The use of Detection Space clarifies which species and life stages
have received suitable field effort so that a zero record represents a true
current absence. The wider utility of this approach is to behavior and camouflage that strongly reduce conspicuousness. Specific applications are to
impacts of midsize predators in the eastern US suburbs. The depauperate
condition of those communities is variously ascribed to one of three factors:
roads, habitat fragmentation, and farm loss. We can take the biological step
of asking if those three factors also increase the local densities of a handful
of common midsize predators, reptile and mammalian. What that trophic
level has in common is elusiveness as individuals and their impacts by
generalized field methods. As an indication of density and potential impacts, I return to the snapping turtle example because they, once found,
could equal the density of all the amphibians at a site combined.
MEYER, HARRY A.,* JULIANA G. HINTON and KATHLEEN TRAHAN. McNeese State University, hmeyer@mail.mcneese.edu, Lake
Charles, Lousiana. Spatial variability of tardigrade populations in leaf
litter, moss, and lichens.
Tardigrades are common inhabitants of lichens, mosses, and leaf litter. Although tardigrades in these habitats are often said to be very patchy in their
distribution, this assessment has not often been backed by quantitative sampling. In this study we assess spatial variability in tardigrade populations
in leaf litter, and in the mosses and lichens found on tress and rocks. To
assess spatial variability in tardigrade populations in lichens and mosses,

tardigrades were enumerated in 20-30 isolated lichen or moss patches of

varying sizes collected in 2002 and 2003 from adjacent trees in central
Florida and from rocks in the Ouachita Mountains, Arkansas. Variation
among lichen or moss patches within rocks or trees was very high; the
only consistent pattern was that very small patches generally lacked tardigrades. Tardigrade diversity and abundance also varied greatly within
sites when lichens and mosses of the same species were compared from
different rocks and trees (in the most extreme case, comparing two adjacent
trees harboring the same lichen species, one tree had numerous individuals
of three tardigrade species present while the other had no tardigrades). To
assess spatial variability in tardigrade populations in leaf litter, four leaf
litter core samples (10 cm diameter) were collected from each of two sites
in southwestern Louisiana and from one site in central Florida. Tardigrade
species richness and abundance varied greatly not only among sites but
among samples within sites (at one site in 4 adjacent cores tardigrade abundance and richness ranged from 0 individuals in a sample to 86 individuals
representing 5 species). Our quantitative sampling supports the assertion
that tardigrades are very patchy in distribution. Given the considerable time
investment required for the quantitative processing of such samples, this
high spatial variability in tardigrade diversity and abundance may make
them unsuitable for rigorous testing of ecological hypotheses.
MEYER, JUDY L.,1,* MERRYL ALBER,1 WILLIAM DUNCAN,1 MARY
FREEMAN,1 V. CODY HALE,1 RHETT JACKSON,1 CECIL JENNINGS,1
KIM LUTZ,2 MONICA PALTA,1 ELIZABETH RICHARDSON,2 BRIAN
RICHTER,4 REBECCA SHARITZ3 and JOAN SHELDON.1 1 University
of Georgia, Athens, GA; 2 The Nature Conservancy, Savannah, GA; 3 Savannah River Ecology Laboratory, Aiken, SC; 4 The Nature Conservancy,
Charlottesville, VA. Specifying ecosystem flows for the Savannah River.
To balance human and ecosystem needs for water in a regulated river, it is
essential to identify ecosystem flow requirements. We have been engaged
in a collaborative project to identify a flow regime for Thurmond Dam that
would sustain ecosystems in the Savannah River and its floodplain. Using
existing information sources, we characterized the current flow regime and
identified flow requirements for various life history stages of representative
species and ecosystem components in the shoals, river mainstem, floodplain
and estuary. The post-dam hydrologic regime is characterized by much
lower flood peaks, 7-day low flows that are higher and more variable in
timing, and higher baseflows during the driest months. In addition, channel
morphology and floodplain access have been altered by dredging; and water
movement and salinity in the estuary have been impacted by harbor deepening. We identified the magnitude and timing of flows needed for egg and
larval development, juvenile growth and survival, adult foraging, and
spawning migration for fishes such as robust redhorse, Atlantic and shortnose sturgeon, striped bass, and American shad. We documented the flow
requirements for seedling recruitment and tree growth and survival for
floodplain plant species such bald cypress and water hickory, and outlined
the consequences of altered plant communities and flood regime for floodplain birds and mammals. We developed simple conceptual models linking
hydrologic changes with their likely ecosystem consequences. The background information and conceptual models guided the development of recommendations on flow regime, which were formulated during a workshop
with stakeholders.
MEYERSON, LAURA A.,1,* JAMIE K. REASER,2 KRISTINE CIRUNA3
and ANN BARTUSKA.4 1 AAAS Environmental Fellow, Global Invasive
Species Programme, Washington, DC; 2 Global Invasive Species Programme, Washington, DC; 3 Nature Conservancy of Canada, The Nature
Conservancys Freshwater Initiative, 202 - 26 Bastion Square, Victoria, BC,
CA; 4 The Nature Conservancy, Ballston, VA. An assessment of the biological and socio-economic impacts of invasive alien species on freshwater ecosystems.
The Global Invasive Species Programme (GISP), on behalf of the Convention on Biological Diversity, held a workshop in June 2003 to bring together a small group of experts to assess the biological and socio-economic
impacts of IAS on freshwater ecosystems globally. The questions addressed
included: What are the major biological and socio-economic impacts of
IAS on freshwater systems? What variables influence the impacts of IAS
on freshwater ecosystems? What are the best practice measures for the

prevention, eradication, and control of IAS in freshwater ecosystems? How

do the principles/practices differ from other systems? What gaps in knowledge exist and how can they be filled? The outcomes of the workshop are
being used to develop a report to the Convention on Biological Diversity
(CBD) and Ramsar Convention on Wetlands, as well as publications for
resource managers and conservation ecologists. The assessment of the impacts of IAS on freshwater ecosystems is one of four ecosystem-based
assessments to be conducted by GISP for the CBD. A similar workshop
was held in October 2002 in Hawaii to address IAS in island ecosystems.
Funds permitting, future studies will assess the impacts of IAS in arid and
semi-arid grasslands and tropical forests.
MIAO, SHILI* and BRYAN GOASSMAN. South Florida Water Management District, West Palm Beach, FL. Effects of water depth and quality
on the growth of various oligotrophic-wetland plants.
One of the greatest challenges in ecosystem restoration is to restore native
vegetation in disturbed areas, as both biotic and abiotic conditions in these
sites have been altered tremendously. In impacted wetlands, commonly soil
and water nutrients are elevated and hydrological regimes are changed.
Whether native vegetation can be established and restored in such impacted
areas is largely species-specific. The objective of the present study was to
examine the effects of water depth and the outflow water quality of a
Stormwater Treatment Area (STA) on five species (Nymphaea odorata,
Eleocharis cellulosa, Eleocharis elongata, Rhynchospora tracyi, and Cladium jamaicense) without the competition of cattail, a dominant species in
nutrient-enriched wetlands. These species occur naturally in pristine Everglades wetlands with water total phosphorus (TP) of 10 ug/L and represent three dominant native vegetation types with distinct hydrologic regimes. The experiment was set up in nine outdoor fiberglass tanks with a
flow-through system using the outflow water from STA 1-West. This water
had an average TP of 25 ug/L. Seedlings of the five species grown in
nutrient enriched soil were placed in the tanks at two water depths (20 and
60 cm) for one year. Water depth affected the survivorship of the five
species differently. As water depth increased, marsh species experienced
the highest mortality, followed by wet prairie, and finally slough. One
slough species, Nymphaea odorata, experienced zero mortality at both water depths. For surviving plants, total biomass of all five species increased
by a range of 5 to 20 times. The preliminary results suggest that, in the
absence of cattail competition, these native plant species can grow under
the examined water quality condition. However, hydrologic regimes, such
as water depth, are key in the initial establishment and colonization of these
plants.
MICHALETZ, SEAN T.* and EDWARD A. JOHNSON. University of Calgary, Calgary, Alberta, CANADA. Tree mortality following surface fire:
A heat transfer model for predicting vegetative bud necrosis.
We have developed a heat transfer model for predicting vegetative bud
necrosis in the buoyant plume above a surface fire. This is part of a larger
program for post-fire tree mortality. The model indicates that time to bud
necrosis is governed by the bud-plume temperature difference and five
thermal bud properties which vary with bud water content, bud size, and
foliage architecture. This results in a logarithmic relationship between
plume temperature and time to bud necrosis. We validated the model using
buds of twelve tree species collected in the southern Canadian Rocky
Mountains and the prairie-forest transition zone of Wisconsin. The model
explained 69.5 % of the observed variation in time to bud necrosis. To
assess variation in bud thermal properties and its effect on time to bud
necrosis, we measured buds of four species from February to November
2002. The bud-plume temperature difference, bud size, and foliage architecture had strong effects on time to bud necrosis while bud water content
had a negligible effect.
MICHELI, FIORENZA,* KIMBERLY W. HEIMAN, CARRIE V. KAPPEL and ANDREW O. SHELTON. Stanford University, Hopkins Marine
Station, Pacific Grove, CA, USA. Persistence and recovery of depleted
marine invertebrates in marine reserves of central California.
An understanding of the processes that allow populations and communities
to persist and recover following disturbance is critical to many important

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235

questions in ecology and conservation biology. Marine protected areas

(MPAs) represent a powerful approach to conserving and promoting recovery of marine species and habitats impacted by fishing and other human
activities. However, certain scenarios may prevent recovery or lead to further declines, even within MPAs. These include: extreme population reductions prior to protection; protection of unsuitable or degraded habitats;
and intervening natural disturbances, such as extreme weather events, predation or disease outbreaks. We investigated patterns and processes of persistence and recovery of depleted invertebrate populations, red (Haliotis
rufescens) and black (H. cracherodii) abalone, in central California. Abalone densities, size structure, and predation mortality were compared along
a time-series of no take and de facto reserves (sites where access is limited
by private property) established in central California between 1931 and
1997 and protected for varying lengths of time. Longterm persistence was
examined within a permanent site established within a no-take marine reserve in 1972. Protection from human harvesting resulted in greater proportions of large individuals within intertidal but not subtidal habitat. Red
abalone shows remarkably low fluctuations in densities and size structure
over a 30 year period. Intense predation by sea otters in the subtidal habitat,
combined with high recruitment rates may underlie these patterns.
MIDDLETON, ELIZABETH M.,1,* JAMES E. MCMURTREY,2 PETYA
E. CAMPBELL,1 LAWRENCE A. CORP1 and LADEAN M. BUTCHER.1
1
NASA/Goddard Space Flight Center, Greenbelt, MD, USA; 2 USDA Hydrology and Remote Sensing Laboratory, Beltsville, MD, USA. Foliar
reflectance and fluorescence responses for corn and soybean plants.
We are investigating the use of spectral indices derived from actively induced fluorescence spectra and passive optical spectra. We examined the
influence of photosynthetic pigment, carbon (C) and nitrogen (N) content
on the spectral fluorescence and passive optical property characteristics of
mature, upper leaves from plants provided different N fertilizer application
rates: 20%, 50%, 100% and 150% of recommended N levels. A suite of
optical, fluorescence, and biophysical measurements were collected on
leaves from field grown corn (Zea mays L.) and soybean plants (Glycine
max L.) grown in pots (greenhouse + ambient sunlight. Steady state laserinduced fluorescence emission spectra (5 nm resolution) were obtained
from adaxial and abaxial surfaces resulting from excitation at single wavelengths (280, 380 or 360, and 532 nm). For emission spectra produced by
each of these excitation wavelengths, ratios of emission peaks were calculated, including the red/far-red chlorophyll fluorescence (ChlF) ratio
(F685/F740) and the far-red/green (F740/F525) ratio. High resolution (,
3 nm) optical spectra (350-2500 nm) of reflectance, transmittance, and absorptance were also acquired for both adaxial and abaxial leaf surfaces.
Species differences were demonstrated for several optical parameters. A
"red edge" derivative ratio determined from transmittance spectra [as the
maximum first derivative, between 650 -750 nm, normalized to the value
at 744 nm, or Dmax/D744], was strongly associated with the C/N ratio (r2
5 0.90, P # 0.001). This ratio, calculated from reflectance spectra, was
inversely related to chlorophyll b content (r2 5 0.91, P # 0.001) as was
the ChlF (F685/F740) ratio obtained with 532 nm excitation (r2 5 0.76, P
# 0.01). Discrimination of N treatment groups was possible with specific
fluorescence band ratios (e.g., F740/F525 obtained with 380 nm excitation).
Higher ChlF and blue-green emissions were measured from the abaxial leaf
surfaces. Abaxial surfaces also produced higher reflectances, in general, in
the 400-800 nm spectrum.
MILLER, AMY E.,1,* JOSHUA P. SCHIMEL,1 JOHN M. MELACK1 and
JAMES O. SICKMAN.2 1 Dept. of Ecology, Evolution, and Marine Biology, University of California, Santa Barbara, CA; 2 California Dept. of
Water Resources, Sacramento, CA. Response of chaparral soils to episodic rewetting: Microbial controls on N losses.
Nitrogen (N) export from chaparral communities of the southern Sierra
Nevada foothills is dominated by episodic NO3- losses during late-season
rain events, accounting for greater than 90% of annual DIN losses. To
evaluate the role of microbial processes in controlling such losses of N,
we examined variation in soil inorganic N pools and microbial biomass in
a chamise-dominated system during the transition from summer drought to
winter rains. Monthly field measurements showed that net nitrification increased over the course of the summer, ultimately exceeding net N min-

236

Abstracts

eralization, while ambient NO3- concentrations remained low and exchangeable NH4+ increased 4-fold. With the first rainfall event, soil NH4+ concentrations decreased 30%, while NO3- increased 30-fold, indicating a period
of intense nitrifier activity following rewetting. Following the first large
storm producing measurable discharge, soil NO3- concentrations decreased
to previous levels and nitrification rates failed to recover, despite concurrent
increases in net N mineralization. These results, paired with results of laboratory incubations, indicate that mild rewetting events can enhance nitrification in this system, and that while nitrifiers may be competitive only
during periods of initial rewetting, they may have a disproportionate impact
on N losses during subsequent storms.
MILLER, STEPHANIE A.,* KELLY O. MALONEY,* RICHARD
MITCHELL* and JACK W. FEMINELLA. Auburn University, Auburn,
AL. Is coarse woody debris a refuge for attached algae in sandy coastal
plains streams?
Shifting sands in small coastal plains streams provide an unstable, abrasive,
and potentially inhospitable habitat for benthic organisms, especially during
spates. Coarse woody debris (CWD) can reduce the impact of physical
disturbance on organisms by reducing sand particle movement and thus
stabilizing the streambed. We compared algal biomass (as chlorophyll a)
and diatom composition on sand versus submerged CWD substrates in 11
small coastal plains streams at Fort Benning, GA. Physicochemical habitat
variables, including CWD abundance and streambed stability also were
quantified. Algal biomass on sand was higher in streams containing high
CWD. Although algal biomass on CWD was highly variable, diatom diversity and richness were higher on CWD than on sand. Average length
of diatom cells also was higher on CWD than sand substrates because of
a greater proportion of larger species occurring on CWD, and diatom cells
on sand in high-CWD streams were larger than cells in sand substrates in
low-CWD streams. Scanning electron microscopy revealed that crevices
and outer surfaces of sand grains contained mostly adnate, short chains of
the diatom Eunotia, whereas a more diverse diatom assemblage occurred
among deep crevices and pits on CWD surfaces. These observations suggest that CWD not only serves as a substrate-stabilizing factor in coastal
plains streams, but also may provide a refuge for large diatom species illadapted to hydrologic disturbance.
MILLER, THOMAS E.* Department of Biological Science, Florida State
University, Tallahassee, Florida, USA. Invertebrates in Sarracenia purpurea: using communities that occur at a continuum of spatial scales
to address questions of scale.
Scale discontinuities due to habitat or resource patchiness are inherent in
virtually all communities. Patterns of diversity and structure studied at different scales should reveal the variety and relative importance of mechanisms that structure communities at each scale. The inquiline communities
associated with the carnivorous pitcher plant, Sarracenia purpurea, have a
long history of use in experimental ecology. At the smallest scale, each
leaf provides a habitat for an aquatic community that is largely structured
by relatively short-term biotic (migration, competition, predation) and abiotic (disturbance, climate) forces. Manipulative experiments demonstrate
that the local dynamics are affected by resource availability (primarily ants
captured by the leaf) and the presence of the top predator (larvae of the
mosquito Wyeomyia smithii). At the among-pitcher scale, communities in
newly opened leaves are migration-limited as they undergo succession.
Migration into established communities also increases diversity through
small mass effects and can change the community structure through species-sorting processes. Community patterns at larger spatial scales will be
determined by processes that occur over longer time scales: biogeographic
patterns formed by long-distance dispersal and establishment, and, potentially, evolution through drift and local adaptation. Data from across N.
America demonstrate that this community type is remarkably consistent
despite significant variation in habitat and climate; most species can be
found across the entire geographic range of S. purpurea. While this community has almost certainly expanded from south to north, bacterial and
protozoan species richness actually increases with latitude, while mosquito
abundance declines. I discuss the relative importance of different forces at
different scales, the interactions among these forces, and the applicability
of these results to other communities.

MILLS, APRIL V.,1,* ANDREW M. BJORN,1 PAUL A. WADDELL2 and

MARINA ALBERTI.1 1 Department of Urban Design and Planning, University of Washington, Box 355740, Seattle, WA, USA; 2 Evans School of
Public Affairs, University of Washington, Box 353055, Seattle, WA, USA.
Valuing proximity to forest patches at the neighborhood scale: A hedonic model using landscape metrics.

MINOCHA, RAKESH,1,* DAVID HOLLINGER,1 STEPHANIE LONG,1

BRYAN DAIL,2 ALISON MAGILL3 and JOHN D. ABER.3 1 USDA Forest
Service, Durham, NH, USA; 2 University of Maine, Orono, ME, USA;
3
University of New Hampshire, Durham, NH, USA. Effects of nitrogen
addition to the canopy at the Howland Ameriflux Site, ME and to the
forest floor at the Harvard Forest LTER Site, MA on the foliar carbon
and nitrogen metabolism and forest productivity.

It has previously been shown that many homebuyers express preferences

for houses located near natural features and in fact are willing to pay a
premium for being located near open spaces, parks, and water features.
This paper asks if homebuyers affinity for natural landscape features at a
neighborhood scale is reflected in the marginal benefit for houses located
in natural settings over those located in more built environments. In this
paper we test the hypothesis that single-family residents in King County,
WA prefer houses closer to large forest patches and further from large
patches of impervious surface. We use a hedonic pricing model that estimates the effect of structural, jurisdictional, locational, and spatial landscape attributes on housing prices. We specifically determine whether the
marginal benefit of spatial landscape attributes significantly effect housing
prices. Spatial attributes are measured at two scales (300m and 600m) using
five landscape metrics: percent area of paved land, percent area of forested
land, mean patch size, contagion, and the Shannon-index. Our preliminary
results show that at the 600m scale, percent area of forested land has a
significant marginal effect on housing prices. These results indicate that
there is economic value in maintaining green space in urban environments
while simultaneously reducing the ecological disturbance of sprawling development. One way to attract people away from sprawling suburban areas
and closer to cities may be to design more natural areas within higher
density urban environments.

Atmospheric nitrogen (N) deposition may cause forest decline or enhance

its productivity depending on the initial N status of the soil, and the rate
and duration of N deposition. Continuous deposition of N can move ecosystems towards nitrogen saturation, a sate that can lead to forest decline.
Previous studies from our group show a strong correlation between soil
nutrient deficiencies and/or chronic N addition and increased foliar free
putrescine (an organic polyamine) levels, indicating that foliar putrescine
can potentially be used as a marker of general stress in visually healthy
trees. Free amino acids, especially arginine, also increased in response to
chronic N input. Arginine could thus be an indicator of excess nitrogen
inputs in otherwise nitrogen limited forest soils. The objective of the present study was to determine changes in foliar physiology (cellular polyamines, amino acids, proteins and chlorophyll levels) at the Howland Experimental Forest following a relatively low level of nitrogen application
to canopy (18 Kg N ha-1 y-1) since 2001 and at the Harvard Forest LTER
Site where a similar experiment underway since 1989 is adding 50 and
150 Kg N ha-1 y-1of N to the forest floor. We predicted that at the Howland
site, the smaller foliar N addition will not exceed the capacity for N uptake
by vegetation and may lead to greater C sequestration. This is unlikely to
put physiological stress on trees. Our preliminary data for first two years
at Howland show no change in the physiology of the needles due to canopy
N application. However, significant changes in foliar physiology were observed after 6 years of chronic N addition at Harvard Forest. The longterm goal is to be able to better understand the N deposition levels that
elicit a stress response in a forest where N availability is assumed to limit
net primary productivity.

MINCHIN, PETER R.* Southern Illinois University, Carbondale, Illinois,

USA. Stepwise ANOSIM: Finding the subset of species that optimally
discriminates groups of sampling units.
ANOSIM is a multivariate, nonparametric test for differences in community
composition among groups of sampling units. It is based on a statistic
computed from the ranks of dissimilarities, with a significance test derived
by random permutation of group membership. Higher values of the statistic
indicate better discrimination among groups. Traditionally the test is performed using all species in the data matrix. To determine the subset of
species that optimally discriminates among groups, a stepwise algorithm
has been developed. Species are removed from the data matrix, one at a
time, and the ANOSIM test recomputed. That species whose removal resulted in the largest improvement in discrimination is then identified and
permanently removed. This process is then repeated on the reduced data
matrix. Typically the ANOSIM statistic increases initially, as nondiscriminatory species are sequentially removed. Eventually the statistic begins to
decline, as it becomes necessary to select from species that all help to
separate the groups. The application of stepwise ANOSIM to research questions in community ecology is illustrated by selected case studies.
MINER, BENJAMIN G. Univerisity of Florida, 223 Bartram Hall, Gainesville, FL, USA. How to test whether phenotypic plasticity is adaptive.
Biologists have become increasingly interested in the evolutionary and ecological role of phenotypic plasticity in natural systems, and many studies
assert that plasticity is adaptive. However, the current theory to test whether
phenotypic plasticity is adaptive can be confounded by a number of factors.
For example, in the typical "performance test" commonly used, performance of phenotypes and costs of experiencing different environments are
confounded yet not discussed. These problems with the current study of
phenotypic plasticity make it difficult to know whether plasticity is indeed
adaptive. Here, I discuss some of the problems with current approaches
and develop a framework to directly test whether plasticity is adaptive. I
also discuss some indirect tests that biologists are using to address this
question.

MINTON, MARK S.* and RICHARD N. MACK. Washington State University, Pullman, Washington, USA. Cultivation counters the long-odds
on plant naturalizations by mitigating environmental stochasticity.
Immigrant plant populations may be especially vulnerable to environmental
stochasticity; they are often small and cannot withstand the repeated reductions in population size that result from random environmental events
in a new range. Cultivation (e.g. irrigation, protection from parasites, predators and grazers) buffers a founder population from a wide array of abiotic
and biotic threats; it could provide a prolonged opportunity for population
growth, dispersal, and eventual persistence in a new range. Using a series
of factorial experiments, we investigated the effects of different regimes of
cultivation on the persistence of the founder populations of four alien species. The cultivation removed vertebrate and insect seed predators and grazers and provided different levels of irrigation. Treatments also examined
the role of size and density of the founder populations in influencing persistence. The fate of founder populations within and across the three years
of experimentation was highly variable and underscored the precarious environment in which plants become established. Irrigation and the exclusion
of vertebrate seed predators and grazers resulted in a highly significant (P
, 0.001) increase in germination and survival. These forms of cultivation
produced a corresponding highly significant (P , 0.001) increase in R0,
although the magnitude of R0 varied significantly among the species. The
size of the founder population significantly increased R0 (P , 0.001) in
most years for all species. Cultivation significantly increased the probability
of naturalization; however, the minimum level of cultivation for R0 $ 1
varied across years and species. These results underscore the importance
of environmental stochasticity in determining the fate of founder populations and the potential of cultivation in countering these long odds against
naturalization.
MIRIK, MUSTAFA,1,* JACK E. NORLAND,2 ROBERT L. CRABTREE,3
MARIO E. BIONDINI2 and GERALD J. MICHELS.1 1 Texas A&M University, College Station, TX; 2 North Dakota State University, Fargo, ND;
3
Yellowstone Research Center, Mozeman, MT. Relationship between fine
resolution imaging data and Big Sagebrush (Artemisia tridentate Nutt.)
phytomass.
We studied the prediction power of fine spatial resolution narrow band
vegetation indices along with simulated multispectral vegetation indices for

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237

estimating biomass in a big sagebrush (Artemisia tridentate Nutt.) region

in Yellowstone National Park. We used the custom-built normalized difference and simple ratio vegetation indices of 1 m2 spatial resolution of
PROBE-1 hyperspectral imagery acquired in August 1999. PROBE-1 imaging system collects information of the earth surface from 423 to 2507
nm region of the spectrum with an average, minimum, and maximum bandwidth of 15, 10.7, and 19.8 nm, respectively. Ground data were collected
in thirteen 1 m2 plots by clipping to ground level all herbaceous vegetation
and stripping green leaves from big sagebrush plants which were then dried
and weighed. A woody vegetation index (WVI) was developed from digital
pictures of the clipped plots. Ground plots were matched to the corresponding image picture elements in order to extract reflectance data and perform
the regression analyses. We found a wide range of coefficients of determination (R2) by regressing biomass components on either custom built
vegetation indices or narrow band reflectance data. The R2 values ranged
from 0.15 to 0.91 for the simple regression models. The multispectral simulation of vegetation indices showed lower R2 values. The Multiple regression models with two variables improved the prediction power and
produced R2 values of 0.48 for WVI, 0.72 for big sagebrush green biomass
only, 0.86 for total biomass, and 0.94 for total biomass excluding big sagebrush green biomass. We concluded that high spatial and spectral resolution
remotely sensed data contain information to predict biomass components
in big sagebrush areas.
MIRITI, MARIA N. The Ohio State University, Columbus, OH. Responses of prairie natives and exotics to addition of soil nitrogen and calcium.
Variation in soil nutrient levels and in plant resource use efficiency has
been linked to community invasibility. Nutrient rich ecosystems tend to be
dominated by plants with high concentrations of leaf nutrients. Plants that
can efficiently convert high levels of one or more resources to growth and
reproduction are capable of invasive population growth. On the other hand,
plants that most efficiently use low levels of the same resource or resources
are often slower growing species whose populations are replaced by invasives as nutrient levels increase. Competitive outcomes between strong
competitors and ruderals in response to experimentally reduced soil nutrients have not been widely studied. Prairie restorations present an opportunity to test differential performance between prairie natives and exotics
since reestablishment of prairie natives to abandoned agricultural areas is
impeded by the greater competitive ability of old field exotics. Addition of
nitrogen and calcium are common agricultural practices because nitrogen
increases plant growth, but acidifies the soil. Calcium is commonly added
to raise pH. Calcium has recently been implicated in promoting success of
invasives, but the interaction between nitrogen and calcium addition is not
generally studied. I measured the growth responses of 7 native and exotic
species including grasses and legumes that commonly occur in prairies and
abandoned agricultural fields in Central Ohio to three nutrient levels in a
greenhouse experiment. Fertilizer treatments consisted of nitrate soda and
calcium nitrate. Nitrogen addition was 50kg/ha. Control plants received no
added nutrients. Height, biomass, and root:shoot investment were measured
and analyzed using ANOVA. Native and exotic species showed differential
responses to treatments with exotics showing a greater response to nitrogen
addition. Responses to nitrogen and calcium addition did not distinguish
between functional groups. Results suggest that reduction of soil nitrogen
may accelerate restoration of prairie natives to abandoned agricultural areas.
MITAL, STEPHEN* and DANIEL UDOVIC. University of Oregon, Eugene, OR. Linking academics, skill development, and community involvement: a service learning model for environmental programs.
Traditional academic courses provide a framework for understanding environmental problems but dont provide students structured opportunities
to respond to the issues they are learning about. While conventional internship programs offer the opportunity to work on real-world problems,
they often fail to provide a standardized set of skills that complement academic curricular goals. The Service Learning Program was created by the
University of Oregon Environmental Studies Program to provide supervised practical learning experiences that support the environmental studies
program curricula. Thoughtfully designed projects provide undergraduate

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students supervised opportunities to apply the knowledge they are gaining

in more traditional courses by acting as consultants to local agencies and
non-profit organizations. Our team-based approach mimics the professional
world. Our year-long project timelines provide students the time needed to
acquire skills - data collection, data analysis, use of specialized software
programs, report writing, oral presentation, and teamwork - that not only
complement their academic interests, but prepare students to enter the
workforce. Students receive project specific training and one-on-one interaction with project managers and community partners. Student teams meet
weekly with the program coordinator to review assignments and discuss
progress. Projects tap students natural motivation to work on real world
issues such as sustainability and invasive species management. Most students devote more energy towards this course than other courses simply
because they believe their work will have a positive impact on the environment. We have found that the direct benefits of our projects to our
community partners have led to substantial community interest and publicity. This, in turn, is helping the Environmental Studies Program build positive relationships with community-based organizations, attract funding to
support the program, and secure student interest in the program. Our poster
presentation will include examples of student projects.
MITCHELL, CHARLES E.* and ALISON G. POWER. Cornell University,
Ithaca, NY. Plant community dynamics, disease, and productivity under global change.
In a field experiment in which biotic community composition was manipulated, nitrogen addition doubled aboveground productivity of grass mixtures except in communities that both included Avena fatua and were inoculated with barley yellow dwarf virus strain PAV (BYDV), where there
was no effect on productivity. In contrast, a simple model predicted that
nitrogen addition would increase productivity of all communities because
$75% of species in all communities were relatively resistant to BYDV
based on monocultures (,10% of individuals infected). Virus incidence
was over twice as high in communities including Avena and inoculated
with BYDV as in the other communities, suggesting that high virus incidence constrained the response of productivity to nitrogen addition. High
virus incidence in these communities resulted in part because the presence
of highly susceptible Avena increased virus incidence in the other species
relative to when they were grown without Avena ("pathogen spillover").
Additionally, pathogen spillover from Avena decreased the abundance of
two of the three more resistant species, indicating apparent competition
mediated by BYDV. Together, these two processes caused communities
including Avena and inoculated with BYDV to more resemble communities
composed entirely of highly susceptible species than expected from monocultures. Finally, nitrogen addition tended to increase virus incidence in the
more resistant species, perhaps further limiting their ability to increase
productivity in response to nitrogen addition. Virus incidence in the highly
infected experimental communities including Avena and inoculated with
BYDV was the same as the mean incidence in local wild grass populations
(c. 40%). These results suggest additional assumptions to be built into
models of the joint effects of nutrient addition and trophic structure on
primary productivity. Moreover, the effects of atmospheric nitrogen deposition, a major component of global change, on grassland ecosystems may
depend on plant pathogens.
MITCHELL, KATHERINE A.,1 CURTIS MONGER,2 JEFFREY HERRICK1 and DEBRA P. PETERS.1 1 Jornada Experimental Range USDA/
ARS, Las Cruces, NM; 2 New Mexico State University, Las Cruces, NM.
Simulation modelling of soil inorganic carbon in arid environments.
Pedogenic carbonate, or soil inorganic carbon, is an important constituent
of subhumid to arid soils throughout the world. Although terrestrial carbon
cycling science has placed more emphasis on organic carbon, arid and
semi-arid soils store approximately 10 times more inorganic carbon than
organic. It has been estimated that more than 8 x 1014 kg of C are sequestered globally in calcic and petrocalcic horizons of Aridisols and Mollisols.
Our goal was to examine and test hypotheses regarding the relationship of
calcium carbonate (CaCO3) in arid lands to atmospheric CO2 sequestration.
The magnitude of soil inorganic carbon flux with the atmosphere is difficult
to estimate at local, regional or global scales. We developed a conceptual
model and an associated process-based model to simulate the rate of for-

mation and vertical distribution of CaCO3 in soil horizons in the arid southwestern U.S. The model depicts short term processes of CaCO3 precipitation and dissolution that occur in periods of days to centuries to millennia.
We use the model to illustrate the environmental and biotic conditions that
control the CaCO3 source-to-sink relation. Initial findings show that pedogenic carbonate can serve not only as a sink, but also as a source for CO2.
Our simulation modelling framework helps illustrate the historic and future
conditions under which soil carbonates alternate between storing and releasing CO2 to the atmosphere.
MITCHELL, RANDALL J.,1 JEFFREY D. KARRON,2 KARSTEN G. HOLMQUIST2 and JOHN M. BELL.2 1 University of Akron, Akron, OH,
USA; 2 University of Wisconsin-Milwaukee, Milwaukee, WI, USA. Patterns of multiple paternity in fruits of Mimulus ringens.
Multiple paternity within fruits arises when pollen from more than one
donor successfully fertilizes ovules in a single fruit. This can occur in two
different ways. 1) A single probe by a pollinator may deposit a mixture of
pollen from several donors. 2) A sequence of several separate probes may
deposit pollen from different donors. These alternatives have distinct consequences for pollen competition and mate choice. However, there is as yet
little evidence on the relative importance of these two mechanisms, and
few studies on the overall rate of multiple paternity. We report on patterns
of multiple paternity in Mimulus ringens, and use observations of pollinator
behavior to infer the mechanisms generating multiply sired fruits. Multiple
paternity was very common for M. ringens, with over 95% of 204 fruits
having two or more outcross pollen donors in progeny arrays of ten genotyped seeds. The number of sires per fruit averaged 4.63 + 0.10 (mean
+ SE), including selfs. From our detailed observations of pollinator behavior, and the limited pollen carryover for this species, we infer that the
observed levels of multiple paternity could not have been solely the result
of single probes that delivered mixed pollen loads. Multiple probes to flowers, each delivering pollen from one to three different sires is more likely
to have caused the observed patterns, This suggests that both mixed pollen
loads and sequential visitation contribute to multiple paternity in this species.
MITCHELL, RICHARD M.,1,* KELLY O. MALONEY,1 STEPHANIE A.
MILLER,1 JACK W. FEMINELLA1 and PATRICK J. MULHOLLAND.2
1
Department of Biological Sciences, 331 Funchess Hall, Auburn University, Auburn, AL, United States; 2 Environmental Sciences Division, Oak
Ridge National Laboratory, Oak Ridge, TN, 37831. Relationships between coarse woody debris, particulate organic matter, and crayfish in
southeastern coastal plain streams.
The purpose of our study was to assess the role of instream coarse woody
debris (.2 cm, CWD) and coarse/fine particulate organic matter (,2 cm,
POM) on density, biomass, and size frequency of crayfish (Cambaridae)
within small sandy-bottom streams of the Southeastern Coastal Plain. CWD
and sand are the predominant substrates in these low-gradient streams, but
only limited work has been done to assess the degree to which CWD and/
or POM influence crayfish population dynamics, despite the high bioenergetic importance of crayfish. Three streams containing varying amounts
of CWD (4 -10% of stream bottom surface) were sampled for crayfish,
POM, and other physicochemical parameters at the Fort Benning Military
Reservation, Georgia. Crayfish (kick seine) and POM (substrate cores) were
sampled monthly and seasonally, respectively, at 5 sites per stream. Surprisingly, abundance of submerged CWD alone did not predict crayfish
density, but rather density was a combined function of CWD and POM
levels, with highest densities (up to 11 crayfish/m2) occurring in sites with
highest CWD and POM. Our preliminary findings suggest that CWD may
only indirectly increase crayfish density and biomass by influencing retention of POM, a potential food resource, within the streambed. In addition
to quantifying the direct and/or indirect importance of CWD on crayfish
populations, we also are studying the influence of CWD structure (debris
dams vs snags) and the nature of instream wood (debris vs live rootwads)
on POM retention and crayfish size, diet, growth, and production in these
streams.

MOELLER, DAVID A. and MONICA A. GEBER. Cornell University,

Ithaca, NY, USA. Coexistence among plants that share pollinators: Facilitation by conspecifics and heterospecifics in Clarkia.
Understanding the mechanisms of species coexistence has been a challenging problem for ecologically similar species that share resources. While
the underlying assumption for most models of coexistence is that competition for limiting resources is the dominant force regulating diversity, a
growing number of studies on plant communities have found evidence that
direct plant-plant interactions may vary from competitive to facilitative.
Relatively little is known of indirect positive interactions among plant species, such as through the attraction or maintenance of shared pollinators.
Although it is clear that plants can compete for (or through) pollinators,
co-flowering of plants that share pollinators is surprisingly common. We
examined the nature of reproductive interactions among Clarkia species
pollinated by a group of pollen specialist bees. For intensive studies, we
focused on the effect of community, population, and local factors on the
reproductive ecology of a target species, Clarkia xantiana. We found that
congeners frequently coexist with C. xantiana across its geographic range
and that the frequency of coexistence declines sharply at the species range
margin. A comparison of observed patterns of flowering phenology to a
null model revealed that communities range from aggregated to segregated.
The effect of congeners on pollinator abundance and pollen limitation of
reproduction in C. xantiana was positive to neutral where pollen limitation
declined with increasing numbers of coexisting Clarkia species. Further,
we found strong evidence of Allee effects suggesting that both the abundance of conspecifics and heterospecifics positively affected reproductive
success through increased mate and pollinator availability, respectively. Detailed surveys of bee visitor communities confirmed that the availability of
specialists was much greater to C. xantiana in communities with multiple
Clarkia species. Both the patterns and processes examined in this study
indicate that positive interactions between conspecifics and heterospecifics
may influence the population dynamics and distribution of Clarkia xantiana.
MOFFETT, J. MINCY,1,* ROBERT S. BOYD2 and MICHAEL C. WOOTEN.3 1 Auburn University, moffejm@auburn.edu, Auburn, AL; 2 Auburn
University, Auburn, AL; 3 Auburn University, Auburn, AL. Post-hurricane dune restoration techniques on vegetation recovery at Bon Secour
National Wildlife Refuge, Alabama.
Coastal beach and dune systems of the U.S. Gulf Coast are frequently
impacted by tropical storms and hurricanes. We studied the effects of sand
fencing and annual fertilization on beach/dune vegetation at Bon Secour
National Wildlife Refuge, a barrier peninsula system near Gulf Shores, AL,
for five years (1998-2002). Pairs of sampling grids were established at three
locations in the Refuge: two in an area of mature relatively stable tertiary
dunes (Perdue Unit) and one in a more dynamic and exposed area of secondary dunes and swales (Ft. Morgan Unit). One member of each pair was
sand fenced, with each sand-fenced grid subdivided into three fertilized
and three unfertilized areas. Fertilized areas were treated with 13-13-13
fertilizer (rates ranging from 140-260 Kg/Ha/yr) applied in early April of
each year. Sand fence broken-chevrons were installed at the seaward edge
of the primary dune. Total plant cover, species richness, and diversity
(Shannon-Wiener Index) were determined each year and analyzed for treatment effects. Fertilizing significantly increased total plant cover and species
richness, but not species diversity, over time in the Perdue Unit. Fertilizing
had no significant effect on any variable in the Ft. Morgan Unit. Sand
fencing in the Perdue Unit significantly increased plant cover and diversity,
but species richness was unaffected. These effects were localized in a narrow zone bordering the sand fence. In the Ft. Morgan Unit, sand-fencing
did not significantly affect any variable. Our results show these restoration
treatments can significantly accelerate post-disturbance vegetation recovery, but that site-specific factors can influence the overall success of these
treatments.
MOHAN, JACQUELINE E.1,* and WILLIAM H. SCHLESINGER.2 1 Harvard University, Cambridge, MA, jmohan@oeb.harvard.edu; 2 Duke University, Durham, NC. Global change and southeastern ecosystems.
The southeastern United States is replete with rich biological diversity in
ecosystems ranging from bald cypress bayous and fire-dependent longleaf

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239

pine savannas, to high elevation spruce-fir forests. Yet global change will
likely impact these systems in numerous ways. Temperatures are predicted
to increase by 1 to 2 oC in the southeast, which may enhance productivity
and lead to expansions of some forest types, but might also trigger the
reduction or elimination of high-elevation Appalachian ecosystems. At the
same time, rises in sea level in response to global warming will impact
rare systems such as barrier island maritime forests and the Florida everglades. Changes in precipitation are less certain, with the Hadley model
predicting a 3% increase for the southeast by year 2030, and the Canadian
model projecting a 20% decrease. Increased occurrence and severity of
drought would affect the many southeastern wetlands, and has been proposed to heighten regional fire frequency thus prompting vegetation conversions from forests to savannas. Negative consequences of NO3 and SO4
deposition in the southern Appalachians have received wide attention and
include declining forest health, nitrogen saturation, and stream acidification.
Similarly, nutrient runoff from intensified agriculture and development
greatly alters rare, low-nutrient southern Appalachian bog systems. Finally,
increased occurrences of exotic and invasive species, such as kudzu, Japanese honeysuckle, hemlock woolly adelgid, and native but overly-abundant white-tailed deer are already affecting ecosystem functioning and regeneration dynamics. Elevated CO2 in Duke Forest alters productivity of
canopy trees and successional trajectories of the understory. When browsed
by deer, this forest exhibits no understory growth and a paucity of tree
recruits even in the presence of high carbon dioxide. The many factors of
global change that are expected to influence the future functioning of southeastern ecosystems require further study and political action to effectively
propose management and mitigation measures.
MOHAN, SANDHYA* and JONATHAN R. CUMMING. West Virginia
University, Morgantown, West Virginia. Analysis of the urban forest
structure of Morgantown, WV.
Urban forests, defined as assemblages of trees and associated vegetation in
an urban setting, have structural and functional components that provide
distinct ecosystem benefits to the city residents such as storm water retention and pollution abatement. Information about urban forest structure of a
city such as species composition, tree density and tree health is a prerequisite for planning, designing and managing city vegetation on a local and
regional scale. Small cities such as Morgantown, WV, (population
,50000), tend to be under-represented in urban forest assessments but
constitute approximately 70% of the U.S. cities and therefore need to be
studied. Utilizing WV Digital Orthophoto Quarter Quads data for Morgantown, land-use classes (high and low density residential areas, commercial
area, institutional area, forest, farm, and urban vegetation corridor) were
determined, 120 plots (0.4ha) randomly located within these classes and
assessed for percent tree cover and ground cover type. Trees were measured
for DBH, height, and crown dieback. Tree species diversity, abundance,
number of trees and basal area per hectare were calculated for each land
use class. The most abundant tree species in Morgantown are Acer saccharum, Prunus serotina, Robinia pseudoacacia and Acer rubrum which
reflect the composition of the native forests of the area. Residential areas
had the highest species diversity, though ornamental species such as Viburnum sp and Pyrus sp were most abundant. Forest and urban vegetation
corridor classes had the highest values for tree cover, basal area and number
of trees per hectare. Invasive and pioneer species such as Ailanthus altissima, Lonicera sp and R. pseudoacacia were more abundant in commercial
and in institutional areas. Such urban forest structure information is important for small cities like Morgantown to aid in invasive species control,
tree planting decisions, monitoring vulnerable species for signs of decline,
and other management purposes.
MOKOS, JENNIFER T.* and CYNTHIA FOWX. Beczak Environmental
Education Center, Yonkers, NY. Adaptation of a wetland rapid assessment method to teach 3rd grade students the functions and values of
wetlands.
We describe an experiential learning curriculum designed to help grammar
school students gain an understanding of the functions and values of wetlands. We utilized a wetland rapid assessment method, the Army Corps of
Engineers Wetland Functions and Values: A Descriptive Approach, as a
framework to discuss wetland functions and the processes behind how a

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Abstracts

wetland provides those functions. The method was modified to increase its
suitability for 3rd grade students. Eight wetland functions are evaluated in
the adapted method: Wildlife Habitat, Fish and Shellfish Habitat, Food
Chain Support, Floodflow Alteration, Shoreline Stabilization, Sediment/
Toxicant Retention, Nutrient Removal/Transformation, and Groundwater
Recharge/Discharge. The students used the method to evaluate a highly
disturbed tidal marsh along the Hudson River, NY. After the fieldtrip, the
students presented the data they collected for each wetland function. Handson classroom activities designed for each of the wetland functions demonstrate the key processes involved in each function and help the students
understand the data they collected on the field trip. The activities also serve
to reinforce or introduce concepts and skills from the science curriculum
standards for their grade-level. The results of the wetland assessment are
recorded on a chart in their classroom. The students will apply the method
to a less-disturbed tidal marsh on the Hudson River on a second field trip,
which will allow them to comparatively study the two tidal marshes and
to observe how a natural wetland functions. The project will culminate
with the students assisting with the restoration of the disturbed tidal marsh
that they evaluated at the beginning of the year.
MOLL, JASON D.* and JOEL S. BROWN. University of Illinois at Chicago, Department of Biological Sciences, Chicago, IL. Evolutionary coexistence in multiple life history stages.
We model the ways that ontogenetic niche shifts and complex life cycles
might affect the number of species that can coexist at an evolutionarily
stable strategy (ESS), using the G-function approach. This approach consists of both an inner game in which the players get payoffs from their
choice of strategy (similar to traditional evolutionary game theory), and an
outer game, where the frequencies of strategies change as a result of those
payoffs. We expand on previous single life history stage Lotka-Volterra
models, where the strategy affects the carrying capacity and the competition coefficients between the different strategies. Depending on the form
of those relationships and the choice of parameters, the ESS can consist of
either one species or coalitions of multiple species in the single-stage model. In the multi-stage model, there are three different ways that the life
stages strategies could be related. They could be unconnected, positively
correlated, or negatively correlated. That is, the trait could have the same
effect in both stages or there could be a tradeoff between the stages. Unrelated strategies and positively related strategies result in outcomes that
are similar to the single life history stage model, with ESSs of both single
and multiple strategies, depending on the choice of parameters. Negatively
related strategies can result in ESSs in which there are some strategies that
are only found in one life history stage or the other.
MOMEN, B.,1,* P. D. ANDERSON,2 J. H. SULLIVAN1 and J. H. HELMS.3
University of Maryland, College Park, MD; 2 USDA-Forest Service, Corvallis, OR; 3 University of California, Berkeley, CA. Using multivariate
statistics to predict mature tree performance from seedling characteristics.

A common objective in commercial reforestation is to plant seedlings that

will produce superior adult trees. Long-term management planning horizons and life spans of most forest trees complicate the prediction of growth
and performance of adult trees originating from planted seedlings of unknown genetic origin. We used multivariate statistics to identify and rank
a set of seedlings characteristics that might predict the performance of
mature trees in Pinus ponderosa. Results indicated that metabolic heat rate
(q), a measure of total metabolism, of one-yr-old foliage during the peak
growth in May, was the most important seedling characteristic that predicted mature-tree performance. Increased metabolic heat rate in seedlings
corresponded with greater vigor of mature trees. Additionally, seedling basal stem diameter (D), height (H), and needle length (NL), measured in
November, were, in order of importance, other variables that defined the
vigor class of the mature clones. However, these seedling morphological
characteristics correlated negatively with vigor classification of the mature
clones, contradicting the notion that greater D and/or H during the seedling
stage may indicate a greater vigor at maturity.

MONACO, THOMAS A. USDA-ARS, Utah State University, Logan, Utah,

84322. Physiological ecology of Isatis tinctoria suggests mechanisms of
invasiveness.
Isatis tinctoria is a biennial herb currently invading numerous semi-arid
plant communities in the Intermountain West and is listed as a noxious
weed in numerous western U.S. states. It proliferates in disturbed patches
and in dense native plant communities. I evaluated how below and aboveground resources control photosynthesis parameters and growth of 60 day
old I. tinctoria plants in a greenhouse experiment. Plants were grown under
low and high levels of light (250 and 500 mmol m-2s-1), soil moisture (50
and 100 ml day-1), and soil N (5 and 50 mg N kg-1 soil) in 8 L pots. Carbon
assimilation and shoot dry mass were significantly lower in low soil water
than in high soil water. The ability to dissipate light stress (FPSII and Fv/
Fm) was greater (P , 0.001) under high soil N than low soil N. An
interaction (P , 0.001) between light and water was observed for root dry
mass and specific leaf area. For example, specific leaf area increased with
soil water only at low light, and root dry mass increased with soil water
only under high light. These results suggest that disturbances that make
soil N more available may increase the ability of I. tinctoria to overcome
excess light stress in open, dry patches. High light, characteristic of open
patches also appears to favor root growth, but only if soil moisture is
available. However, it is unclear how this herb can also slowly invade intact
dense native vegetation. Perhaps a closer look at how increases in specific
leaf area in dense canopies provide a competitive advantage when water is
not limiting, will reveal additional mechanisms associated with I. tinctoria
movement into dense native vegetation.
MOODY, AARON1,* and ROSS K. MEENTEMEYER.2 1 University of
North Carolina at Chapel Hill, Chapel Hill, NC; 2 Sonoma State University,
Rohnert Park, CA. Life history, xylem function, and drought stress interact to control spatial distributions of chaparral shrubs.
Spatial patterns of shrub life-history categories, and in the distribution of
five Ceanothus species were examined in relation to topographically-mediated differences in drought severity within three watersheds on the coastal
and inland flank of the Santa Ynez Mountains, California. Spatially distributed fields of soil water tension were simulated for the studied watersheds using high-resolution digital terrain data and daily climate data in
combination with a process-based hydro-ecological model. Field samples
of species composition were spatially integrated with the distributed
drought data for analysis of ecological relationships. Species that depend
on fire for recruitment were increasingly represented with increasing soil
water tension. Sites that experience moderate drought severity permitted
co-dominance of species from both recruitment modes (obligate seeders
and facultative sprouters). The distribution of five Ceanothus shrub species
strongly tracked differences in soil-water tension in accordance with their
resistance to water stress-induced xylem dysfunction.
MOODY-WEIS, JENNIFER M.* and HELEN M. ALEXANDER. University of Kansas, Lawrence, Kansas. Role of disturbance and seed density
on seedbank formation and population dynamics in Helianthus annuus.
The role of seedbanks in determining plant population abundance, structure,
and dynamics is beginning to be realized. However, we know little about
what factors influence seedbank formation. In annual, weedy species, it is
clear that soil disturbance plays a major role in determining the fate of
seeds in the soil. Helianthus annuus is a weedy species that is native to
the Great Plains, is commonly found in disturbed habitat, and forms persistent seedbanks. The goal of this study is to determine how different
disturbance regimes and seed densities influence the formation of seedbanks and, in turn the role of the seedbank in the population dynamics of
H. annuus. Specifically I am addressing the following questions:1.) How
does the timing of disturbance relative to seed dispersal influence the formation of a persistent seedbank? 2.) Does the density of seeds dispersed
influence seed bank formation? 3.) Are populations limited by the presence
of safe-sites (i.e. disturbance) or by the availability of seeds (i.e. seed density)? In 2001, I set up 210 experimental populations, representing 21 treatments of different combinations of seed density, pre-dispersal disturbance,
and post-dispersal disturbance. Data were collected on the numbers of seedlings that emerged, the number of plants surviving to reproduction and the

reproductive output per population. Soil samples from each population

were collected and sieved to estimate the number of seeds remaining in
the soil. In 2002, more seedlings germinated in plots that had a large disturbance compared to those with no disturbance or a small disturbance.
Also, higher seed density lead to higher seedling emergence. In most cases,
the differences in seedling density appear to carry over into number of
flowering plants and the number of flowers produced. In fall of 2002, all
plots were disturbed, which should create ideal germination conditions.
Data will again be collected in 2003. Germination and seed sieving results
from both 2002 and 2003 will allow me to evaluate the mechanisms that
lead to variation in seedbank formation. This study is important as it experimentally examines the role of disturbance in seed bank dynamics.
MOONEY, HAROLD A.* Stanford University, Stanford, CA. The Multiscale approach and the Millennium Ecosystem Assessment.
The Millennium Ecosystem Assessment (MA), modeled after the IPCC
(Intergovernmental Panel on Climate Change), is a response to governments, the private sector, non-governmental organizations and scientists for
an integrated assessment of the consequences of ecosystem change for human well-being. The MA focuses on ecosystem goods and services, the
consequences of changes in the delivery of these ecosystem services for
human well being, as well as the significance of changes in ecosystems for
other life on earth. It is an analysis of the options available that can enhance
the sustainability of ecosystems in providing for human needs. This multiscale assessment consists of inter-linked assessments undertaken at local,
regional and global scales. Sub-global assessments will directly meet the
needs of decision-makers at the scale at which they are undertaken,
strengthen the global findings with on the ground reality, and augment the
local findings with global perspectives, data and models. The MA will
strengthen the capability of regional, national and local institutions to bring
the best science to bear on decisions concerning ecosystems and their impacts on people. Scenarios are being developed that will indicate the potential trajectories of ecosystem service provision and sustainability under
differing global developmental regimes. The final results of this international effort, involving hundreds of scientists, will be released in 2005. The
main focus of this symposium is to present the conceptual underpinnings
of the Millennium Ecosystem Assessment.
MOONEY, KAILEN A.* University of Colorado, Boulder, Deptarement of
EPO Biology, Boulder, CO, USA. Symmetry and strength of bird-ant
interactions in ponderosa pine canopies.
In ponderosa pine (Pinus ponderosa) forests of the Colorado Front Range,
canopy foraging birds caused a sixty-fold reduction in the abundance of
the aphid Cinara schwartzii when they were tended by the ant Formica
podzolica, but had no measurable effect on aphids in the absence of ants.
To better understand this result, I conducted two experiments to characterize the strength and symmetry of bird-ant interactions in this system. In
the summer of 1999 I observed the foraging patterns of birds on sixteen
pairs of control and ant-excluded trees. In the summer of 2002 I observed
the foraging patterns of ants on ten pairs of control and bird-excluded
branches equipped with ant feeders that provided a consistent food resource. There was not a significant difference in mean bird foraging time
between control and ant-exclusion trees, indicating that ants did not deter
foraging birds. However, birds reduced ant abundance by 25%. I propose
that the asymmetry of this bird-ant interaction may explain, in part, the
dramatically strong negative effects of birds on ant-tended aphids; ants can
not deter these predatory birds, and birds negatively effect aphids not only
directly through predation, but also indirectly by reducing the abundance
of mutualist ants.
MOORE, DAVID J.,1,* SUSANNE AREF,1 ADRIEN FINZI,2 JASON
HAMILTON,3 JEFFREY PIPPEN,4 MOON-HO HO1 and EVAN H. DELUCIA.1 1 University of Illinois - Urbana-Champaign, Urbana, IL; 2 Boston
University, Boston, MA; 3 Ithaca College, Ithaca, NY; 4 Duke University,
Durham, NC. Inter-annual variation in the response of tree growth and
productivity to exposure to elevated [CO2].
Elevated atmospheric [CO2] enhances growth in plants through an increase
in photosynthetic efficiency. Our ability to predict the capacity of future

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241

forests to sequester atmospheric carbon depends on understanding how

variations in climate will modulate that response at the tree level and forest
level. In the longest running experiment of its kind, an intact Pinus taeda
forest was exposed to +200 p.p.m. CO2 for 6 years using Free Air Carbondioxide Enrichment (FACE). We used a non-linear growth model to determine the initiation and cessation of diameter growth. Stimulation of diameter growth rate, expressed in relative basal area increment (RBAI) was
sustained over time (ranging from 17.2- 26.2%) and correlated well with
seasonal temperature totals. These results agree with previous modeling
and empirical studies of photosynthetic response. There was a sustained
increase of productivity caused by the elevated [CO2] treatment (193
gm22y21 or 25%). There was no interactive effect of elevated [CO2] and
seasonal rainfall; the productivity of both ambient and elevated [CO2] treatments showed a strong parallel reduction in low precipitation years. Predicted rises in [CO2] and temperature will likely stimulate forest growth
but water availability will remain a key growth limiting factor.
MOORE, GEORGIANNE W., BARBARA J. BOND and JULIA A.
JONES. Oregon State University, Corvallis, OR. Does diversity of overstory trees affect transpiration?
One of the greatest differences between natural and managed forests lies
in overstory species diversity. While much attention is paid to plant species
diversity in terms of ecosystem resilience or animal habitat, less attention
has been paid to impacts of species composition and diversity on fluxes of
matter and energy. In this study we asked: Do mixed-species forests transpire more or less water than the average of similar-density monocultures
of the same species? The answer to this question is vital to understanding
how co-occurring species interact in an ecosystem and to gain insights into
how diverse forest types impact the hydrologic cycle. This study took place
at the H.J. Andrews Forest in the Western Cascades of Oregon. Transpiration was measured using constant heat sap flow sensors in 15-year-old
red alder and Douglas-fir planted at 1100 trees per hectare as either monocultures or admixed 50:50 plantings in 20 x 20 meter plots. We hypothesized that total annual transpiration is greater in mixed-species plots than
the average of single-species plots because of altered timing of peak transpiration, less sensitivity to stresses such as high vapor pressure deficit and
low soil moisture, and better ability to utilize soil moisture. Further, because
red alder is a nitrogen fixing plant, we used nitrogen isotopes to evaluate
whether Douglas-fir received nutrient benefits from co-occurring red alder,
which would provide an additional explanation for increased transpiration
in mixed-species plots.
MOORE, JEFFREY E.* and ROBERT K. SWIHART. Department of Forestry and Natural Resources, Purdue University, West Lafayette, IN, USA.
Consequences of forest fragmentation for regeneration of animal-dispersed hardwood tree species in the Midwestern United States.
Hardwood tree species (e.g., oaks Quercus, black walnut Juglans nigra,
hickories Carya) in the Midwestern U.S. depend on granivorous rodents
and birds, principally tree squirrels (Sciurus spp.) and blue jays (Cristata
cyanocitta), to disperse their seeds for regeneration. However, forest fragmentation predictably alters the community composition of granivores
within isolated habitat patches, increasing the biomass of species that function primarily as seed predators (e.g., white-footed mice Peromyscus leucopus, eastern chipmunks Tamias striatus) and reducing the biomass of
species that also function as seed dispersers. We conducted a multi-factorial
experiment to investigate how forest fragmentation, by directly altering the
communities of important seed dispersers, may indirectly affect hardwoodtree dispersal and regeneration. We established 12 study sites in forest
patches differing in granivore composition. Within each site, we placed
metal-tagged acorns from three oak species (Q. alba, Q. rubra, Q. palustris), shagbark hickory nuts (C. ovata), and black walnuts in four types of
semi-permeable exclosures that allowed access to different groups of granivore species (1 5 access to all granivores, 2 5 blue jays excluded, 3 5
blue jays and Sciurus excluded, 4 5 accessible only to Peromyscus). We
used metal detectors to locate eaten and cached seeds following removal
by granivores. We recorded rates of seed removal from exclosures, proportion of seeds consumed vs. cached after the fall caching period, and the
dispersal distance of cached seeds. We analyzed these data with respect to
granivore composition of each patch, and to the exclosure treatment from

242

Abstracts

which seeds originated. Results and implications of fragmentation for granivore-mediated forest succession are discussed.
MOORE, JOHN C.,1,* KATHY COCRHAN,1 MARISA CHELIUS1 and
JRENE RHAM.2 1 Math and Science Teaching Institute, University of
Northern Colorado, Greeley, CO, USA; 2 Department of Educational Psychology, Univeristy of Montreal, Montreal, Quebec, CANADA. Models
of research mentoring: How do K-16 students, teachers, and researchers perceive ecological research?
Collaborations require that all participants have a clear understanding of
the project objectives and that they speak with a common language. Our
work to connect K-16 students and teachers to the ecological research community revealed gaps in both, in that what students and teacher view as
authentic science differs from the perspectives of the researchers. We make
the case for an emergent notion of authenticity of science based on systems
theory and neo-Piagetian thought. Because of the contextual nature of
learning, the influence of the learning environment, and the roles that different experiences have on learning, the science of a scientist is different
from the science of a student and that of a teacher. Duplicating real-life
science experiences in the classroom may be more myth than fact, as it
creates a unique form of science and authenticity. As such, what constitutes
authentic science can neither be taught in the traditional didactic modes
nor through simulations of scientists science in the classroom, but requires
an immersion through mentoring. We draw from two different face-to-face,
teacher/student-scientist partnership programs to illustrate the notion of authenticity that emerges as teachers, students and scientists come to interact.
MOORE, JONATHAN W.,* DANIEL E. SCHINDLER and MARK D.
SCHEUERELL. Univeristy of Washington, Seattle, Washington. Ecosystem consequences of disturbance by spawning salmon in Alaska.
Pacific salmon return to their natal freshwaters to spawn and die, acting as
vectors of marine-derived nutrients and energy to coastal ecosystems.
While much research has focused on the importance of marine-derived
nutrients to coastal food webs, relatively little attention has been devoted
to the impact of the disturbance that salmon create when digging their redds
(nests). Spawning sockeye salmon (Oncorhynchus nerka) dig redds that
cover over 2 m2 and are 20 cm deep, on average. In areas with healthy
populations of salmon, sockeye often spawn at extremely high densities,
with networks of redds covering entire spawning reaches. Thus, we hypothesized that this bioturbation is a critical source of disturbance to benthic habitats. We investigated the impacts of disturbance caused by nestdigging salmon in two streams and two lakes in Alaska by experimentally
excluding salmon from areas where they traditionally spawn. We sampled
exclusions and control plots before, during, and after the peak spawning
time. Salmon displaced silt, some of which subsequently accumulated in
areas without spawning. However, this silt displacement is not only a local
impact. The particulate load of water flowing out of creeks with spawning
salmon indicated that disturbance by salmon leads to the system-wide export of enormous quantities of silt. In addition, salmon spawning locally
decreased algal biomass by 5x as compared to exclusion plots. Spawning
salmon also impacted the benthic invertebrate assemblage. Therefore, habitat modification by spawning salmon can be a substantial disturbance to
coastal freshwater ecosystems. Removing populations of strong interactors
like salmon, or other ecosystem engineers, modifies disturbance regimes
and could lead to unforeseen ecosystem consequences.
MOORHEAD, DARYL L. and ROBERT SINSABAUGH. Modeling impacts of N- and polyphenolic inhibition on litter decay.
A semi-mechanistic model of litter decay was used to explore the negative
impacts of polyphenolic compounds on turnover of labile compounds and
high N concentrations on polyphenolic decay. Microbial N:C was varied
with relative availability of both elements. When turnover of labile compounds was inhibited by polyphenolic compounds, decay of polysaccharides was stimulated but polyphenolic turnover slowed; these changes had
small impacts on overall decay rate. However, negative effects of N on
polyphenolic turnover reduced overall litter decay so severely that a reduction in N was necessary to achieve reasonable rates. Simulated leaching
stimulated decay rates, but produced variable patterns of mass loss sensitive

to patterns of precipitation. Altering N:C ratios of microbiota according to

the availability of C and N generated a rapid replacement of a high N:C
decomposer community in early decay, with a low N:C group during intermediate decay, followed by an increase in N:C in the latter stages of
decay; consistent with shifts between bacteria and fungi-dominated communities. In all, these modeling experiments produced patterns in decomposition consistent with qualitative reports from literature, although few
studies provide sufficient data for rigorous quantitative validation.
MORALES, MANUEL A.* and ANGUS L. BEAL. Williams College, Williamstown, MA. The effects of host-plant quality in an ant-membracid
mutualism.
A relatively unexplored question in ecology is the degree to which the
population dynamics of mutualism is mediated through bottom-up regulation, including host-plant quality. We report results from an experiment that
tested the consequences of host-plant quality (manipulated with NPK fertilization) for the mutualism between Publilia concava treehoppers (Hemiptera: Membracidae) and ants in the genus Myrmica and Formica. Publilia concava is a phloem feeding insect found on the host-plant Solidago
altissima that excretes large quantities of honeydew. Ants collect this honeydew and in exchange, protect these treehoppers from predators and may
improve feeding efficiency. During the summer of 2002, we established a
two-treatment experiment in Hopkins Memorial Forest, Williamstown MA,
that manipulated host-plant quality and ant presence in a factorial design.
Predators were excluded from all treatments. We censused plants for anttending level and treehopper population size throughout the summer. Results confirm that NPK fertilizer had a significant effect on host plant quality: fertilization significantly increased leaf surface area and nitrogen content. Fertilization and ant tending significantly increased treehopper survivorship, but there was no significant interaction between these factors.
Similarly, fertilization had a strong positive effect on the weight of falladult treehoppers and ant tending marginally increased treehopper weight,
but with no significant interaction between these factors. Although both
host-plant quality and ant tending increase the performance of Publilia
treehoppers, our results do not support the hypothesis of strong bottom-up
mediation in this mutualism.
MORAN, MATTHEW D.* Hendrix College, Conway, AR. Effect of bison
grazing on arthropod abundance and diversity in a tallgrass prairie.
I investigated the effect of grazing by American Bison (Bos bison) on an
arthropod community during an entire growing season in tallgrass prairie
habitat. Bison grazing increased the overall abundance of arthropods, an
increase that was present in both herbivorous and carnivorous arthropods,
but not in detritivores. The increase in herbivores resulted in an arthropod
herbivore load almost three times higher in bison-grazed plots. Among
herbivores, the phloem-feeding guild was dramatically more abundant
while chewing herbivores were not affected by grazing. Among carnivores,
both generalist and specialists were more abundant in grazed plots. When
comparing insect abundance on individual C4 grasses and C3 forbs, I found
that the grazing affect appeared stronger than the effect of plant type. Herbivorous and carnivorous arthropod richness was higher in grazed plots,
although the response was strongest among herbivores. In general, the
greatest difference in abundance and diversity between grazed and ungrazed plots was present during the early samples, while late season samples
tended to converge between treatments. Published research indicates that
ungulate grazing, while decreasing available plant material to other consumers, enhances plant quality by increasing nutrient cycling. The results
of this study suggest that this higher plant quality outweighs the potential
negative competitive effect of plant biomass removal by grazing. Further,
the response of arthropods on individual plants supports the hypothesis that
these changes plant quality are more important than the changes in plant
community structure (increase in forbs), at least in terms of overall arthropod abundance and diversity patterns. Since arthropods are among the most
diverse and abundant organisms in North American grasslands, and since
they are a food source for many other consumers, bison may represent
valuable management tools for maintaining biodiversity in this system.

MORENO-CASASOLA, PATRICIA. Institute of Ecology A.C., Xalapa,

Veracruz, Mexico. Coastal development in rural regions: mechanisms
for planning within the scenarios of sea level rise.
The effect of sea level rise on coastal dunes should be approached within
the general trend of beach and dune erosion. Management of many coastal
areas in the tropics are dealing with more everyday problems and there is
little concern with probable scenarios 20 years from today. Planning for
development and for education in coastal areas should take into account
these trends. There is also a general lack of data, which is needed for
effective planning. In the rural region of La Mancha, Veracruz, along the
central region of the Gulf of Mexico we have been working with local
authorities along three lines: land planning along the coast for touristic
development, redefining areas with different degree of risks and redefining
the federal owned land along the coast based on geomorphological and
ecological criteria, a base line monitoring program for the beach, dunes,
mangrove and freshwater wetlands which will help us understand the functioning of these ecosystems and their ability to withstand sea level rise, an
educational demonstrative program with children, women, farmers, cattle
ranchers and fishermen for increasing their awareness on environmental
problems, including greenhouse effects and sea level rise. Finally, in a 750
km state with only three touristic coastal cities, one industrial port and one
gas industry along the coast, we have developed a state program focused
on coastal county majors to make them aware of both the potentials and
the risks of their coastal areas
MORGAN, JACK A.,1,* DIANE E. PATAKI,2 JOSE M. GRUENZWEIG,3
CHRISTIAN KOERNER,4 PASCAL A. NIKLAUS,4 H. W. POLLEY,5
ROBERT S. NOWAK6 and PAUL NEWTON.7 1 USDA-ARS, Fort Collins,
CO, USA; 2 University of Utah, Salt Lake City, UT, USA; 3 Weizmann
Institute of Science, Rehovot, Israel; 4 University of Basel, Basel, Switzerland; 5 USDA-ARS, Temple, TX, USA; 6 University of Nevada, Reno, NV,
USA; 7 AgResearch, Palmerston North, New Zealand. Grassland productivity responses to rising atmospheric carbon dioxide are driven primarily by water relations.
Increases in CO2 and other trace gases are implicated in global warming
that occurred in the past century, and are predicted to cause substantial
increases in temperature and alter precipitation patterns in this century.
However, the direct effect of CO2 on plant and soil responses may be just
as important, and in some cases, more important than their predicted responses to climatic change. Thus, an understanding of how ecosystems
respond directly to rising CO2 concentrations is key to predicting their
future functioning and impacts in a CO2 enriched world. A number of field
and microcosm CO2 enrichment studies have now been completed in natural and semi-natural grassland ecosystems, and a survey of the pertinent
literature revels a wide range of responses to increases in CO2, from no
production responses in alpine grasslands or for wet years in the sub-humid
tallgrass prairie to consistent and substantial production responses every
year in semi-arid shortgrass steppe. We propose that most of the variation
in grassland production to CO2 may be accounted for by variability in
precipitation and soil/plant water dynamics, with production being enhanced relatively more by increased CO2 in drier than in wet systems (or
years). To evaluate this notion, experimental results from several CO2 enrichment studies from contrasting natural or semi-natural grasslands (temperate and Mediterranean grasslands, tallgrass prairie, shortgrass steppe)
and one desert ecosystem were compared. We found that relative aboveground plant biomass enhancements due to increasing CO2 concentrations
were generally stronger as annual precipitation declined, and that these
biomass enhancements were associated with increases in soil and/or plant
water status. The results suggest that CO2-induced production responses in
natural and semi-natural grasslands may be predicted moderately well by
considering only the effects on plant/soil water relations.
MORRIS, JAMES T.1,* and P.V. SUNDARESHWAR.2 1 University of
South Carolina, Columbia, SC; 2 Duke University, Durham, NC. Effects
of nitrogen loading on salt marsh metabolism and sedimentation: interactions with rising sea level.
Long-term fertilization (18 years) of experimental salt marsh plots with
nitrogen and phosphorus has led to an increase in productivity and standing

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243

biomass of Spartina alterniflora, and an increase in the relative elevation

of the marsh surface. While an increase in sediment metabolism decreased
the soil inventory of soil macroorganic matter, the increase in standing plant
biomass increased mineral sediment trapping, leading to a net increase in
marsh elevation in fertilized plots. Interestingly, the stem density in fertilized plots did not decrease, despite an increase in plant stature and standing
biomass. Changes in sediment metabolism were ascribed to changes in root
respiration and production, production and degradation of labile organic
matter, and/or degradation of existing soil carbon. In contrast to primary
producers, which are limited by nitrogen and secondarily by phosphorus,
sediment bacteria, including diazotrophs, appear to be limited primarily by
phosphorous. This implies that interactions between N and P biogeochemical cycles are likely. Thus, while nitrogen loading should have a net positive effect on marsh stability, interactions between plant production, nitrogen loading and soil organic matter transformations are complex and could
alter a number of ecosystem processes.
MOSES, MELANIE E.* and HILARY M. LEASE.* University of New
Mexico, Albuquerque, NM, USA. Ant EnergeticsThe scaling of foraging costs with colony size.
Ant colonies have population sizes ranging from tens to millions. The ant
genus Pogonomyrmex contains species with maximum population sizes
ranging from 100 to 10,000 individuals per colony. We examine what factors might generate these extreme differences in colony size. We investigate
how the energetic cost of foraging scales with colony size in these seed
harvesting desert ants. The energetic cost of foraging is the distance foraged
times the metabolic expense per distance. Ants in larger colonies travel
greater distances to obtain sufficient food for the colony. However, the cost
incurred by traveling longer distances must somehow be compensated for
in large colonies. Here, we test the hypothesis that increased metabolic
efficiency (per ant, per distance traveled) also occurs in larger colonies. We
make field observations of forager population size and foraging distance,
and laboratory measurements of oxygen consumption and ant mass to assess the metabolic cost of foraging in colonies of different sizes. We use
this data to construct a metabolic argument to explain why species can
exist at radically different colony sizes.
MOSIER, ARVIN R.1,* and GARY A. PETERSON.2 1 USDA/ARS, Fort
Collins, CO; 2 Soil and Crop Science Department, Colorado State University, Fort Collins, CO. Trace gas exchange and agroecosystem management in nonirrigated, semiarid cropping systems in northeastern Colorado.
Agroecosystems play an important role in the biosphere-atmosphere exchange of the greenhouse gases CO2, CH4, and N2O. Little data exist to
define how management of different agroecosystems impact trace gas exchange and net global warming potential. In the semiarid portion of the
Great Plains dryland crop production is highly dependent upon precipitation from both snow and rainfall. The typical farming practice in this region
was winter wheat/fallow employing intensive tillage during the summer
fallow period. This practice allows crop production only every second year
and has depleted soil organic matter and soil productivity. In 1985 a field
research site was established to develop more sustainable dryland agroecosystem management techniques for the region. A variety of crop rotations
were imposed under no-till management over a soil gradient at this site. In
the spring of 2002 three systems were selected for study, perennial grass,
wheat/corn/fallow rotation (WCF) and opportunity cropping (OPP) that was
effectively annual cropping of either corn, soybeans or wheat. Gas flux
measurement sites were established at mid and toe slope positions of the
field within grass, WCF and OPP plots and gas flux measurements were
made one or two times per week to attempt to quantify the impact of
management on net GWP through measurement of CO2, CH4, N2O flux
and soil carbon storage. Both WCF and OPP were cropped with corn, but
the lowest amount of precipitation recorded in the last 85 years in eastern
CO was received from September 2001to July 2002, and crop production
was minimal. During this extremely dry year, crop management did not
impact trace gas exchange. Methane oxidation rates were lower in the perennial grass than in cropped fields while N2O emissions were higher in
both corn rotations than in grass.

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Abstracts

MOURA, CATARINA F.,1,* ANDREW J. MCELRONE,1 JONATHAN

WATKINSON,2 RUTH GRENE2 and ROBERT B. JACKSON.1 1 Duke
University, Durham, NC; 2 Virginia Tech, Blacksburg, VA. Genome-wide
effects of elevated CO2 in the loblolly pine FACE experiment.
Rising atmospheric CO2 concentrations are altering the present environment
and creating new challenges for forest tree species. Physiological and structural responses to environmental change are controlled by molecular processes including changes in the expression of a large number of genes.
Studies to date generally involve a non-woody plant (Arabidopsis), greenhouse conditions and/or a limited number of genes. There is a special need
to understand global cellular responses in more complex organisms and to
clarify the role of certain genes in their natural context. This study provides
a larger, genome-wide view of the responses of an important and fieldgrown woody species, loblolly pine (Pinus taeda), to changes in atmospheric CO2 concentration. We compared the patterns of gene expression
in needles of loblolly pine exposed to Ambient and Elevated (Ambient +
200ppm) CO2 levels in the Duke Forest FACE experimental site. For this
purpose we used microarray slides each containing ;2000 loblolly pine
ESTs (expressed sequence tags). Among the ESTs there are some coding
for enzymes involved in photosynthesis and nitrogen metabolism, two processes important for the response and potential acclimation of loblolly pine
to elevated CO2. Understanding the molecular mechanisms underlying
physiological acclimation will allow us to better predict responses of forests
under future climatic scenarios.
MOYER, MELISSA A.,1,* JAMES W. MCCOWN,2 THOMAS H. EASON3 and MADAN K. OLI.1 1 University of Florida, Gainesville, FL,
U.S.A.; 2 Florida Fish and Wildlife Conservation Commission, Bear Management Section, Gainesville, FL, U.S.A.; 3 Florida Fish and Wildlife Conservation Commission, Bear Management Section, Tallahassee, FL, U.S.A.
Home range dynamics of the Florida black bear in Ocala National
Forest.
An understanding of how animals distribute themselves across an available
landscape is necessary for conservation and management planning, particularly for large carnivores existing in an increasingly human dominated
landscape. The human population of central Florida is expanding rapidly
which will result in loss and fragmentation of black bear habitat. Implementation of effective conservation measures will require an understanding
of their space use patterns. Black bears were captured within a 50,000 ha
study area in the Ocala National Forest (ONF) as part of a broader Florida
Fish and Wildlife Conservation Commission (FWC) study. All females and
several males were fitted with radio-collars and then located with ground
or aerial telemetry 2-3 times per week. Locations were compiled to estimate
annual and seasonal home ranges using the minimum convex polygon. For
each bear, the sizes of the annual and seasonal home ranges were quantified.
A simultaneous display in ArcGIS of each home range estimate indicated
significant overlap between bears. Overlapping areas were quantified. The
space use pattern and how it may adjust with seasonal change could have
important consequences for management and conservation, particularly as
space becomes more limited in the future.
MOYES, ANDREW B.1,2,* and JOHN A. GAMON.1,2 1 California State
University, Los Angeles, Los Angeles, CA; 2 CEA-CREST, CSULA, Los
Angeles, CA. Prescribed fire effects on a California annual grassland.
This project was designed to assess the effects of a prescribed April burn
on efforts to restore a California annual grassland dominated by the introduced annual grass Bromus diandrus. The fire reduced the number of viable
B. diandrus seeds by around 96%, but did not significantly affect forb seed
abundance or composition. The destruction of grass seed led to replacement
of B. diandrus by introduced forbs, especially black mustard (Brassica
nigra), which corresponded with delayed productivity and altered microclimate variables, including soil moisture and air and soil temperature. A
second study is being conducted to determine how these burn effects and
nutrient mineralization by the burn impact the reintroduction of native species (the perennial grass Nassella pulchra and the shrub Artemisia californica). Results from this second study will also be included in the poster.

MULDER, C. P.H.1,* and B. A. ROY.2 1 University of Alaska, Fairbanks,

AK; 2 University of Oregon, Eugene, OR. Climate change and invertebrate herbivory on boreal understory plants: A survey.
Rapid changes in climate are predicted for interior Alaska. Much attention
has been focused on the potential physiological responses of boreal plant
species, but changes in invertebrate population dynamics may also alter the
growth and reproduction of plants. To examine climatic influences on herbivore damage, we surveyed damage on two species that are common
across a range of climatic conditions in interior Alaska: Alnus crispa (green
alder) and Vaccinium vitis-idaea (lowbush cranberry). Ten individuals of
each species were examined at each of nine sites: three cold and wet, three
moderate and three warm and dry. Diversity of herbivores was high for
alder (a mean of 4.3 per individual, 11 species total) but only one herbivore
was found on cranberry. For alder, distance to nearest conspecific neighbor
was negatively related to total percent damage, and plant size was positively related. Alders at the moderate sites had significantly higher levels
of damage (% leaf damaged) than those at cold and wet or warm and dry
sites. However, individual herbivore species differed in their overall pattern
of abundance across the different sites. Cranberry showed increasing levels
of damage from cold and wet to moderate to warm and dry sites. For alder,
increased herbivory significantly reduced reproduction (as measured by
number of cones). We conclude that both the environmental conditions and
local density of plants affect diversity of herbivores and damage to leaves
in these species. Since invertebrates are more mobile than plants, potential
changes in herbivore-plant dynamics should be included in predictions of
altered vegetation distributions resulting from climate change.
MULLAN, CAITLIN P.,* BRIAN R. SILLIMAN and MARK D. BERTNESS. Brown University, Providence, RI, USA. Spatial segregation of
New England marsh plants across an estuarine salinity gradient.
While it has long been recognized that marsh plant community composition
shifts across estuarine salinity gradients, the mechanisms responsible for
these species shifts have never been experimentally examined. We conducted a series of experiments in southern New England marshes to investigate the roles of physical and biotic interactions in driving species
segregation along this gradient. Greenhouse studies revealed that all of the
common plants in this system grow better in fresh water than in full
strength salt water. Field transplant experiments, using 10 common plants
in the system, showed that when salt marsh plants were transplanted to
tidal freshwater marshes, in the absence of neighbors (without competition)
they grew better than they did in salt marshes, but were strongly suppressed
when neighbors were present. In contrast, when freshwater marsh plants
were transplanted to salt marshes, they did poorly and generally died with
or without neighbors (competition) present. These results suggest that while
fresh marsh plants are limited from salt marshes by physiological stress,
salt marsh species are limited from fresh marshes by competitive displacement. Thus, theory developed along vertical intertidal gradients, that superior competitors dominate physically benign habitats and displace weaker
competitors to physically harsh habitats, can be further generalized to horizontal landscape-scale gradients. Experimental results contribute to our
understanding of the organization and assembly of the full range of estuarine marshes and have important implications for predicting how marsh
plant communities will respond to altered hydrology (e.g. freshwater diversions, sea level rise), coastal eutrophication, and invasive species.
MULLIGAN, MARGARET R.* and DAVID L. GORCHOV. Miami University, Oxford, Ohio. Quantifying population loss of goldenseal, Hydrastis canadensis L., in Ohio.
Goldenseal, Hydrastis canadensis L. (Ranunculaceae), is a slow-growing
perennial herb that is harvested for the medicinal properties of its rhizome.
Goldenseal has become increasingly rare throughout its range, and was
listed on the Convention for International Trade on Endangered Species
(CITES) in 1997, but it is not clear to what extent this is due to overharvesting versus habitat loss or degradation. In Ohio, the core of its range,
there is minimal information on the status of goldenseal, but the number
of populations is believed to be declining. Voucher specimens from eight
herbaria in Ohio were examined to determine what proportion of sites that
historically supported goldenseal populations still support them today. Out

of 269 voucher specimens reviewed, 71 (dated 1845 -1998) had sufficient

detail for the collection site to be relocated. Eight of the sites (11%) were
deforested and no longer supported populations. The remaining 63 sites
were searched for goldenseal in summer 2002 and the number of plants
(ramets) present recorded for each. Goldenseal was found on 40 (63%) of
these forested sites. Most (54%) of these populations had , 200 plants,
19% had 200 to 1000 plants and 27% had .1000 plants. Stratifying sites
by age of the record, 20% of early (1845-1933), 50% of intermediate
(1957-1971) and 58% of recent (1977-1998) sites still contained goldenseal. Among sites that no longer contained goldenseal, none of the early,
75% of intermediate and 83% of recent sites were still forested, indicating
deforestation has played a minor role in recent extinctions. This study revealed that nearly half of documented goldenseal populations have gone
extinct, suggesting an overall decline in numbers of goldenseal populations
in Ohio. Most of the population loss is due to factors other than habitat
loss, suggesting overcollection has been an important cause of this decline.
MUNGUIA, PABLO.* Deartment of Biological Sciences, Florida State
University, Tallahassee, Florida, United States. Testing metacommunity
concepts with density manipulations of Pen shell (Atrina rigida) communities.
Mechanisms acting on different spatial scales have not been easy to understand because of difficulties in conducting large-scale experiments. Further, species vary in their perception of space (related to behavior) and time
(related to life history), making it difficult to interpret patterns of diversity
and richness. Therefore, a community must be understood as a melange of
species, where forces acting at the community level will have different
effects depending on the individual and combined responses of component
species. Here I present a model benthic system used to address communitylevel mechanisms acting at different spatial scales. At St. Joe Bay, Florida,
the pen shell (Atrina rigida) is one of the few sources of hard substrate,
serving as a habitat for both sessile and mobile invertebrates, as well as
fish. Habitat was constructed at different scales by anchoring empty shells
in arrays of varying density. Treatments included age of the community,
distance from the nearest community, as well as simulated destruction of
communities. The effects of these treatments varied depending on the group
of species present on a pen shell. Pen shell communities exhibit species
successional patterns that affect the relationship between spatial scales.
Furthermore, these patterns seem to differ depending on the life history of
the group of species. In order to understand how mechanisms that produce
patterns of relative abundance and species richness change with scale, we
must take into account the natural history of the component species.
MURDOCH, WILLIAM,1,* SUSAN SWARBRICK1 and CHERYL
BRIGGS.2 1 University of California, Santa Barbara, CA; 2 University of
California, Berkeley, CA. Biological control of red scale.
In most cases of successful biological control there is likely to be more
than one natural enemy that attacks the pest. Credit for control is sometimes
given to just one of these enemies, and in others it has been argued that
the multiple enemies are in some way complementary, and perhaps even
essential to control. This is a difficult issue to resolve. We created experimental outbreaks of California red scale, a pest of citrus whose successful
control in southern California requires Aphytis melinus. We have shown
that Aphytis responded to increased scale density and that the mortality it
imposed can account for outbreak suppression. It appears that the other
agents present did not respond, or not adequately, to increased scale density. The other enemies may, however, have an important effect on control
by increasing red scale ?background? death rate. We are investigating this
possibility in a model of the system.
MURPHY, C. A.,* B. L. FOSTER, M. E. RAMSPOTT, K. P. PRICE, T.
HILDEBRAND and L. EVANHOE. University of Kansas, Lawrence, Kansas. Spatial scaling of plant species richness in grasslands: relationships to productivity, soils and management.
Management regimes of varying types and intensities can have profound
impacts on grassland biodiversity at various spatial scales. We examined
plant species richness in relation to productivity and soil characteristics at
several spatial scales in grasslands of Northeastern Kansas. Plant species

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245

richness, biomass, soil bulk density, soil carbon and nitrogen were measured in four managed grassland regimes (cool season hay and grazed,
warm season hay and grazed). In 2001, data were collected within fiftyfour, 1m2 quadrats placed systematically within an area of 14,400m2 in each
of nine fields. The program, EstimateS, was used to generate species accumulation curves for each field and estimates of species richness at different spatial scales (1-54m2). Among fields, a unimodal, quadratic relationship was found between species richness and biomass at all spatial
scales. Species richness was negatively correlated with soil bulk density,
but positively correlated with soil carbon and nitrogen. The variance in
richness explained by soil variables, and the slopes of these relationships,
increased with spatial scale, reflecting large differences in spatial heterogeneity among fields of contrasting management. The observed relationships of plant species richness to productivity and soils reflect a complex
gradient of human-induced disturbance associated with current and historic
agricultural land-use.
MURPHY, KATHRYN A.* and RACHEL OMALLEY. California State
University - San Jose, San Jose, CA. Pollination ecology of a self-compatible endangered annual plant: Chorizanthe robusta var. robusta.
Pollinators can be essential to the survival and evolution of most plant
species, ensuring the exchange of genetic information and reproductive
success. For highly endemic and endangered species, the pollinator-plant
relationship may be even more crucial. Yet, studies of endangered plant
pollination ecology are few. The robust spineflower (Chorizanthe robusta
var. robusta), a small annual plant found originally throughout the central
coast of California, is now limited to seven known populations, all occurring in Santa Cruz and Monterey Counties. In order to better understand
the pollination ecology and breeding system of this federally endangered
plant, this research evaluates the frequency, importance, ranges and habitat
needs of likely pollinators through extensive observation and experimental
bagging of inflorescences. It also correlates pollinator visitation frequency
with environmental conditions and with characteristics of two robust spineflower populations in Santa Cruz County. C. robusta var. robusta appears
to support a generalist pollination system, with over 15 families of insect
visitors at each population. Spatial and temporal variability in the pollinator
assemblage and flowering phenology indicate that the wide array of pollinators may function to provide pollinator services for the robust spineflower throughout the flowering season and in varying climate conditions
at the sites. In addition, although the robust spineflower is self-compatible,
pollinator access to flowers increased seed set significantly, indicating that
insect pollinators increase plant fitness. At one population, a high density
of other plant species had a negative correlation with the frequency of
visitation to C. robusta var. robusta flowers. Based upon habitat correlations for important pollinators as well as literature research of their habitat
requirements, this project recommends that local agencies and the US Fish
and Wildlife Service monitor and potentially control non-native plant species growing within patches of robust spineflower plants. This project also
recommends restricting insecticide use in adjacent properties to protect the
plant-pollinator relationship.
MURPHY, PETER J.1,* and KAREN LIPS.2 1 Idaho State University, Pocatello, ID, USA; 2 Southern Illinois University, Carbondale, IL, USA. UVB and water temperature unlikely cofactors in Central American amphibian declines due to pathogenic chytrid.
The pathogen Batrachochytrid dendrobatidis has been linked with multispecies amphibian declines in Australia, the United States, and Central
America. We sought to identify environmental cofactors that may predispose upland and cloud forest amphibians to an epidemic caused by this
pathogen. Two cofactors, temperature and UV-B exposure, have both exhibited mean increases during the past two decades in Central America.
We tested the effect of these cofactors on tadpole infection in a mesocosm
experiment in Chiriqui, Panama. We measured infection in Smilisca phaeota (Hylidae) tadpoles, a species whose adults and tadpoles were susceptible
to the pathogen in preliminary trials. We measured infection as the proportion of keratin lost in tadpole mouthparts due to fungal infection. Two
levels of three treatments were crossed in the factorial experiment: ambient
vs. reduced UV-B, ambient vs. increased water temperature, and ambient
vs. increased pathogen loads. The water source for the shallow, flow-

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Abstracts

through mesocosms was a stream in which infected tadpoles were found

both summer 2001 and 2002. Our results indicated that increased temperature strongly protected tadpoles from infection, UV-B did not change tadpole infection rates, and adding pathogen increased tadpole infection. Interestingly, tadpoles were protected at water temperatures at which the chytrid grew optimally under lab conditions. Our study suggests that increasing
temperatures and UV-B are not linked directly to epidemic outbreaks of B.
dendrobatidis, especially because warmer water protects tadpoles. The
study is particularly convincing because it was conducted at a site where
species declined due to this pathogen, hence UV-B and temperature treatments were balanced by ambient controls. Moreover, we used locally isolated B. dendrobatidis for the experiment, hence the pathogen strain used
was that which had initially driven the declines, not an exotic with which
the frogs and tadpoles had no prior exposure.
MURPHY, SHANNON K.1,* and DAVID M. RIZZO.2 1 skmurphy@
ucdavis.edu, University of California, Davis, CA; 2 University of California, Davis, CA. Surveying two Phytophthora species in California plant
communities: Comparing an invasive with a native pathogen.
Phytophthora ramorum, causal agent of Sudden Oak Death, has caused
extensive mortality of Quercus spp. and Lithocarpus densiflorus in California forests and woodlands. This invasive pathogen also infects over 15
other native species including Umbellularia californica, Arbutus menziesii
and Sequoia sempervirens. We recently have discovered another Phytophthora, tentatively named P. nemorosa, that has a nearly identical host
range and distribution as P. ramorum. Phytophthora nemorosa causes similar disease symptoms as P. ramorum on many hosts but does not cause
landscape level mortality of oaks and tanoak as does P. ramorum. P. nemorosa appears to be indigenous to California forests and woodlands. A
total of 120 circular plots (18m radius) were established at ten sites in
northern California (distance between furthest sites ;130 km east to west;
;220 km north to south) to determine incidence of P. ramorum and P.
nemorosa and relate pathogen incidence to plant community structure.
Three types of native forest and woodland communities were surveyed:
northern mixed evergreen forest, coast live oak woodland, and Oregon oak
woodland. All woody plant species were examined for the presence of
Phytophthora. Phytophthora species were identified using both cultural and
molecular detection techniques. We evaluated plant species composition,
density, and environmental variables, including slope, aspect, and elevation.
Preliminary results confirm P. ramorum at five of ten sites and within 20
of 120 plots, and P. nemorosa presence at nine sites and within 47 of 120
plots. The two species co-occurred on eight plots with one co-occurrence
on the same individual L. densiflorus tree. This study is the first to examine
the ecological associations between these two pathogens and will lead to
further studies concerning competition and possibly hybridization.
MURRAY, TANYA. Syracuse University, Syracuse, NY. Survey of the
effects of ungulate grazing on the relationship between VAM fungi and
plants in Yellowstone National Park.
Comprehensive understanding of vesicular-arbuscular mycorrhizae in systems that support grazers may offer valuable insight to the mechanisms by
which plants respond to grazing pressure. A survey of seven sites in Yellowstone National Park compares the species diversity and structural characteristics of VAM fungi collected inside and outside long term grazing
exclosures. These sites are located in the northern range of YNP where
ungulates (Bison bison and Cervus elaphus) graze during the winter
months. The sites vary in slope and moisture from dry upland to wet
lowland soils. Data obtained from random samples are compared to data
from dominant plant samples to determine landscape and grazing influences
on mycorrhizal composition and structural characteristics. For each site
VAM species diversity is determined from spore density and diversity.
Plant benefit from the mycorrhizal association is assessed using a supplier
ratio based on the relative investment in arbuscules vs. other components
of internal colonization. Finally, mechanisms of mycorrhizal function with
and without grazing pressure is analyzed by comparing internal and extramatrical hyphal abundance from soil and root samples collected inside and
outside grazing exclosures.

MURRELL, KATHREN E.,1,* ANN DENNIS2 and MICHAEL BARBOUR.3 1 University of California at Davis, kemurrell@ucdavis.edu, Davis,
CA; 2 CAL FLORA, Albany, CA; 3 University of California at Davis, Davis, CA. Patterns of species and habitat richness and diversity in
grazed meadows in the eastern Sierra Nevada.
The Golden Trout Wilderness study examined how impacts on physical
characteristics of meadows affect biodiversity. Our hypothesis was that
meadow species richness is related to the range of habitats represented, and
that richness within specific affinity groups is related to the representation
of the habitat those species depend upon. An alternate hypothesis was that
meadow area was more important than habitat in determining species richness. Species richness and habitat richness were positively correlated at
R25 .53 and Prob.[t]5.0109. Neither the correlation between meadow
size and species richness nor the correlation between meadow size and
habitat richness was significant. Species diversity peaked at an intermediate
level of habitat diversity (R25 .69 and Prob.[t]5.0023), suggesting that
some habitats did not contribute substantially to species diversity. As a
result of these analyses, we hypothesized that habitats in recently incised
or degraded channels contribute less to species diversity than non-channel
habitats. When analyzed separately, non-channel habitat diversity and species diversity displayed a positive linear correlation with an R25 .43 and
Prob.[t]5.02. According to Tukeys comparison of means, non-channel
transects contributed more to species richness than channel transects, and
stringers had significantly higher species richness than meadow sampling
units. Results also indicated that a group of species missing in grazed
meadows in the Southern Sierra Nevada were also absent in grazed meadows in this study where sod banks, willows and stringers provided the only
remaining habitat. We concluded that in these meadows, habitat richness
was more important than meadow area in determining species richness, and
that non-channelized habitats and stringers contributed the most to plant
diversity.
MUSS, JORDAN D.,* DIANNE V. OWEN and JOHN C. VOLIN. Florida
Atlantic University, Davie, FL. Predicting the landscape spread of Lygodium microphyllum in South Florida.
Invasions by non-indigenous plant species can profoundly disrupt the original composition, structure and function of native plant communities. In
Florida, over 4,000 plant species have been introduced, 125 of which are
listed as invasive by the Florida Exotic Pest Plant Council. Among these,
Lygodium microphyllum is of major concern because of its recent rapid
spread. In 1978, 14 infested sites were documented in Martin County, Florida. By 1999 it had spread throughout the ten southernmost counties of
peninsular Florida, occurring at nearly 1,200 documented sites. To understand the rates and patterns that characterize the spread of L. microphyllum,
we have developed a predictive, spatially explicit, cellular model of its
dispersal and establishment within the South Florida landscape. Parameter
values estimated from field and laboratory data on habitat susceptibility to
invasion, growth habits of L. microphyllum, spore production rates and
dispersal profiles were used to calibrate the model for the period from 1978
to 1993. Model validation was performed using data on documented invasion sites for the period from 1995 through 1999, and will be extended
to 2001 when the data is released. Validation of model output, using a
permutation procedure that is not biased by spatial autocorrelation, showed
good agreement between the model and data, with correlations generally
higher than 0.5 across a range of spatial grains. Model runs initialized to
the documented 1999 invasion sites show that L. microphyllum will spread
to most of the suitable habitats found in South Florida within the next
decade.
MUTCHLER, TROY,1,* MICHAEL J. SULLIVAN1 and BRIAN FRY.2
1
Mississippi State University, Mississippi State, MS; 2 Louisiana State University, Baton Rouge, LA. Response of seagrass community food webs
to nutrient enrichment with a 14N labeled fertilizer.
Studies using stable isotopes to delineate trophic pathways in aquatic systems have become increasingly common. While most studies have been
descriptive in nature, greater efforts are being made to couple tracer additions with experimental manipulations. We used 14N tracer additions coupled with multiple fertilization strategies to investigate the potential for

cultural eutrophication of coastal waters to alter food web dynamics in

seagrass communities. Two levels of fertilization and a no-fertilization control were employed in small seagrass beds off Horn Island, Mississippi. In
the low fertilization addition, the fertilizer contained a known d15N signature that allowed us to trace the fate of our added nitrogen into the primary
producers and consumers. Both the seagrass and epiphytes were strongly
labeled by our 14N tracer in beds with fertilized sediments (as low as -90
and -68, respectively), while beds receiving water column fertilization
were labeled to a lesser degree (-24 and -17, respectively). Control
seagrass and epiphytes had d15N values of +5. This isotopic label was
then transferred to the consumers within the system. The uptake of label
by the primary producers and the subsequent transfer of label to consumers
indicates that nitrogen additions to these systems are in fact entering the
food web and differential uptake of these additions could lead to shifts in
the relative importance of individual primary producers within these communities.
MUTH, CHRISTINE C.,* AMBER VOGEL, BETTY W. BROWN and
WALTER E. BOLLENBACHER. Department of Biology, University of
North Carolina at Chapel Hill, Chapel Hill, NC, USA. An ecosystem model for science education reform.
Science education occurs not in an isolated classroom but within a complex
educational ecosystem. This ecosystem includes not only science teachers
and students, but also teachers of other disciplines, school administration,
local government, the physical school environment, learning resources and
tools, funding, and school location (e.g., urban vs. rural). Furthermore, this
ecosystem extends into the larger community to students families, community organizations, colleges and universities, and even businesses. In
order to understand the context in which students learn science, we have
developed an ecosystem model of science education. Our model identifies
components of the education ecosystem, describes the flow of knowledge,
resources and decision-making throughout the system, and defines interrelationships at multiple levels. Insights provided by the model can be used
to identify best practices, organizationally and educationally, and thus contribute to reforming science education. Widening Horizons in Science Education (WHISE) is a research project investigating the efficacy of the
ecosystem model as a tool for science education reform. WHISE is being
conducted at the secondary level with four North Carolina school systems,
all rural, low-wealth, and low-achieving in science education. Based on our
ecosystem model, we have developed and implemented initiatives to
achieve specific outcomes (e.g., student performance, students pursuing
science careers, administration support, changing how science is taught)
that broadly impact the science education ecosystem. Programmatically,
investigations include teacher professional development, interdisciplinary
curricula, community awareness, policy maker collaboration, and marketing techniques. Goals are to influence the ecosystem at multiple levels of
organization and function in ways that synergistically increase the quality
of science education. Ongoing external evaluation and research findings
enable the project to evolve in optimal directions and yield insights into
how secondary science education can best be improved to meet a pressing
national need. This work is supported by grants from NIH (RR016306)
and GlaxoSmithKline.
OZ, ALEJANDRO A.* and MARY T.K. ARROYO. Universidad de
MUN
Chile, Santiago, Chile, RM, Chile. Negative impacts of vertebrate predators on insect pollinator visitation and seed output in Chuquiraga oppositifolia.
Studies on plant-pollinator interactions have mostly neglected the potential
negative effects of predators of pollinators on seed output, even though
pollinator anti-predatory behaviour may affect visitation patterns, pollen
transfer, and ultimately plant reproductive output. We tested the hypothesis
that lizards and insectivorous birds, by reducing pollinator visitation, can
have significant negative effects on seed output in the insect-pollinated,
self-incompatible shrub, Chuquiraga oppositifolia (Asteraceae), in the lower alpine belt in the central Chilean Andes. This alpine belt possesses a
high density of Liolaemus (Tropiduridae) lizards, and is characterized by
the dominance of low shrubs interspersed among rocks that are inhabited
by lizards and used by flycatchers Muscisaxicola (Tyrannidae) as perching
sites. In a 2x2 factorial predator exclusion experiment, visitation rates of

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247

the most frequent pollinators, a satyrid butterfly and a syrphid fly, the
duration of pollinator visits, and C. oppositifolia seed output, were 2-4
times greater when lizards were excluded, while birds had no effect. Further, in a natural experiment, visits by syrphids were 9 times shorter, and
pollinator visitation rates of satyrid butterflies and syrphid flies, and C.
oppositifolia seed output were 2-3 times lower on shrubs growing adjacent
to lizard-occupied rocks compared to those distant from rocks. Our results
suggest that the presence of lizard predators, by reducing pollinator visitation, can elicit strong top-down indirect negative effects on seed output
in plants. Acknowledgements: FONDECYT 2010032 (A.A.M.) and P99103-F-ICM (M.T.K.A)
NAGEL, JENNIFER,1,* TRAVIS HUXMAN,2 KEVIN GRIFFIN3 and
STANLEY SMITH.4 1 Texas Tech University, Lubbock, TX; 2 University
of Arizona, Tucson, AZ; 3 Columbia University, Lamont-Doherty Earth Observatory, Palisades, NY; 4 University of Nevada, Las Vegas, NV. Carbon
dioxide enrichment reduces the energetic cost of biomass construction
in an invasive desert grass.
A central issue in plant ecology is how plant communities will be impacted
by human-induced environmental changes. These community-level changes
will be influenced by the differential responses of constituent species to
changing resources. Evaluating the effects of projected changes in resource
availability on co-occurring nonindigenous and native plant species in invaded ecosystems could provide insight useful in developing reliable predictions of how these plant communities could change in the future. While
deserts presently are among the least invaded ecosystems, they have been
predicted to be one of the most responsive ecosystems to global change.
To examine how increasing atmospheric CO2 partial pressure could influence invasion dynamics in an intact Mojave Desert ecosystem, we compared the responses of energetic properties and growth of invasive and
native grass species to atmospheric CO2 enrichment at the Nevada Desert
Free-Air CO2 Enrichment Facility. Combined with its modest positive influence on photosynthetic activity, elevated atmospheric CO2 was associated with a significant reduction in the energetic cost of aboveground biomass construction in invasive Bromus madritensis spp. rubens (red brome)
without a concurrent cost reduction in native Vulpia octoflora (six-weeks
fescue). Consequently, the invasive grass species grew faster and bigger
and produced more seeds with atmospheric CO2 enrichment than the native
grass species. Because plant growth involves both the assimilation and
expenditure of energy, it has been suggested that the most successful plant
species in a given environment could be those that maximize their energetic
gains while minimizing their energetic costs. As a physiological mechanism
of invasive species success driven by CO2 enrichment, such reductions in
the energetic cost of biomass construction combined with increased photosynthetic activity could trigger a shift in the species composition of this
Mojave Desert ecosystem, and potentially that of other invaded ecosystems,
toward increased invasive species dominance.
NAGELKERKE, KEES. University of Amsterdam, nagelkerke@science.
uva.nl, Amsterdam, Holland, Netherlands. The effect of climate change
on species9 ranges.
The consequences of climate change depend on species9 ability to track it.
I model the effect of a moving North-South climate profile on the latitudinal range of a metapopulation. It is assumed that climate affects one or
more of the main metapopulation parameters (amount of habitat that is
regionally available, local extinction rate and colonization rate). When
ranges shift in response to climate, tracking delays may develop on both
the advancing and the retreating edge of a range. The difference between
the two delays determines the effect on range size. It appeared that a moving climate lowers regional population size. However, ranges may become
larger when populations on the retreating edge take a long time to disappear. Remarkably, global population size may then increase. Species not
able to track the climate fast enough go extinct because their retreating
edge moves faster than, and eventually collides with, the advancing edge.
Species with intermediate rates of patch turnover may be the first to disappear. Results depend very much on which of the parameters climate
actually acts on, especially concerning the retreating edge. Tracking ability
increases with range size, amount of habitat, dispersal rate and dispersal
distance. Conventional predictions based on modeling migration speeds

248

Abstracts

under a homogeneous climate can seriously overestimate the ability to track

a climate profile. Climate change greatly increases vulnerability to habitat
destruction and dispersal impairment.
NAKATANI, NAOKI,* TAKI OKUNO and KOJI OTSUKA. Osaka Prefecture University, Sakai, Osaka, Japan. Evaluation of Carbon and Nutrient Fixation Effect in An Artificial Lagoon.
Quality of seawater significantly depends on quantitative relationship between pollution load and natural purification capacities. Water quality in
an enclosed sea, where the big industrial cities are surrounded, is chronically bad since the pollution load is much higher than the natural purification capacities. In order to purify the seawater, several coastal eco-techniques have been proposed already. An artificial lagoon, having calm water
area surrounded by rubble-mound breakwater, is one of the solutions. Many
researchers have pointed out that the artificial lagoon has several water
purification functions, such as wave breaking re-aeration, biological contact
oxidation in the rubble-mound breakwater, transparency increase due to
sinking of suspended matter, carbon and nutrient fixation of seaweed, oxygen supply of seaweed, creation of sound biological material circulation.
However, the water purification mechanisms by seaweed and its biological
process have not been clarified yet. This paper deals with the water purification mechanism, and also the prediction method of carbon and nutrient
fixation in the artificial lagoon by using the coastal ecosystem model. The
numerical simulation model is constructed as material circulation of carbon,
nitrogen and phosphorus. The predicted results show that the fixation of
carbon and nutrient mainly depend on the seaweed biomass, and that the
total fixation decreases year by year, because of the accumulation of the
sedimentary organic matter. The rock filter is not so effective for the carbon
and nutrient fixation. The fixation efficiency increases as the depth of the
inner sea is decreasing. At first, the results of field investigation are summarized, and then the ecosystem model is introduced as the prediction
method. In order to evaluate the carbon and nutrient fixation effects in the
lagoon, the prediction method, a conservation model of the materials, is
proposed for the evaluation.
NANNINI, MICHAEL A.1,* and MARK C. BELK.2 1 Environmental Protection Agency, Research Triangle Park, NC, USA; 2 Brigham Young University, Provo, UT, USA. A field test of the antipredator responses of
two native cyprinid prey fish exposed to predation by introduced
brown trout.
We examined how differences in antipredatory strategy affected mortality
where introduced brown trout Salmo trutta prey on two native cyprinids,
leatherside chub Gila copei and redside shiner Richardsonius balteatus.
Previous work has determined that leatherside chub and redside shiner respond to both the presence of brown trout and simulated attacks by a model
brown trout. However, the response of these two prey species differs markedly. Redside shiner increase their activity level in the presence of brown
trout, respond earlier and quicker to attacks, and have a more maneuverable
escape response. In contrast, leatherside chub decrease their activity level
in the presence of brown trout, respond to attacks later and slower, and
have a less maneuverable escape response. Antipredatory strategies used
by other similar prey indicate that the strategy used by redside shiner ought
to be more effective against brown trout predation than the strategy used
by leatherside chub. To test this hypothesis we examined the mortality
inflicted upon these two prey species by brown trout in a field setting.
Enclosures were set up so that equal numbers of both prey species were
either exposed to predation by brown trout or not. Brown trout significantly
affected survival of both species. There was also a trout by species interaction in which the effect of brown trout on mortality of leatherside chub
was stronger than the effect of brown trout on mortality of redside shiner.
These results indicate that leatherside chub are more heavily impacted by
brown trout than redside shiner.
NATALI, SUSAN M.,1,* MATT LANDIS,2 JESSICA GUREVITCH1 and
GORDON FOX.3 1 SUNY, Stony Brook, Stony Brook, NY, US; 2 Middlebury College, Middlebury, VT, US; 3 University of South Florida, Tampa,
FL, US. Age and growth of pitch pines (Pinus rigida) in the context of
frequent disturbance.
Fire is an integral factor in the maintenance of pine barren stands along
the Atlantic coastal plains. Pitch pine (Pinus rigida) is the characteristic

and dominant species in the pine barrens of Long Island, NY, which cover
approximately 100,000 acres in south-central/southeastern Long Island.
Historically, pine barren stands on Long Island burned very frequently with
severe enough fires to result in partial or full stand regeneration about
every10-30 years. Since the enormous population increase on Long Island
following WW II and the rise of suburbanization, fire frequency has greatly
decreased due to fire-suppression and increased fragmentation of the pine
barrens. In this study we used a dendrochronological analysis to determine
age distribution, mortality, recruitment, and growth of P. rigida in 15 sites
of varying burn histories in the Long Island Pine Barrens. A greater understanding of the forces shaping pitch pine populations promises to offer
insight into the dynamics of disturbance-prone populations and communities.
NEATROUR, MATTHEW A.,1,2,* ROBERT H. JONES1 and STEPHEN W.
GOLLADAY.2 1 Department of Biology, Virginia Tech, Blacksburg, VA;
2
Joseph W. Jones Ecological Research Center, Route 2, Box 2324, Newton,
GA. Nutrient and fine root heterogeneity in forested wetlands with
contrasting hydrologic regimes.
Nutrient heterogeneity and fine root distribution are related in upland forests of southeastern United States. However, it is not clear whether these
relationships exist in forested wetlands where lack of oxygen can limit fine
root growth. We used a combination of univariate, multivariate, and geostatistical techniques to investigate how nutrient heterogeneity affects fine
root distribution in three types of forested wetlands: floodplain swamps,
river swamp sloughs, and depressional swamps. Floodplain swamps and
river swamp sloughs are alluvial ecosystems and receive nutrient subsidies
from streams during flood pulses. Depressional wetlands, in contrast, are
hydrologically isolated from streams and receive most of their nutrients via
precipitation. We measured standing fine root biomass and nutrient availability (N-NO3, N-NH4, and P-PO4) in one representative site of each wetland type and predicted that fine root biomass would be more strongly
related to nutrient availability in alluvial swamps. Fine root biomass was
not related to nutrient availability at any of the sites. Multiple regression
analyses showed that nutrient availability explained little of the variation
in fine root biomass. In addition, fine root variability, as indicated by the
coefficient of variation (CV), was relatively low at all of sites (33-50%)
and did not match nutrient CV (53-209%). Semivariograms showed that
fine root biomass and nutrient availability were spatially autocorrelated.
These data suggest that fine root biomass and nutrients are patchily distributed in these forested wetlands; however, fine root biomass and nutrients
are not congruent.
NELSON, DAVID H.1,* and CYNTHIA B. SCARDAMALIA-NELSON.2
University of South ALabama, Mobile, AL, USA; 2 Providence Hospital,
Mobile, AL, 36688. Vertebrate road-kill survey of the Mobile Bay
Causeway.

A systematic, road-kill survey was conducted (by bicycle or automobile)

on the Mobile Bay Causeway each week during 2001 and 2002 to assess
the numbers of vertebrates killed by vehicular traffic. More than 1000 organisms representing 85 species of vertebrates were encountered: 8 amphibians, 25 reptiles, 39 birds and 13 mammals. Southern leopard frogs
(Rana utricularia) were the most abundant amphibian (n516). Endangered
Alabama red-bellied turtles (Pseudemys alabamensis) were the most frequently encountered reptile (n5131); most specimens were hatchlings.
Laughing gulls (Larus atricill, n582) and American coots (Fulica americana, n562) were the most commonly encountered birds. Raccoons (Procyon lotor, n587), opossums (Didelphis marsupialis, n582), and nutria
(Myocastor coypus, n559) were the most abundant mammals. Data were
analyzed by taxon and season. The most significant finding in this ongoing
study so far has been the emergence of 102 hatchlings of Pseudemys alabamensis during the months of March and April. These data confirm overwintering in the nest and document significant roadkill mortality of hatchlings of the Alabama red-bellied turtle.
NELSON, DAVID M.,1,* FENG SHENG HU,1 JIAN TIAN,1 THOMAS A.
BROWN,2 IVANKA STEFANOVA3 and HERBERT E. WRIGHT.3 1 University of Illinois, Urbana, IL; 2 Lawrence Livermore National Laboratory,
Livermore, CA; 3 University of Minnesota, Minneapolis, MN. Response
of C3/C4 plants to mid-Holocene aridity in west-central Minnesota.
Paleorecords of ecological response to aridity can offer insights into the
consequences of pervasive drought that will likely accompany future cli-

matic warming in the mid-continent of North America. To examine the

effects of mid-Holocene (MH; ;8.0 to 4.0 ka yr BP) aridity on the prairiewoodland ecosystems of west-central Minnesota we analyzed sediment
cores from West Olaf Lake (WOL; 468 379 N, 968 119 W) and Steel Lake
(SL; 468 589 N, 948 419 W) for d13C of charcoal, charcoal abundance, and
pollen assemblages. Mineral composition (aragonite/calcite ratio) at WOL
and d18O of bulk carbonate at SL suggest that the region experienced pronounced droughts during the MH, and that drought severity decreased with
time. By applying a mixing model to charcoal d13C data we estimate that
the abundance of C4 plants averaged 50% at WOL and 43% at SL during
the MH. At WOL the abundance of C4 plants was negatively correlated
with the aragonite/calcite ratio (r5-0.63). Severe and variable aridity suppressed C4 species in favor of weedy C3 species such as Ambrosia, and
reduced fuel loads and fire importance. As aridity ameliorated, C4 abundance increased (from ;35 to 65%), weedy species decreased, fuel availability increased, and fire became more important (charcoal accumulation
rates increased from ,10 to .60 charcoal pieces cm-2 yr-1). In contrast,
further east at SL C4 abundance showed no correlation with d18O-inferred
aridity. Climatic conditions were less arid at SL than at WOL, which resulted in a higher abundance of woody C3 species such as Quercus, and
lower biomass flammability (,5 charcoal pieces cm-2 yr-1) at SL throughout
the MH. Our results imply that C4 plants may not be adapted to climatic
conditions characterized by high severity and variability of aridity.
NELSON, JENNIFER A. and WILLIAM G. WILSON. Duke University,
Durham, NC. Importance of predator and prey territory size on stability within a Rosezweig-MacArthur framework.
Population dynamics of both predators and prey are affected by predator
foraging behavior, specifically the scale of each species territory relative
to the predators foraging scale. Previous theoretical models of predatorprey interactions, such as the Rosenzweig-MacArthur model, assume that
predator and prey territories are of equal size, and both are equal to the
predator foraging scale. Using deterministic models based on the ubiquitous
Rosenzweig-MacArthur model, we analyzed the effect of a change in the
spatial extent of predator foraging on the predator and prey population
dynamics by allowing a predator forgaging site to overlap multiple prey
sites in our model. We demonstrate an increase in the number of possible
equilibria and changes in the stability of the preexisting ones. We also
demonstrate, interestingly, that there is no change in the predator-prey coexistence region.
NELSON, KAREN,* MARGARET A. PALMER and BROOKE A. HASSETT. University of Maryland, College Park, MD, karen-nelson@erols.com. Stream ecosystem structure and function at multiple
scales: Effects of land use and impervious surface.
An important issue in both theoretical and applied ecology is understanding
the scale at which landscape processes affect ecological response. In this
talk we examine the impacts of land use change on stream ecosystem structure (concentrations of nitrate, ammonia, and phosphate) and function (uptake velocity of ammonia and phosphate). We identify 3 possible scales of
impact - whole watershed, riparian corridor, and in-stream features - and
determine their relative importance. Our dataset includes measurements of
nutrient concentrations (4 seasons per site) and uptake (1 measurement per
site), as well as extensive in-stream habitat and GIS-based land use information, at 71 sites arranged hierarchically within 4 watersheds in Maryland,
USA. We found that concentrations responded most to land use, less so to
riparian status, and least to in-stream features, suggesting that nutrient loading rather than uptake is the dominant factor determining concentrations.
In addition we found that impervious surface and residential/commercial
land use were associated with lowered nutrient concentrations, probably
due to decreased agricultural nutrient inputs and/or availability of metals
to which nutrients may adsorb. Uptake velocities had lower association
with land use, and higher associations with in-stream features, particularly
the presence of fine sediment.
NENZEN, HEDVIG* and CHERYL SWIFT.* Whittier College, Whittier,
CA. Seedling recruitment and seed bank densities across ecotones betwen coastal sage scrub and grassland.
Local patterns of plant community distribution are a function of disturbance
regime, microclimate, soil type and biota, seed dispersal, seedling recruit-

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249

ment and competition. Coastal sage scrub often occurs as patches within a
matrix of native perennial grasses, and non-native invasive annuals and
annual grasses in the Puente Hills in southeastern Los Angeles County. The
matrix of patches of coastal sage scrub and native grasses is typical of
undisturbed coastal sage scrub as is the matrix of coastal sage scrub and
non-native annuals; however, the factors responsible for the maintenance
of the boundaries between these patches are not well understood. This study
compared the ability of shrubs, perennial grasses, and non-native annuals
to invade across shrub-native grassland and shrub-invasive grassland ecotones. Coastal sage scrub and native grasses rarely penetrated more than 1
m across the ecotone; however, invasive grasses and annuals penetrated
more than 5 m into coastal sage scrub. The ability of invasive grasses and
annuals to invade coastal sage scrub is correlated with the amount of bare
ground present within the coastal sage scrub patch. The results of this study
suggest that in the absence of invasive, the large-scale disturbance shifted
the boundary between coastal sage scrub and grassland. However, when
invasive annuals are present, small disturbances leading to patches of bare
ground within the larger coastal sage scrub patch are easily invaded by
invasive annuals leading to fine scale fragmentation within the coastal sage
scrub patch. These results also suggest that invasive annuals produce seeds
that disperse farther in higher numbers than native shrubs or grasses. This
is confirmed by an examination of the seed banks across ecotones since
invasive seeds penetrate at least 15 m into coastal sage scrub patches.
NESS, JOSHUA H.,1,* JUDITH L. BRONSTEIN,1 ALAN N. ANDERSEN2 and J. N. HOLLAND.1 1 University of Arizona, Tucson, Arizona;
2
CSIRO Tropical Ecosystem Research Centre, Winnellie, NT, Australia.
Disruption of native ant/seed interactions by invasive ants is predicted
by body size.
Ecologists have long sought to link the traits of individual species to their
influences on communities and ecosystems. This challenge becomes more
urgent as an increasing array of exotic species disrupts processes within
invaded communities. We demonstrate that the mean and maximum distances that ants transport seeds adapted for ant dispersal increase exponentially with ant body size. These patterns occur in three of the four largest
ant subfamilies (Dolichoderinae, Formicinae, and Myrmicinae, but not Ponerinae). Ant body size is a useful predictor because it incorporates many
of the variables known to influence seed dispersal distance, including worker-to-seed mass ratio, group vs. solitary foraging, foraging range, ant nest
distribution, as well as the likelihoods that ants will collect seeds or drop
them in mid-transport. We suggest that many of the benefits plants receive
from this mutualism increase with dispersal distance. Such benefits include
reduced competition between related plants, patch colonization, access to
different microhabitats, and escape from high density patches where seeds
are vulnerable to density-dependent natural enemies (e.g. small mammals
and pathogens). Invasive ants are smaller than most seed dispersing ants,
and seed dispersal distances are reduced in sites occupied by invasive ants,
relative to sites with other ants. The link between ant body size and seed
dispersal distance, combined with the dominance of invaded communities
by these typically small ants, provides a simple, mechanistic explanation
for the disruption of native ant-seed dispersal mutualisms in habitats invaded by exotic ants.
NEUBAUER, SCOTT C.,1,* PATRICK MEGONIGAL1 and DAVID EMERSON.2 1 Smithsonian Environmental Research Center, Edgewater, MD,
USA; 2 American Type Culture Collection, Manassas, VA, USA. Microbial iron oxidation in the wetland plant rhizosphere.
The formation of iron plaque on the roots of wetland plants can occur
when oxygen leaking from plant roots oxidizes soluble ferrous iron in
wetland porewaters. Although the chemical oxidation of Fe(II) proceeds
very rapidly at circumneutral pH, lithotrophic Fe(II) oxidizing bacteria
(FeOB) are common on the roots of many wetland plants and can be a
significant fraction of the total root-associated microbial community, indicating that they may play a role in rhizosphere Fe(II) oxidation. To assess
the significance of microbial iron oxidation, a series of laboratory experiments was performed using a purified strain of FeOB (strain BrT) that was
isolated from the roots of Typha latifolia. The first set of lab experiments
used batch cultures of strain BrT grown in bioreactors under tightly controlled pH, O2, and Fe(II) supply conditions. A second set of experiments

250

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examined iron oxidation in hydroponic microcosms that contained surfacesterilized roots of live wetland plants that were subsequently inoculated
with live BrT cells or autoclaved iron oxides. In the microcosm experiments, the only oxygen source to the hydroponic solution was radial oxygen loss from the plant roots. In both sets of experiments, rates of iron
oxidation were measured by monitoring the disappearance of dissolved
Fe(II) over periods of hours to days. Results from these experiments indicate that iron oxidizing bacteria can accelerate rates of Fe(II) oxidation
and may account for over 50% of total Fe(II) oxidation. Additionally, the
FeOB were able to successfully compete with chemical oxidation for Fe(II)
(i.e. grow), despite evidence that total Fe(II) oxidation was limited by the
Fe(II) supply (bioreactor experiments) or the rate of Fe(II) diffusion to the
root surface (microcosm experiments). Taken together, these experiments
indicate that iron oxidizing bacteria are likely to contribute to iron plaque
formation in the rhizosphere.
NEVAI, ANDREW L.* and RICHARD R. VANCE. University of California at Los Angeles, Los Angeles, California, USA. Competition for
light between plant species with different foliage profiles: Analysis.
Two coexisting plant species that deploy their foliage at different heights
above the ground experience contrasting benefits and costs. By placing its
leaves high above the ground, the taller species intercepts full sunlight that
produces rapid photosynthesis, but at the cost of growing and then supplying maintenance metabolic energy for tall stems. The shorter species, by
contrast, requires less energy for stem growth and maintenance, but its
leaves intercept less light and consequently experience slower photosynthesis. Can this height difference by itself make stable coexistence of these
competing species possible? How does extensive overlap in foliage profiles
affect the outcome of competition? To find out, we examined a two-species
Kolmogorov competition model whose specific growth rate functions incorporate realistic descriptions of plant structure and function. The differential equations do not yield explicit equations for the two species nullclines, but implicit arguments reveal nullcline placement and hence the outcome of competition. Depending upon the magnitude of the species height
differences and the amount of vertical foliage overlap, four outcomes are
possible: the tall species persists alone, the short species persists alone,
neither species persists, or the two species persist together at stable equilibrium abundances. This last result appears to occur throughout a large
portion of parameter space. Founder-controlled competitive exclusion is not
possible. These findings establish that, at least in principle, a difference in
foliage height can promote stable competitive coexistence of plant species
that differ in no other way.
NEWBOLD, T. A. SCOTT* and JAMES A. MACMAHON. Utah State
University, Logan, UT. Prey specificity and diet selection in a variable
environment: Desert horned lizards (Phrynosoma platyrhinos) and ants.
Diet selection is influenced by a complex set of factors that include prey
availability, prey size, and prey defenses, among others. In conjunction with
a larger study assessing horned lizard habitat associations, we examined
the relationship between availability and selection of ants by the desert
horned lizard, Phrynosoma platyrhinos, to determine: 1) the nature of
horned lizard preference for harvester ants (e.g., generalist vs. specialist),
and; 2) how spatial and temporal variability of ant abundance affects lizard
prey preference. Ant species composition and abundance and lizard prey
preference were determined using pitfall sampling and lizard scat dissection
on 80 plots established along a 12km transect in western Box Elder Co.,
Utah, U.S.A. Ant availability varied with location (soil type and shrub
community) and, to a lesser extent, season (late spring through late summer). Across the entire site, 14 of 24 ant species were represented in the
lizard diet. However, 86% of that diet consisted of just 4 ant species (n563
scat). Other insects, mostly beetles, comprised less than 5% of scat items.
In contrast to previous studies (e.g., Rissing 1981, Collins 1988; but see
Suarez et al. 2000), lizards did not show a disproportionate preference for
harvester ants. Instead, larger ants (.5mm; including harvester ants) were
preferentially selected over medium-sized ants (2-5mm) and smaller ants
(,2mm) (73% vs. 26% vs. 1%, respectively). Of the medium-sized ants
selected, species that were particularly abundant comprised larger proportions of the diet than less abundant species. Also, while diet composition
varied considerably based on spatial location, diet selection on plots mon-

itored through time remained relatively constant. Our results suggest that
desert horned lizards in our area should be considered opportunistic foragers rather than harvester ant specialists. Moreover, ant body size and ant
abundance largely determine desert horned lizard diet choice.
NEWINGHAM, BETH A.,1,* JORGE M. VIVANCO,2 HORMOZ BASSIRIRAD,3 WILLIAM GOOD4 and RAGAN M. CALLAWAY.4 1 US Geological Survey, Moab, UT, USA; 2 Colorado State University, Fort Collins,
CO, USA; 3 University of Illinois at Chicago, Chicago, IL, USA; 4 University of Montana, Missoula, MT, USA. Root herbivory affects nitrogen
uptake and secondary chemical production in the invasive plant, Centaurea maculsa.
Centaurea maculosa is one of the most invasive plants in the northwestern
United States. Previous research has shown that C. maculosa compensates
for root herbivory by biological controls, which sometimes results in increased negative effects on native grasses. We investigated the effects of
two root-boring biocontrol species on C. maculosa, and tested two mechanisms by which C. maculosa may increase its competitive ability. In neither experiment did herbivory affect the growth of C. maculosa. In our
first experiment, we tested the hypothesis that C. maculosa increases nitrogen uptake after herbivory by Agapeta zoegana. In a 15N labelled experiment, we examined the effects of herbivory on the 15N content of the roots
and shoots of C. maculosa. Root herbivory decreased 15N in roots, but did
not affect 15N in shoots. Since root herbivory decreased 15N, this experiment
indicates that nitrogen uptake is not responsible for the increased competitive ability of C. maculosa experiencing herbivory. Studies have shown
that C. maculosa produces a secondary chemical, (-)-catechin, which has
powerful allelopathic effects on other plant species. In a greenhouse experiment, we subjected C. maculosa to shoot herbivory by Trichoplusia ni
or root herbivory by Cyphocleonus achates and measured the amount of
(-)-catechin produced by C. maculosa. Shoot herbivory had no effect on
(-)-catechin production; however, root herbivory increased (-)-catechin production four-fold compared to uneaten and shoot damaged plants. Similar
results were demonstrated in C. maculosa plants transplanted outdoors.
These results suggest that root herbivory may increase the allelopathic effects of C. maculosa, and suggest caution when using some insect species
as biocontrols to control C. maculosa.
NEWMAN, GREGORY S.,1,* STEPHEN C. HART,1 DANIEL R. GUIDO1
and STEVEN T. OVERBY.2 1 School of Forestry and Merriam-Powell Center for Environmental Research, Northern Arizona University, Flagstaff,
AZ; 2 Rocky Mountain Research Station, USFS, Flagstaff, AZ. Wildfire
effects on soil microbial activity and community-level physiological
profiles in a ponderosa pine ecosystem.
Forests of the western United States have received increasing incidence
and extent of wildfire during the past decade as a result of increased tree
densities and buildup of forest floor fuels due to fire suppression policies.
Evidence of the direct effects of wildfire on soil microbial activity and
composition is scarce and these effects may have a greater influence than
prescribed burns due to the greater intensity of disturbance. Ponderosa pine
plots in New Mexico associated with the national Fire - Fire Surrogate
network were burned by wildfire in September 2002 following pretreatment
measurements and prior to the initiation of treatments. We resampled these
plots one month following the wildfire and one year following pretreatment
measurements, removing any seasonal influence on soil activity. Wildfire
homogenized bacterial community-level physiological profiles (CLPPs) and
had the opposite effect on fungal populations. The bacterial CLPP plate
activity doubled following wildfire while there was no effect on fungal
plate activity. Conversely, basal and substrate-induced respiration were substantially lower than in a nearby unburned stand which was similar to the
wildfire stand based on pretreatment measurements. Although b-glucosidase activity declined following wildfire, galactase, xylosidase, cellobiohydrolase, alkaline phosphatase, and sulfatase activites all increased significantly, consistent with the amplified bacterial metabolism observed in
CLPPs. Wildfire and subsequent microbial activity resulted in 23x increase
in net N mineralization, an 80% decline in net nitrification, and nearly 2x
and 19x increases in soil available NO3- and NH4+ pools, respectively.
Altered soil microclimate and organic matter transformations, as well as
the removal of plant competition, following wildfire create a more favor-

able environment for microbial populations, which may facilitate the regeneration of forest stands through increased mineralization of plant available nutrients.
NGAI, JACKIE* and ROBERT L. JEFFERIES. University of Toronto,
Toronto, Ontario, Canada. Contrasting nutrient limitations in arctic
coastal marshes: Implications for goose foraging.
Marshes grazed by snow geese on the Hudson Bay coast were examined
to determine whether a switch from nitrogen- to phosphorus-limited plant
growth occurs between the salt marshes and adjacent freshwater marshes.
Experimental plots, which were fertilized with N and P in a factorial design,
were set up in both a salt and freshwater marsh, and plant biomass and
tissue nutrient content responses to the fertilization treatments were measured. In the salt marsh, it was found that, when added alone, N and P had
non-significant effects on aboveground biomass. However, there was a significant increase in biomass when the nutrients were added together, with
nitrogen being the most important element determining the response. In
contrast, in the freshwater marsh, the addition of N did not significantly
increase aboveground biomass, although there was a significant effect of P
additions. No NP interaction was observed in the freshwater site, while in
the salt marsh, P does become limiting when N availability is supplemented. N:P stoichiometry also indicates that plant growth in the salt marsh
is mainly N-limited, while that in the freshwater marsh is P-limited. In
recent decades, destructive feeding by geese has destroyed much of the salt
marsh, and the geese have been forced to forage in the freshwater site.
This difference in nutrient limitation between the salt and freshwater marshes means that vegetation available for geese during the post-hatch period
is now P- rather than N-limited. The potential implications of this change
in forage quality will be assessed in terms of the observed decline in gosling growth.
NGUGI, MOFFATT K.* and RICHARD T. CONANT. Colorado State University, Natural Resources Ecology Lab., Fort Collins, Colorado, U.S.A.
Remote Sensing of grazing intensity: Case studies in the short- and
midgrass steppes using MODIS data.
Improved grassland management that leads to increased production or rehabilitation of degraded lands may lead to C storage in soils. Without
knowledge about how various grasslands are managed, however, the potential for influencing soil C dynamics cannot be well quantified. Remote
sensing enables collection of land characterization data that are spatially
explicit, broad in extent, and repeatable. One goal of this project is to
develop techniques to characterize grassland management using remote
sensing data from EOS-MODIS and LANDSAT. This work is focused on
detection and assessment of grazing management at Central Plains Experimental Range (CPER, in eastern Colorado), which has grazing of varying
intensity, and at Crescent Lakes National Wildlife Refuge (CLNWR, in
central Nebraska) where recent history of grazing varies between different
paddocks. Ground-truth measurements of aboveground net primary productivity (ANPP) and leaf area index (LAI) were used to validate estimates
derived from remotely sensed surface reflectance. Ground-based results indicate that ANPP and LAI are strongly related during the wetter part of
the growing season, but not during drier periods. At CPER, remotely sensed
average LAI and the seasonal distribution of LAI can be used to distinguish
heavily grazed plots from ungrazed plots, while at CLNWR, those paddocks that are regrowing are discernible from those that are grazed using
remote sensing. Overall, our results suggest that remotely sensed data can
be used to characterize management impacts at divergent sites across the
Great Plains and this information is likely to be useful for driving ecosystem models to assess management impacts on soil C stocks.
NIEMI, GERALD J.,1 NICHOLAS P. DANZ,1,* RONALD R. REGAL,1,2
TOM HOLLENHORST,1 VALERIE BRADY,1 LUCINDA B. JOHNSON,1
TERRY BROWN1 and GEORGE HOST.1 1 University of Minnesota Duluth, Duluth, MN; 2 . Sampling design for linking stress with response
in the Great Lakes coastal ecosystem.
The goal of our work is to develop indicators that both estimate ecological
condition and suggest plausible causes of ecosystem degradation. Thus, the
indicators developed must be based on their responsiveness to anthropo-

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251

genic stress. To evaluate the association of potential indicators with stress,

it is necessary that the sample be distributed across the stress gradients of
interest. Although there are many examples of sampling designs for the
goal of estimating mean condition such as in monitoring programs, a sampling framework has been lacking for relating stress to ecological response.
Our project consists of five individual subcomponents to investigate different types of biological responses (e.g. birds, diatoms) with different sampling methodologies and sample size requirements. Previous work in the
Great Lakes coastal region has identified many influential anthropogenic
stresses. The major challenges for our sampling design were to distribute
the sample across many stress gradients and to allow for integration of
indicators from individual project subcomponents by maximizing site overlap. We used a GIS to compile over 200 data layers representing six types
of anthropogenic stress: agriculture, atmospheric deposition, land cover,
human populations, point source pollution, and shoreline modification. We
divided the coastal region into 762 units using geomorphological criteria,
and calculated stresses for each unit. We used principal components analysis to remove redundancy within the stress categories and cluster analysis
was used with pc scores as input to create groups of coastal units having
similar stress profiles. Random selection of sites from the clusters provided
an unbiased sample of sites that spanned the range of stress conditions in
the region and provided a high degree of overlap for project subcomponents. Sampling of the sites is being completed in 2002 and 2003.
NIESENBAUM, RICHARD A.* and EMILY C. KLUGER. Muhlenberg
College, Allentown, PA. Where is spicebush the spiciest? Variation
in leaf quality, chemistry, and herbivory in Lindera benzoin.
The aim of this study has been to assess how rates of herbivory vary with
light environment in populations of Lindera benzoin (spicebush). Specifically we have focused on how plant growth rates, leaf toughness, plant
chemistry and nutrition, and insect biology and ecology vary with light
environment and influence these rates. Over the entire season, plants from
shade locations experienced greater levels of herbivory than plants in sun
locations while the abundance of herbivores did not differ across habitats.
Shorter term feeding trials with the lepidopteron herbivore Epimecis hortaria also showed that leaves in the shade are consumed at a greater rate,
and that younger leaves are preferentially eaten. Also, herbivores experienced greater rates of weight gain when feeding on leaves in the sun.
Leaves originating from high light environments were significantly tougher
and had a stronger fragrance (and perhaps more aromatic compounds) than
leaves from low light environments. Gas chromatography spectra and solid
phase micro-extraction revealed that plants in high light environments had
a greater concentration of lower molecular weight compounds while shade
plants had a greater relative abundance of heavier compounds. Plants in
the shade had significantly more nitrogen per unit weight of leaf than plants
in the sun. The relationships between leaf age, light environment, leaf defense, and fine scale variation in rates of herbivory are considered.
NILES, JOHN O.,1,2,* CLAIRE KREMEN3 and JOHN P. FAY.4 1 Climate
and Biodiversity Alliance, Conservation International, 1919 M St, NW,
Suite 600, Washington, DC; 2 University of California, Berkeley, Energy
and Resources Group, 310 Barrows Hall, Berkeley, CA; 3 Prinecton University, Ecology and Evolutionary Biology, Guyot Hall, Washington RD,
Princeton, NJ; 4 Stanford University, Center for Conservation Biology, Gilbert Hall, Stanford, CA. A tropical forest meta-analysis: Ecological and
conservation findings.
We conducted a meta-analysis of ecological and socio-economic parameters
within the tropical forest biome. We built and analyzed a GIS database
using layers describing biological importance, carbon, threat, impact, ecosystem type, and historical project implementation by country. The goal of
our research was to demonstrate broad ecological patterns, such as the
relationship between perceived biological value and carbon across the tropical forest landscape. We also revealed important conservation findings
such as: 1) areas of congruence and discrepancy within global carbon and
biological-importance models, 2) specific areas of robust conservation importance and the estimated above ground carbon they contain, 3) areas of
consensus threat, and 4) important differences by country of historical project implementation rates. We confirmed our hypothesis that high-impacted
areas have lower carbon than low-impact areas. We also demonstrated that

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Abstracts

the consensus areas of threat do not have signifcantly different carbon

values than areas not imminently threatened. For conservation, we identified almost 7 million hectares of tropical forest that: 1) are in countries
with relatively high project implementation success rates, are considered
consensus areas of high biological importance, have relatively low impact
and are threatened. These areas contain almost 1.5 billion tons of carbon
and should be global priorities for conservation. Conservation policy recommendations are discussed based on these new understandings of the
ecology, history and social context of the tropical forest biome.
NIPPERT, JESSE B.,1,* ALAN K. KNAPP,1 MELINDA D. SMITH2 and
JOHN M. BRIGGS.3 1 Division of Biology, Kansas State University, Manhattan, KS, USA; 2 National Center for Ecological Analysis and Synthesis,
735 State St., Suite 300, Santa Barbara, CA, USA; 3 Department of Plant
Biology, Arizona State University, Tempe, AZ, USA. The influence of
precipitation amount vs. pattern on grassland net primary production.
Both rainfall amount and variability (seasonal timing and event size) have
been shown to influence grassland aboveground net primary productivity
(ANPP) in controlled field experiments. Using long-term data sets from the
Konza Prairie LTER program, we analyzed patterns of natural variability
in precipitation (event size, timing, event variability and length of intraevent dry periods) to assess their influence on ANPP in a tallgrass prairie
ecosystem. Data were analyzed from an annually burned watershed comprising both upland and lowland plots and an experimentally irrigated site.
Our results showed that seasonal precipitation amount was the best predictor of total and grass ANPP in three of the four long-term datasets analyzed
(r2 $ 0.656). In the lowland burned watershed site, there were no significant
correlations with any of the variables measured. Significant relationships
were found between total and grass ANPP and several precipitation timing
and event size variables, but these relationships were each weaker than for
precipitation amount alone (r2 # 0.664). No significant relationships were
found between forb productivity and any of the precipitation variables measured (P . 0.05). These results imply that while precipitation variability
can influence total and grass ANPP in tallgrass prairie under controlled
conditions, variation in other abiotic and biotic factors in these long-term
experiments makes it difficult to detect the impact of variability in precipitation patterns under natural conditions.
NORBY, RICHARD J.,1,* SARA S. JAWDY,1 JOANNE LEDFORD,1
JOHNNA D. SHOLTIS2 and DALE W. JOHNSON.3 1 Oak Ridge National
Laboratory, Oak Ridge, TN; 2 Texas Tech University, Lubbock, TX; 3 University of Nevada-Reno, Reno, NV. Increased nitrogen uptake supports
higher NPP but not carbon storage in CO2-enriched sweetgum trees.
The free-air CO2 enrichment study in a Liquidambar styraciflua (sweetgum) plantation on the Oak Ridge National Environmental Research Park
provides the opportunity to evaluate effects of elevated atmospheric CO2
on nitrogen cycling in an established forest ecosystem in which the N cycle
is dominated by recycling. Net primary productivity (NPP) in this stand
has increased 24% in response to CO2 enrichment over 5 years, a consistent
response attributable to a sustained increase in photosynthesis and despite
an 8% decline in foliar N concentration. Does this increase in photosynthetic nitrogen-use efficiency imply a negative feedback on organic matter
production because of the requirement for C and N within relatively constrained proportions? A stand-level analysis of C and N budgets sheds light
on this question. Although elevated CO2 has caused reduced N concentration in both green leaves and leaf litter, there has been no effect of elevated
CO2 on whole-canopy N content or the amount of N returned to soil in
leaf litter. NPP increased linearly with N uptake, and N uptake was significantly higher in CO2-enriched plots. N uptake increased linearly with increasing fine-root length duration, which also was significantly higher in
elevated CO2. However, most of the additional N taken up by trees in CO2enriched stands, as well as most of the additional NPP, was used in fineroot production rather than being invested in increased photosynthetic capacity (leaf area duration did not increase) or production of wood (no
significant increase after the first year). Hence, there has not been a sustained increase in C or N sequestration in the CO2-enriched trees compared
to trees in ambient air. There also has been no indication that the decomposition rate of leaf or fine root litter has been altered by elevated CO2,
nor has there been any detectable changes in soil microbial N cycling, so

the system-level response of accelerated carbon and nitrogen cycling should

be sustainable.
NORRIS, MARK D.,1 PETER B. REICH,1 SARAH E. HOBBIE2 and DAVID A. WEDIN.3 1 Department of Forest Resources, University of Minnesota, St. Paul, MN, USA; 2 Department of Ecology, Evolution and Behavior, University of Minnesota, St. Paul, MN, USA; 3 School of Natural
Resource Sciences, University of Nebraska, Lincoln, NE, USA. Soil nutrient dynamics across a fire frequency gradient in oak savanna.
Savanna ecosystems provide an opportunity to examine the dynamic competitive balance between two life forms, an herbaceous understory and
woody overstory, and how this balance is influenced by global change.
This research evaluates species effects on ecosystem functions and the
mechanisms controlling observed patterns of nitrogen cycling in a midwestern oak savanna. The fire frequency gradient (0-0.8 fires per year) at
Cedar Creek LTER, Minnesota, drives a gradient in oak canopy dominance
which in turn leads to substantial differences in N cycling rates. Available
soil N pools and cycling rates from multiple years are negatively correlated
with fire frequency as annual net N mineralization rates in savanna plots
of low fire frequency (high oak dominance) are 3 times that of high fire
frequency (high grass dominance) plots (60 vs. 20 kg N/ha/yr). A shortterm lab incubation that standardized microclimatic conditions found that
this pattern of increased N mineralization is likely due in large part to
increased microbial biomass and activity as fire frequency decreases. An
ongoing reciprocal transplant decomposition experiment (oak vs. grass litter
in oak vs. grass dominant plots) suggests that decay rates of aboveground
material do not vary greatly after two seasons with respect to fire frequency,
however belowground patterns indicate that litter decomposes quicker in
its native habitat. Results thus far indicate that short-term patterns of N
cycling are closely linked to microbial activity irrespective of plant community composition and fire effects.
NORRIS, RYAN D.,1,* PETER P. MARRA,2 KURT T. KYSER,3 THOMAS
W. SHERRY4 and LAURENE M. RATCLIFFE.1 1 Department of Biology,
Queens University, Kingston, ON, Canada; 2 Smithsonian Environmental
Research Center, Edgewater, MD, United States; 3 Department of Geological Sciences, Queens University, Kingston, ON, Canada; 4 Department of
Ecology and Evolutionary Biology, Tulane University, New Orleans, LA,
United States. Tropical winter habitat limits reproductive success on
the temperate breeding grounds in a migratory bird.
Isolating the factors that control population dynamics in migratory animals
has been constrained by our inability to track individuals throughout the
annual cycle. Using stable-carbon isotopes, we show for the first time that
the reproductive success of a long-distance migratory bird is influenced by
habitat quality thousands of kilometers away on tropical wintering grounds.
For male American redstarts (Setophaga ruticilla), winter habitat quality
influences arrival date on the breeding grounds, which in turn affects key
variables associated with reproduction, including number of young fledged.
Females occupying high quality winter habitat produce up to two additional
young and fledge offspring up to a month earlier compared to females
wintering in poor quality habitat. Differences of this magnitude are highly
significant considering redstarts are single brooded, lay clutches of only
three to five eggs and spend only two and a half months on the breeding
grounds. Results from this study provide fundamental evidence for the
importance of understanding how periods of the annual cycle interact for
migratory animals. Importantly, continued loss of tropical wintering habitat
could have negative effects on migratory populations the following breeding season, minimizing density-dependent "buffer" effects on the breeding
grounds and leading to further population declines. If conservation efforts
are to be successful, strategies must, therefore, incorporate measures to
protect habitat throughout the entire annual cycle in migratory animals.
NORTON, ANDREW P.,1,* GREG ENGLISH-LOEB2 and STEVE HESLER.2 1 Colorado State University, Bioagricultural Sciences and Pest Management, Fort Collins, CO; 2 Cornell University, NYSAES, Department of
Entomology, Geneva, NY. Assessing interaction strengths in a plant arthropod - plant pathogen interaction web: Do factorial manipulations predict real world patterns?
Determining interaction strengths in multi-trophic systems has been a challenge for applied and basic ecologists for decades. In multi-trophic systems

both indirect effects (i.e. species A alters the density of species B which
in turn alters the density of species C) and interaction modifications (i.e.
species A alters the per-capita impact of B on C) may be common. Factorial
experimental manipulations and analysis with a combination of ANOVA
and path analysis are powerful tools for the examination of interaction
webs. However, practical considerations often limit the spatial and temporal
scale over which we are able to manipulate organism abundance, potentially limiting the predictive ability of experimental results. We used a
combination of three years of short term factorial manipulation experiments
with a large scale, multiple year field trial to examine interaction webs in
a plant - mite - plant pathogen system. The riverbank grape, Vitis riparia,
possesses acarodomatia which separately increase the densities of two taxa
of beneficial mites: the mycophagous Orthotydeus lambi and the predaceous Amblyseius andersoni. These mites reduce the abundance of the plant
pathogen grape powdery mildew and plant feeding spider mites respectively. However, these two mites negatively effect each other, potentially
limiting the value of acarodomatia to the plant. ANOVA and path analysis
of factorial manipulations of acarodomatia and the mites indicate that indirect effects are common in this system but that there is little evidence
for interaction modification. Further, interaction strengths in this web were
largely consistent between the different years the experiments were run.
Comparison of these results with a large scale multi-year field trial indicates
that the results from factorial manipulations are a good predictor of organism abundances at a much larger scale.
NOVOTNY, AMY M.,1,* JOHN D. SCHADE,1,2,* ADAM KAY,2 DANIEL
HERNANDEZ,2 MEGAN OGDAHL2 and SARAH E. HOBBIE.2 1 Arizona
State University, Tempe, Arizona; 2 University of Minnesota, St. Paul, Minnesota. Changes in plant elemental ratios in response to increased CO2
and nitrogen enrichment.
Human activities have resulted in increased nitrogen deposition and atmospheric CO2 concentrations, potentially causing significant changes in
nutrient cycling rates in human-dominated ecosystems. Specifically, increased nitrogen and carbon availability may lead to phosphorus limitation
of key ecosystem processes. We tested this hypothesis using a fully-crossed
CO2 and N enrichment experiment (BioCON at Cedar Creek Natural History Area, Minnesota), focusing on variation in tissue chemistry of herbaceous plant species. Here we present results from monocultures of two
species, Solidago rigida and Amorpha canescens, a leguminous forb. We
predicted higher C:P and N:P ratios in plants receiving elevated CO2 and
N deposition, indicating that P has become the limiting resource. We found
significantly higher C:P ratios in Solidago plots receiving both C and N
enrichment than in all other treatments. Monocultures of Amorpha showed
lower C:P in all C enriched plots and showed no response to N additions.
In both species, N:P decreased under C enrichment. In Solidago plots, N:
P increased under N enrichment, while the effects of simultaneous enrichment with C and N were additive. When combined with recent data showing a significant influence of changes in plant C:P and N:P on insect herbivores, these data suggest the potential for an important influence of human activities on food web dynamics mediated by changes in food quality.
NUFIO, CESAR R.1,2,* and DANIEL R. PAPAJ.2 1 Natural History Museum, University of Colorado, Boulder, CO; 2 University of Arizona, Tucson, AZ. Parent-offspring conflict in the walnut fly, Rhagoletis juglandis.
Choosing where offspring will develop is especially important for insects
whose larval stages are restricted to a particular host resource. In such
insects, maternal egg laying decisions may not only involve choosing optimal hosts based on their intrinsic qualities but also avoiding hosts occupied by conspecific brood. In many insects, the ability to discriminate between previously exploited and unexploited hosts is often mediated by the
use of a marking pheromone. Despite engaging in what appears to be hostmarking behavior, the walnut fly Rhagoletis juglandis prefers to deposit
clutches into previously exploited hosts. In this study, we quantified host
reuse in R. juglandis and assessed its impacts on offspring fitness. We also
explored the role that marking pheromone plays in determining the level
to which hosts are reused. Host reuse by the walnut fly was common in
the field, where trees were synchronously infested over a 14 to 17 day
period. Field and laboratory experiments showed that increases in larval

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253

densities within fruit reduced larval survival and pupal weight, the latter
being strongly correlated with the number of eggs a female produced over
her lifetime. In field-cage experiments, fruit that were host marked by females for longer durations were less acceptable to other females. Moreover,
the duration of time that a female marked a fruit was positively correlated
with the size of her clutch. These results indicate that, while females commonly reuse fruit, they nevertheless signal the level of larval competition
associated with a fruit and adjust allocation of eggs to fruit accordingly.
Host reuse by R. juglandis appears to reflect a strategy by which females
attempt to maximize the number of clutches they deposit over a lifetime
at the expense of the per capita fitness of her brood.
NUTTER, DOUGLAS A.* University of Chicago, Chicago, IL. Community development and structure in natural mesocosms: A multitrophic approach.
Although the relationship between sampling area and species richness is
well studied, only recently has trophic structure been incorporated. Organisms of different trophic rank have varying resource requirements, and
community structure should therefore differ systematically as a function of
area. In the rocky intertidal, tidepools are an ideal system in which to test
this theory. They are discrete entities with little between-pool migration,
and yet they are truly open systems. I am using a system of artificial
tidepools to test predictions of the rate and nature of community development and trophic structure. Tidepools provide a practical test of current
theory, relaxing the assumptions of weak top-down effects, closed systems,
trophic specialization, and equilibrium conditions, and allowing determination of those assumptions that are most important. Data suggest that 1)
overall rates of species addition increase with area, 2) sensitivity to area
increases with trophic status, and 3) maximum rate of species accumulation
occurs earlier in community development for resources than for consumers.
These patterns may be highly context-dependent and strongly influenced
by natural variability in interannual recruitment, however. Recruitment variability acted to ameliorate otherwise clear patterns in trophic-group community assembly.
NUTTLE, TIM1,2 and JAMES W. HAEFNER.2 1 Friedrich Schiller University, Jena, Germany; 2 Utah State University, Logan, Utah, USA. A mechanistic model for wind-dispersed seeds in heterogeneous environments:
Design and validation.
Seed dispersal is an important determinant of local and regional community
composition. We present a mechanistic model of seed dispersal by wind
that incorporates heterogeneous vegetation structure within the model landscape. Such vegetation heterogeneity is important because it affects the
horizontal wind profile, which is one of the primary determinants of seed
dispersal distance. The model was developed for hardwood bottomland
forests in the Mississippi Alluvial Valley, but is applicable to any forest
community for which the requisite data are available. The model moves
seeds in 3-dimensional space from the parent tree until it reaches the
ground, by combining local wind speed and seed terminal fall velocity. The
model differs from other mechanistic seed dispersal models in that vegetation, and thus wind speed, may vary along the trajectory of each seed.
The model was validated using source populations of trees in forests and
seed trap data in adjacent reforestation sites of two distinct vegetation structures (younger, open reforestation sites and older, closed reforestation sites).
The model was replicatively and predictively valid for all species and vegetation types, even though patterns of seed density differed greatly between
the two vegetation types. The models sensitivity to vegetation structure,
and its ability to accurately predict seed arrival when this structure is incorporated, demonstrates its utility for modeling seed dispersal in heterogeneous environments, and its potential utility for incorporating mechanistic seed dispersal in a forest simulation model.
NYDICK, KOREN,1,* CHRIS ARP,2 MICHELLE BAKER,2 BOB HALL3
and WAYNE WURTSBAUGH.1 1 Aquatic, Watershed, and Earth Resources
Dept., Utah State University, Logan, UT, USA; 2 Biology Dept., Utah State
University, Logan, UT, USA; 3 Biology Dept., University of Wyoming, Laramie, WY, USA. Comparative retention of nitrogen in stream-lake linkages during snowmelt.
Aquatic ecosystems are important landscape features because of their ability to transform and retain inorganic nutrients such as nitrate. In many

254

Abstracts

landscapes, lakes intersect streams, and this hydrologic connectivity may

alter watershed capacity for nutrient uptake. As a part of a study to understand effects of stream-lake interactions on catchment biogeochemistry,
we performed a two-week release of 15Nnitrate and bromide during snowmelt to the inflow stream of Bull Trout Lake, Idaho. After two weeks we
accounted for 89% of the 15N. During this period of high discharge and
low temperature, the inflow stream benthos and seston took up only 3% of
the 15N. An additional 29% was removed from the inflow by floodplain
exchange. This water, but not 15N, returned to the stream 1-4 weeks later,
suggesting substantial nitrate storage in the floodplain. Lake seston, epiphytes, and zooplankton accounted respectively for 24%, 8%, and ,1% of
added 15N. Despite sedimentation of an additional 8% to the benthos, 15N
increments in the lake sediment were too dilute to measure. Lake nitrate
accounted for 7% of the 15N, but no 15N-nitrate left the outflow and only
6% was transported out of the lake as seston. Thus, the lake and flood
plain wetland, but not the stream channel, were effective in retaining nitrate
during snowmelt.
OCONNELL, LISA,1,* KERMIT RITLAND1 and JOHN RUSSELL.2
1
University of British Columbia, Vancouver, British Columbia; 2 British
Columbia Ministry of Forest, Mesachie Lake, British Columbia. Factors
affecting the mating system of an inbreeding conifer, Thuja plicata (Cupressaceae).
Conifers are predominantly outcrossing, but in a review of the literature
25% of 52 species showed selfing rates above 20%. For a conifer, western
redcedar (Thuja plicata) shows high self-fertility, and population outcrossing rates estimated using one isozyme locus varied significantly among six
natural populations (weighted mean 5 71.5%). We used microsatellite assays of bulked DNA to obtain fine-scale estimates of outcrossing and study
the variation in selfing within and among trees. Outcrossing rates did not
vary significantly among crown positions within trees, but decreased with
tree height in all four populations studied. Taller trees probably receive
higher proportions of self-pollen because they contribute a larger proportion
of self-pollen to the surrounding pollen cloud. The occurrence of multiple
embryos within the same ovule (polyembryony) is a mechanism in conifers
that can potentially increase the proportion viable and/or outcrossed seeds.
Using controlled crosses, we found evidence that with high self-pollination
rates, embryo competition can decrease the proportion of selfed seeds.
However, pre-pollination mechanisms and the degree of self-fertility are
more important than embryo competition in determining selfing rates.
OETTER, DOUG R.,1,* OLGA N. KRANKINA,2 WARREN B. COHEN3
and THOMAS K. MAIERSPERGER.2 1 Georgia College & State University, doetter@gcsu.edu, Milledgeville, GA; 2 Oregon State University, Corvallis, OR; 3 USDA Forest Service, Pacific Northwest Research Station,
Corvallis, OR. Estimating carbon storage for the boreal forest of St.
Petersburg, Russia using satellite imagery.
The boreal forests of northwest Russia play a critical role in sequestering
global atmospheric carbon. Estimates of their current and potential carbon
storage are important for modeling worldwide carbon sinks and sources.
Using detailed forest inventory data in conjunction with multiple-date
Landsat satellite imagery, we predicted regional forest age and aboveground biomass for Russias 76,860 km2 St. Petersburg Region. The remote
sensing analysis included multiple-image rectification and normalization,
creation of a land cover map, and application of advanced regression techniques to estimate forest characteristics. Model development and map error
assessment were based on field measurement of over 1500 forest ground
plots, collected in 1992-93 by the Northwest State Forest Inventory Enterprise. Twelve Landsat Thematic Mapper (TM) images were used to map
the full region. Following geographic registration and scene-to-scene radiometric normalization, unsupervised classification was used to create a
13-class land cover map. For the forest classes, forest inventory data were
used to build regression models to predict mean forest age and aboveground biomass for each forested pixel. A canonical correlation analysis
index of the TM bands was regressed against the vegetation variables using
the reduced major axis technique. Regional estimates of forest cover and
total biomass were in close agreement with ground-based estimates. Results
were assessed against an independent subset of the forest inventory plots.
Error assessment indicated that predictive accuracy varied by forest com-

position, and that the overall error was acceptable. This satellite-based forest mapping method proved to be an effective tool for estimating carbon
storage for a large boreal forest region.
OGLE, KIONA* and JAMES F. REYNOLDS. Department of Biology,
Duke University, Durham, NC. The importance of precipitation seasonality to the growth dynamics of a desert shrub.
Climate models predict shifts in precipitation to winter months in the American Southwest. To elucidate how such shifts may influence desert plants,
we are developing a plant growth model to explore the effects of seasonality on the growth of the desert shrub, Larrea tridenata (creosotebush).
The model was parameterized for Larrea growing at the Jornada LongTerm Ecological Research site in southern New Mexico. The Jornada is
characterized by three distinct seasons: hot, dry springs (Apr-Jun); hot,
moist summers (Jul-Oct); and cold, moderately dry winters (Nov-Mar).
Average long-term precipitation is around 240 mm/yr, of which approx.
65% occurs in the summer, 25% in the winter and 10% in the spring. We
employed the model and Jornada weather data from 1997-2000 (mean precip. 5 249 mm/yr) to examine the importance of seasonality to Larreas
growth. In a simulation experiment, we redistributed rainfall across the
seasons while maintaining total annual rainfall. The greatest predicted
change in peak biomass (26.2% increase over 4 years) occurred when 65%
of the rain fell in the winter. Under drought conditions, growth dynamics
were more variable and growth was greatest when summer rainfall was
high. However, relative growth rate (RGR), peak biomass, growing season
length, and timing of growth were more strongly determined by interactions
between rainfall in different seasons and were less coupled to precipitation
in a single season. For example, a shift in precipitation to favor spring
months resulted in increased RGR, especially if the following summer was
drier than the spring. Larreas greatest growth increment is expected during
years of average to above average rainfall that is distributed such that the
winter-spring period is relatively wet, followed by a relatively dry summer.
This suggests that a re-distribution of precipitation favoring winter-spring
months would enhance Larreas performance.
OGLE, STEPHEN M.,1,* DENNIS OJIMA,1 DAVE CLAUSNITZER2 and
ROBERT SANFORD.3 1 Colorado State University, Fort Collins, CO;
2
USGS, Forest & Rangeland Ecosystem Science Center, Corvallis, OR;
3
University of Denver, Denver, CO. Livestock impacts on soil organic
carbon storage in Great Basin ecosystems.
Livestock production has been shown to have dramatic impacts on Great
Basin ecosystems. Our objective was to evaluate the impact of livestock
use on soil organic carbon (SOC) storage using a combination of field
sampling and computer modeling. To quantify livestock impacts, we sampled plots at 75 and 2640 m from 4 watering holes in Oregon and Utah,
representing high- and low-use by livestock. Plots near watering holes had
significantly more soil organic carbon for 3 of the 4 sites, with high-use
plots having 406 to 510 gC m-2 more SOC than low-use plots. Consistent
with SOC, high-use plots had significantly more root biomass, ranging from
19 to 21 g m-2. Modeling the increase in root biomass accounted for 25%
of the enhancement in SOC storage near water holes. In addition, livestock
have redistributed carbon to the high-use areas, estimated at 0.1 to 1 gC
m-2 month-1, and modeling this effect accounted for an additional 30% of
the increase in carbon storage near the water holes. This research suggest
that SOC storage increases in areas surrounding water holes across Great
Basin ecosystems over time frames ranging from 20 to 50 years, and both
redistribution of carbon by livestock and shifts in plant carbon allocation
favoring root growth in those locales have contributed to this landscape
pattern.
OJIMA, DENNIS S.,1,* G-J LEE,2 N. SALIENDRA,3 D.A. JOHNSON,3,4
Y-H TANG,5 K. AKSHALOV6 and TOGTOHYN CHULUUN.7 1 Colorado
State University, Fort Collins, CO; 2 Tsukuba University, Tsukuba, Japan,
Japan; 3 Utah State University, Logan, UT; 4 USDA-ARS Forage and Range
Research Lab, Logan, UT; 5 National Institute of Environmental Sciences,
Tsukuba, Japan, Japan; 6 Barayev Kazakh Research Institute of Grain Farming, Shortandy, Kazakhstan, Kazakhstan; 7 Mongolian Academy of Sciences, Ulaanbaatar, Mongolia, Mongolia. Shared templates for modeling
carbon eddy flux and nitrogen cycling on Asian and American grasslands.
Ecologists and atmospheric scientists are concerned with understanding
how biospheric characteristics of the land surface are being modified in

response to changing climate, CO2 concentrations, and land use. Our understanding of the biological controls of carbon fluxes between the atmosphere and the land surface (referring to the soil, vegetation, water system)
is critical to our estimation of net terrestrial carbon fluxes. Recent advances
in modeling and observation systems of surface fluxes of CO2 and other
trace gases have improved our ability to understand seasonal and interannual dynamics of the biosphere. These technologies are proving to be important for cross continental comparisons of carbon and other biogeochemical fluxes. Data from US, Asian, and Central Asian grasslands will be
presented, along with analyses of the climate and land use effects on the
biosphere and atmosphere interactions. The Century model of Great Plains
short and tall grass prairie can be compared with SimCYCLE Eddy for
central Japanese (Tsukuba) meadow and northern and southern Chinese
steppes. A recent assessment indicated that salinization of soils due to the
drop of the Aral Sea level occurred over 4.9 million hectares, having an
adverse impact on the regional economy and human health. The 30-year
study of carbon balance of the chernozem soils in northern Kazakhstan
conducted at the Barayev Institute of Grain Farming (Shortandy, Kazakhstan) indicated a 25 to 30 percent reduction of humus reserves due to cultivation. Cumulative daily net CO2 fluxes during measurement periods
(174-237 days) for three years (1998-2000) indicated that Central Asian
rangelands were sinks for atmospheric CO2 in this order: Shortandy
(1.5860.26) . Karnap (1.3660.55) . Karrykul (0.3260.43 t C ha-1 season-1).
OLAPADE, OLA A.* and LAURA G. LEFF.* Kent State University, Kent,
OH. Response of stream bacterial populations in biofilms to inorganic
nutrients and dissolved organic matter.
The response patterns of bacterial taxa in stream biofilms developed on
ceramic clay pots amended with dissolved organic matter (DOM) and inorganic nutrients were examined. We determined the abundance of different
taxa, including Domain Bacteria, Cytophaga-Flavobacteria, and alpha, beta,
and gamma Proteobacteria. Additionally, population sizes of bacterial species commonly found in stream biofilms (i.e. Pseudomonas putida, Acinetobacter calcoaceticus and Burkholderia cepacia) were examined. Clay
pots amended with algal exudates, leaf leachate, glucose, phosphate, nitrate
or a combination of phosphate and nitrate were placed in the Mahoning
River during the summer and fall of 2002. After 3 weeks, samples were
collected and preserved for fluorescent in situ hybridization (FISH), a technique used for the enumeration of different taxa. The responses among the
three bacterial species were not significantly different to the presence of
DOM and nutrients in both seasons. Among the three proteobacteria taxa,
beta proteobacteria predominated, accounting for more than 40% of proteobacteria in all treatments, including controls. The results from the study
reflect the need for a seasonal examination of community composition, in
order to accurately represent the dynamics of response of bacterial populations to environmental variables, such as DOM and inorganic nutrients,
in aquatic ecosystems.
OLI, MADAN K.1,* and KENNETH B. ARMITAGE.2 1 Dept of Wildlife
Ecology, University of Florida, Gainesville, FL, USA; 2 Dept of Ecology
and Evolutionary Biology, The University of Kansas, Lawrence, KS, USA.
Yellow-bellied marmot population dynamics: Demographic mechanisms of growth and decline.
Multiple environmental factors may act synergistically to influence demographic characteristics, and ultimately the dynamics, of biological populations. Using prospective and retrospective analyses of demographic data
from a 40-yr study of individually-marked animals, we investigated the
demographic mechanisms of the temporal and spatial dynamics of a yellow-bellied marmot (Marmota flaviventris) population. Prospective elasticity analyses indicated juvenile survival rate would have the largest relative
influence on the projected population growth rate. Relative magnitudes of
elasticities did not differ between years characterized by positive and negative population growth. However, retrospective analyses of life table response experiment (LTRE) revealed that changes in age at first reproduction, followed by fertility rate, made the largest contributions to observed
annual changes in population growth rate. Changes in age at first reproduction made the largest contributions to annual declines in population
growth rate most frequently, whereas fertility rate made the largest contri-

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255

butions to increases in population growth rate most frequently. Population

dynamic differences among marmot colonies were due primarily to spatial
variations in age at maturity and juvenile survival rates. Broader implications of our findings to population ecological studies are discussed.
OLSON, JERRY S., * TAKEHISA OIKAWA, GILJAE LEE and WENHONG MO.2 1 Universityof Tennessee-Knoxville, Knoxville, Tennessee,
USA; 2 Tsukuba University, Tsukuba, Ibaraki 305, Japan. Equations and
rates of change for C3 and C4 grasses and classic sand dune successions.
1,

Holistic and reductionist models nearly merge in abstractions for pioneer

ecosystems: e.g. dunes with few species. Many models simulate rates of
change of leaves, stems, and roots, (state variables C1/C2/C3), and litter
and humus pools (C4,C5 residues), initially encompassing their decomposers. Marram grasses (Ammophila spp.), lyme grasses (Leymus spp.), and
some Elymus spp. use cool seasons for rapid expansion (positive feedback).
These thrive during and soon after rapid sand buial (Olson 1958a-d; 1997).
Lee and Oikawas (2002) "Sim-CYCLE Eddy" has an explict compartment
C6 here for C4 grasses (e.g. sand reed, Calamovilfa longifolia; little bluestem Schizachyrium scoparium 5 Andropogon scoparius) which overtake
the C3 pioneers on classic Great Lake and Great Plains dunes. Here concepts of succession reached America from Denmark (Cowles 1899, 1901;
after Warming 1891, 1895). Ishikawa compared C3 and C4 behavior on
beach dunes of Ibaraki, Japan. Sea oats (Uniola paniculata)), like sand
reed, resists erosion as well as burial by waves and wind. Sim-CYCLE
Eddy has an explicit compartment for reproductive parts (C7), peaking in
summer. One or more, or all, herbivores may sometimes be aggregated in
pool #C8. Animal residues, in C9, preferably treated collectively, suggest
estimated total consumpton. Do C8,C9 paths often hasten recycling significantly in pioneer or later successions of Shelfords (1913) Indiana age
sequence of Animal Communities in Temperate North America? Fire does
hasten carbon cycling in oak savannas (Quercus velutina..) and open dune
woodlands (Pavlovic et al.). Fuller modeling calls for a) explicit loops for
burning (Itoh and Oikawa 2002, Itoh et al. 2003 this session); b) slow and
rapid partitions of stem, root, litter and humus compartments; separate decomposers (e.g. GTEC of Post et al. 1997, 2002); c) multi-element cycling
(Parton et al. 1987, 1989; Ojima et al. 2002, 2003 this session; Olson
1963a,b; 2002); and d) grid arrays of modules for landscapes, regions or
the biosphere. Model projections have broader implications for management choices: for park, educational, amenity, forest and grazing resource values, and harbor/strand/resort protection.
ORIANS, COLIN,1,2,* SIGRID SMITH1 and LAWREN SACK.2 1 Tufts
University, Medford, MA; 2 Harvard University, Cambridge, MA. How
are leaves plumbed on a shoot? A novel hydraulic approach to quantifying species-specific sectoriality.
Most vascular plants exhibit "sectoriality"the restricted movement of water, nutrients, photosynthate, chemical signals, and other substances along
specific vascular pathways, known as Integrated Physiological Units
(IPUs). From qualitative dye studies we found that leaves on a shoot are
arranged in discrete sectors. However there is variation among species in
sectoriality. We employed a novel hydraulic technique to quantify patterns
of sectoriality in current year shoots of 1-2 species of Acer, Betula, Castanea, Fraxinus, Liriodendron and Quercus with at least six fully expanded
leaves. We measured the hydraulic resistance of the pathways between
leaves on the shoot. We severed each shoot under solution (10mM KCl),
and excised the terminal apex and 6-8 successive leaves, leaving only 2
cm petiole stubs. The base and tip of each shoot were sealed with acrylicbased glue, and petiole stubs were attached to tubes filled with KCl solution. KCl solution was applied to one petiole stub at a known pressure,
and flow was measured out of the other petiole stubs, and hydraulic resistance was quantified as the applied pressure divided by the flow rate (normalized by leaf area). As expected, resistance of flow between leaves within an IPU was low in all species. Resistance to flow between IPUs varied
among species, indicating variation in shoot sectoriality. Resistance between IPUs was significantly higher relative to resistance within IPUs for
ring-porous species (Castanea, Fraxinus and Quercus), indicating greater
sectoriality than for diffuse-porous species (Acer, Betula and Liriodendron). In Betula resistance between IPUs was the same as resistance within

256

Abstracts

IPUs. Species-differences in internal plumbing have potential implications

for the coordination of long-distance transport, as well as for responses to
heterogeneity in resource supply and herbivory.
ORLAND, MARY C.1,* and DOUGLAS A. KELT.2 1 University of California, Davis, Department of Entomology, Davis, CA, USA; 2 University
of California, Davis, Department of Wildlife, Fish and Conservation Biology, Davis, CA, USA. Hierarchical resource distribution lowers species diversity in a desert rodent assemblage: Impacts of individual-level
processes on community structure.
Individual-based models have shown that a hierarchical distribution of resources among individuals in a population has the potential to significantly
impact population persistence. We investigated the community-level consequences of a hierarchical resource distribution with a food supplementation experiment in a heteromyid rodent community in the Sonoran Desert
of southern California. Reproduction and biomass increased dramatically,
but there was no change in the density of adult rodents. There was also a
decrease in species diversity and a corresponding increase in the presence
of the dominant pocket mouse species with the addition of resources. Combined with the lack of change in home range size, the disproportionate
numerical increase of the dominant species suggests that resources are distributed hierarchically among individuals in the community. This indicates
that individual behavior can affect community level processes, and specifically that a hierarchical distribution of resources may decrease species
diversity in a system with spatially heterogeneous resources.
ORR, STUART B.,* DEBORAH L. MARR and ANDREW F. SCHNABEL. Indiana University - South Bend, South Bend, IN. A comparison of
plant species diversity between restored tallgrass prairies and native
prairie remnants.
The objective of this study was to compare plant species diversity and
abundance in restored tallgrass prairie sites with nearby remnant prairies
and to monitor the success of the restorations over time. In the summers
of 2001-2002, plant diversity and abundance were measured in three restored areas and compared to three nearby remnant prairies. The three restored areas were planted in 1997, 1998, and 1999 at The Nature Conservancy Efroymson Restoration in Newton Co., Indiana. The Shannon-Weiner index values for the restorations were 4.68 (1997), 4.54 (1998), and
4.01 (1999). The Shannon-Weiner index values (H) for the older two restorations were similar to the values for the two remnant sites analyzed (H
values for the remnant sites were 4.46 and 4.66). Although overall species
diversity was similar for the older restored and remnant sites, the restored
sites had lower numbers of native species present than the remnants (76%
native species in the restorations vs. 87% in remnant sites). Restored sites
tended to have a lower proportion of grasses to forbs in both 2001 and
2002 (range 10-59% forbs in restored sites vs. 45-58% forb in remnant
sites). There was also greater variability in grass:forb ratios among restored
sites compared to remnant sites. In addition, we found that native plant
species diversity and abundance in restored sites increased, while the remnant sites showed little change between 2001 and 2002. Further monitoring
of restored and remnant sites will continue to provide information on how
closely restorations mimic remnant prairies in plant species diversity and
on how plant species composition changes over time in both restored and
remnant prairies.
ORR, TERI J.,* WILLIAM L. GANNON, ERIN C. GREENLEE and GABOR R. RACZ. University of New Mexico, Albuquerque, New Mexico,
USA. A continuously operating acoustic monitoring station for
Bosque-flying bats in central New Mexico.
Landscape and even continental-wide perturbations may affect survivorship
and distribution of bats. Since all North American vespertilionid insectivorous bats echolocate, monitoring bat calls is an efficient way to detect
their presence. It is also possible to identify most bat species by examining
a clear call sequence. In this project we deployed one acoustic monitoring
system to record calls of free flying bats in the Bosque of the Rio Grande
in Albuquerque, New Mexico at the Rio Grande Nature Center. The monitoring system consisted of an Anabat acoustic detector, an interface module, and two DOS driven computers. A laptop computer collects, stores,

and permits download of call data to the other PC for analysis. Call file
headers were cleaned and saved and standard analysis was conducted on
all calls using Analook software. One advantage of using a monitoring
system of this type is that large amounts of data can be continually generated for long periods of time and stored. In this paper, we investigate
temporal variation (e.g., within a night, season, year) and activity of bats
surrounding the detector site. Analysis of call file data has revealed trends
in total calls per night (abundance) and changes in number of species detected during monitoring. Using natural sonic categories, sound space was
divided in order to partition different groups of calls based on minimum
frequency (kHz). Three groups were created: high frequency (40-60 kHz),
low frequency (20-32 kHz), and intermediate frequency bat calls. Several
interesting temporal, activity, and biodiversity patterns will be discussed.
ORROCK, JOHN L.1,* and ELLEN I. DAMSCHEN.2 1 Ecology and Evolutionary Biology Interdepartmental Program, Iowa State University, Ames,
IA; 2 Department of Zoology, North Carolina State University, Raleigh,
NC. Rodent seed predators may offset benefits of corridors for birddispersed plants.
Some of the detrimental effects of habitat fragmentation may be offset by
corridors, strips of habitat that promote movement of individuals and propagules among patches. However, most work in corridors has neglected how
corridors may impact predator-prey interactions. We demonstrate that corridors affect seed predation, resulting in negative effects on seeds of a birddispersed tree, black cherry (Prunus serotina). In fall 2002, we placed 20
seeds of P. serotina in each of 80 experimental exclosures located within
a large-scale, replicated landscape with connected and unconnected patches.
After four weeks, seeds were collected and the experiment was repeated.
Seed predation by arthropods was negligible, as arthropods removed , 1
% of P. serotina seeds. Conversely, rodents were significant agents of seed
predation, removing 54% of P. serotina seeds in unconnected patches and
84% of P. serotina seeds in connected patches. The significant difference
in seed predation between connected and unconnected patches suggests
that, although bird-dispersed seeds may experience increased dispersal
when corridors are present, this benefit may be offset by corridor-mediated
changes in seed predation.
ORTIZ-BARNEY, ELENA* and JULIET C. STROMBERG. Arizona State
University, Tempe, AZ. Heat tolerance of grass seeds in desert grasslands and their vulnerability to prescribed burning.
Over the last 100 years, large areas of desert grassland in the US Southwest
have shifted to shrublands. Land managers are experimenting with the use
of prescribed burning to restore these areas to grasslands. A previous study
of ours indicated that there is potential for regeneration of the herbaceous
community from the soil seed bank. However, within the soil seed bank,
the majority of seeds were found in the litter layer rather than the soil. Due
to their location, the available seeds will likely be exposed to lethal temperatures during a fire. In this study, we measured maximum fire temperatures during prescribed burns at two field sites and at two depths (soil
surface and 2-3 cm below surface). During prescribed burning, maximum
fire temperatures were extremely variable. However, the maximum temperature on the surface of the soil was higher than below the surface. Most
temperature values recorded on the soil surface ranged between 508 and
3508C whereas, sub-surface most temperature values were between 508 and
1008C. We also tested the heat tolerance of seven southwestern range grasses by placing dry seed in an oven for 2 minutes at a range of temperatures
(508 to 3008C). Of the grass seeds that germinated under laboratory conditions, none germinated after being exposed to temperatures above 1008C.
These results indicate that prescribed burning may damage viable seed
reserves on the surface of the soil, although the patchy nature of the burns
may allow for local survivorship.
OSTERTAG, REBECCA,1,* LINH VUONG2 and SUSAN CORDELL.3
University of Hawaii at Hilo, Hilo, HI; 2 University of Puget Sound, Tacoma, WA; 3 Institute of Pacific Islands Forestry, Hilo, HI. Cost of leaf
construction among native and invasive plant life forms within and
across biomes.
1

The cost of construction measures the amount of energy required to build

new leaf tissues and has the potential to be an integrative variable linking

leaf level traits, plant growth rates, and resource use efficiency. Using bomb
calorimetry, we evaluated cost of construction across different ecosystems
in Hawaii, comparing across life forms and between ecologically similar
native and invasive species. Cost of leaf construction on an area basis
(CCarea) was consistently lower for invasive species (101.3 kJ/m2 6 11.6
SE) than for native ones (235.8 kJ/m2 6 17.7 SE). CCarea was also lowest
in herbaceous species (121.2 kJ/m2 6 22.5 SE), intermediate in shrubs
(163.0 kJ/m2 6 15.8 SE), and highest in trees (279.1 kJ/m2 6 24.0 SE).
Across ecosystems, the alpine shrubland had much higher CCarea regardless of life form and notably lacked any invasive species. Dry forest, lowland wet forest, and sub-alpine forest did not differ in their overall mean
CCarea, but individual species that could be found in several ecosystems
due to the wide ecological amplitude of Hawaiian flora did differ in CCarea, with highest costs in the alpine ecosystem. For all life forms, CCarea
was highly correlated ( r . 6 0.6) with several leaf traits, including specific
leaf area (SLA), N concentration, and C:N of plant tissues, with strongest
patterns emerging for trees. Previous research has demonstrated that invasive species tend to exhibit traits such as high SLA, high N concentrations, and lower C:N, and we suggest that CCarea data may provide one
explanation of the mechanism by which invasive species can outcompete
native species for resources across a variety of climatic regimes.
OSTFELD, RICHARD S.,1,* FELICIA KEESING1,2 and KATHLEEN LOGIUDICE.1,3 1 Institute of Ecosystem Studies, Millbrook, NY; 2 Biology
Department, Bard College, Annandale, NY; 3 Biology Department, Union
College, Schenechtady, NY. Community ecology meets epidemiology:
biodiversity loss and elevated Lyme disease risk in fragmented landscapes.
Lyme disease is a tick-borne bacterial zoonosis in which the white-footed
mouse (Peromyscus leucopus) serves as the primary source (or reservoir)
from which ticks acquire the disease agent, Borrelia burgdorferi. Previous
research using both field data and modeling has shown that high species
richness within the vertebrate community results in decreased risk of human exposure to Lyme disease. High vertebrate diversity: (1) dilutes the
impact of white-footed mice by providing ticks with blood meal sources
that are unlikely to result in infection; and (2) appears to directly reduce
the abundance of mice via both predation and competition. We used an
empirically based simulation model to assess the degree to which the sequence of species loss from vertebrate communities influences the proportion of ticks infected with the etiological agent, an important risk factor.
We observed marked differences in the shapes of curves relating vertebrate
biodiversity to disease risk. All potentially realistic disassembly rules resulted in increases in disease risk with decreasing biodiversity, although
shapes of the curves varied dramatically. We extend these results by allowing species to interact with one another within foodwebs. For example, the
loss of grey squirrels can result not only in the loss of blood meals for
ticks, but also in the increased abundance of rodent competitors, which
themselves serve as blood meal sources. Greater interaction strengths within the foodweb result in stronger indirect effects of species loss on tick
infection. To understand how biodiversity loss affects Lyme disease (and
similar vector-borne diseases), it will be necessary to determine: (1) the
order by which species are lost in human-impacted ecosystems (i.e. disassembly rules); and (2) the direct and indirect effects of the loss of key
host species on tick abundance and infection.
OVADIA, OFER, HEINRICH ZU DOHNA and OSWALD SCHMITZ.
School of Forestry and Environmental Studies, Yale University, New Haven, CT, USA. Consequences of trait variation on trophic interactions
and abundances in a food chain.
Classical theory aiming to predict community structure and function assumes that smaller-scale details such as individual traits can be abstracted
safely and that community dynamics can be simply characterized in terms
of net changes in population densities. Here we use a computational model
that consists of a three-level food chain to explore the effect of initial body
size variation among herbivores on community dynamics resulting from
non-linear relationship between population demography and body size. We
illustrate that initial herbivore body size variation has a negative effect on
their survival and consequently a positive effect on the final plant biomass.
We then use trait distribution in combination with body size-survival and

Abstracts

257

-fitness curves estimated through simulations to generate predictions for

comparison with observed food chain effects. We show that, at the population-level, generating accurate predictions requires estimating fitness
curves by incorporating relevant frequency-dependent effects. However, at
the community-level, this frequency-dependence as well as changes in consumer population size can be abstracted safely and community dynamics
can be simply predicted from initial trait distribution in combination with
the survival curve. Our findings suggest a need to revisit classical theory,
opening a new research avenue for community ecology. Doing so will
require mechanistically studying population demography and experimentally testing the effect of trait variation on community dynamics.
OZGUL, ARPAT,1,* LOWELL L. GETZ2 and MADAN K. OLI.1 1 University of Florida, Gainesville, FL; 2 University of Illinois, Urbana, IL.
Demography of fluctuating populations: Temporal and phase-related
changes in vital rates of prairie voles, Microtus ochrogaster.
Small mammal population fluctuations have been an ecological puzzle and
a source of heated debates among ecologists. Identifying the demographic
parameters that closely covary with density changes can help elucidate the
underlying causes of population fluctuations, but few studies have reported
rigorous estimates of these parameters. We applied capture-mark-recapture
analysis to biweekly trapping data from a long-term study of a fluctuating
prairie vole, Microtus ochrogaster population in Illinois, USA to estimate
overall and stage-specific survival and maturation probabilities. We also
estimated population density, fecundity rates, age at maturity, and lifespans.
Survival, maturation and fecundity rates exhibited phase-related changes
during the major density fluctuations, but they showed density-independent
temporal variations during the prolonged low-density phases. Among these
variables, maturation and juvenile survival probabilities most closely covaried with population density. These results suggest that phase-related
changes in maturation and juvenile survival probabilities are likely to be
the main demographic factors driving the dynamics of our study population. Phase-related changes in maturation rates provide a plausible demographic explanation of density fluctuations. Because direct predation is unlikely to influence maturation rates, our results suggest that direct predation
may not be necessary for large-scale fluctuations in prairie vole abundance.
PACKER, ALISSA A.* and KEITH CLAY. Indiana University- Bloomington, Bloomington, IN. Interactions between black cherry seedlings
and the microbial soil community across a successional gradient.
Plant interactions with soil microbes may strongly affect the structure and
dynamics of natural communities. Negative feedback between black cherry
(Prunus serotina) and soil-borne fungal pathogens (Pythium spp.) is known
to result in high mortality of conspecific seedlings close to trees. Based on
these findings we expect uniform, rather than aggregated, spatial distributions of trees. However black cherry trees at early successional sites in the
Midwest are often aggregated. We asked whether the plant/pathogen interaction intensifies with succession, as might be predicted if host-specific
pathogens accumulate over time. We expected that seedling mortality near
conspecific trees in the field would be greater in later successional sites
than earlier successional sites. In greenhouse experiments we compared
seedling survival and growth in sterile and unsterile soil collected beneath
trees at each site, which allowed us to examine the effect of soil biota
independent of confounding factors in the field. Seedling survival did not
vary predictably with successional age in the field. In greenhouse experiments sterilization of field soil improved seedling survival regardless of
successional age, suggesting that the soil microbial community negatively
affects seedlings at all sites. We also examined the rate at which the soil
community develops by repeatedly planting seedlings in soil collected from
conspecific trees at both a young and a mature successional site (four sequential 5-week growth cycles). Host-induced alteration of the soil community was inferred from changes in seedling survival in field soil relative
to sterile soil. Negative effects of soil biota increased with repeated growth
cycles, potentially resulting from changes in microbial composition. In field
soil, survival decreased with increased growth cycles regardless of the site
age. Together these results suggest that negative plant/soil microbe feedback occurs rapidly during plant succession. Aggregated populations of
black cherry trees may occur because seedlings establish in the community
at the same time.

258

Abstracts

PAERL, HANS W.,1,* JAMES L. PINCKNEY,6 LAWRENCE W. HARDING,2 CHARLES S. HOPKINSON,3 EDWARD D. HOUDE,4 DAVID
KIMMEL,2 JAMES T. MORRIS5 and LEXIA M. VALDES.1 1 Univ. of
North Carolina, Institute of Marine Sciences, Morehead City, NC; 2 Univ.
of Maryland, Horn Point Environmental Laboratory, Cambridge, MD;
3
Marine Biological Laboratory, Woods hole, MA; 4 Univ. of Maryland,
Chesapeake Bay Laboratory, Solomons, MD; 5 Univ. of South Carolina,
Columbia, SC; 6 Texas A&M University, College Station, TX. Developing
indicators to assess human vs. climatic impacts on Atlantic coastal water quality and ecological condition.
More than half the US and world population resides within 100 km of the
coast and the absolute numbers and proportions of human inhabitants of
coastal watersheds are steadily increasing. The impacts of human encroachment are evident in estuarine and coastal waters that process a bulk of
land-based nutrients and other pollutants. These ecosystems are under the
influence of both human and natural perturbations such as major storms,
hurricanes and flooding, which may coincide and act synergistically or
antagonistically. The US Atlantic coastal region has seen its share of ecological change in coastal watersheds, with human development and no less
than 6 major hurricanes coincidentally-impacting this region in the past 6
years. Distinguishing and integrating the impacts of natural and human
stressors in time and space are difficult but essential for understanding
environmentally-driven change of biodiversity and function, starting at the
microbial indicator level. Here, we explore the complementary use of analytical (HPLC detection of diagnostic photopigments), trophic structure
(food web size spectral analysis), geomorphological, physiological and molecular indicator tools coupled to remote sensing to characterize microbial
and higher trophic level community structural and functional responses to
a variety of anthropogenic and natural physical-chemical perturbations.
These tools can be adapted to automated surveys by ferries, synoptic monitoring, and remote sensing in order to evaluate environmental controls on
biotic community structure and function over ecosystem to regional scales.
PAGEL, ALISHA L.,1,* FRANK P. DAY,1 BRUCE A. HUNGATE2 and
BERT G. DRAKE.3 1 Old Dominion University, Norfolk, VA; 2 Northern
Arizona University, Flagstaff, AZ; 3 Smithsonian Environmental Research
Center, Edgewater, MD. Influence of elevated atmospheric CO2 on root
biomass accumulation in Florida oak-scrub.
During the first two years after fire disturbance, continuous minirhizotron
observations revealed that elevated atmospheric CO2 stimulated fine root
abundance in a scrub-oak woodland in central Florida. During years 5-6,
however, fine root abundance converged in the two CO2 treatments. To
confirm these findings and to develop relationships between root biomass
and the minirhizotron observations, we removed soil cores (7 cm diameter
by 1 m deep) from each experimental chamber six years after CO2 manipulation began. Live roots, dead roots and organic matter were separated
from the soil, and live roots were sorted into five diameter classes. Similar
to recent trends observed in the minirhizotron data, preliminary core data
suggest that elevated CO2 altered neither root biomass or root length. Specific root lengths also did not appear to differ between ambient and elevated
CO2 treatments for the smaller diameter classes, but for the 2-10 mm roots,
the specific root length of elevated CO2-grown roots was 20% greater than
roots grown in ambient conditions. Elevated CO2 initially stimulated root
growth in this ecosystem, but the effect appears to be transient, disappearing after 3-5 years possibly as CO2 treatments converge on closure of the
root system.
PAHL, JAMES W.* and CURTIS J. RICHARDSON. Duke University Wetland Center, Nicholas School of the Environment and Earth Sciences, Durham, NC, USA. The role of fire in the restoration of Everglades wetland
communities.
Hydrological modification and nutrient loading from surface water inputs
have led to alterations in the community structure of northern Everglades
plant communities, where southern cattail (Typha domingensis) has been
rapidly expanding into the natural mosaic of slough, sawgrass (Cladium
jamaicense) marsh and tree island communities. We are investigating if
community type and extent of soil nutrient enrichment influence whether
fire can be used to control Typha expansion. Managed cover burns (surface

water is present to prevent root or peat burn) encompassing three plant

community types of interest - Cladium-dominated and Cladium-Typha
mixed marshes, and Cephalanthus-Salix shrub communities fringing hardwood tree islands - were conducted by the Florida Fish and Wildlife Conservation Commission from March to October 2002 in Water Conservation
Area 3a (WCA3a). The soils of WCA3a are generally considered to have
a background soil nutrient status (, 500 mg P/kg dry weight). We established sites in these three community types in both burned and non-burned
communities, and sampled for water, soil, and vegetation both before and
after fire events. Surface and interstitial water chemistry response was highly variable both temporally and spatially between and within community
type. We did observe short-term increases in interstitial phosphorus and
both increased and decreased ammonium-nitrogen following burns. Preliminary analysis of vegetation response suggests that fire had mixed success
in removing Typha from the community structure of mixed-species marshes
while maintaining structure in non-Typha communities. We will present
soil chemistry data to illustrate fire-induced changes in the peat matrix and
relate that data to vegetation response.

to provide detailed information about potential impacts of land-use change

and climate change that can be used to assess the reliability of a largerscale regional model, through an assessment of the effects of heterogeneity
on scale and aggregation. The regional modeling component of the research
is designed to construct a regional economic modeling framework that can
be linked to regional ecosystem and hydrology models to conduct integrated assessment of the economic, ecosystem and hydrological impacts of
ecosystem sustainability in Western China. The research areas that have
been chosen include investigating the transition area of Loess Plateau and
Mu Us desert in Yulin Prefecture, the Hunshandake sandy land in the
middle east of Inner Mongolia, the oases of Manas river basin in the northern Xinjiang and the area of water and soil conservation in the upper and
middle reaches of Yangtze River and the Mekong River Valley in the Yunnan province.

PAI, ASWINI* and BRIAN C. MCCARTHY. Ohio University, Athens,

Ohio. Population variability of Acorus calamus L. in patterns of resource allocation with respect to environmental variability.

We assessed competition for water between three invasive Centaurea species and two commonly occurring native bunchgrasses in the field. We
tested the hypothesis that Centaurea would reduce soil water availability
and shorten the active growing season for the co-occurring native grasses.
Intraspecific comparisons of predawn leaf water potential, midday leaf water potential, and gas exchange throughout the growing season revealed
that native grasses have less access to soil water when growing in the
presence of Centaurea. The active growing season duration was quantified
for Festuca idahoensis (C3) and Aristida longespica (C4) and the growing
season for the grasses was reduced where Centaurea occurred. There was
no significant difference however (p5 0.22) in aboveground growth of the
grasses in response to Centaurea. Instantaneous measurements of water use
efficiency (WUE) varied by day but leaf d13C of Festuca idahoensis growing with Centaurea increased (p, 0.005) by approximately 1 over the
course of the growing season (-27.6 in April compared to -26.5 in
June) indicating an increase in WUE. In contrast, there was no change in
leaf d13C of Festuca idahoensis growing in plots where Centaurea had been
experimentally removed (-27.4 in April compared to -27.5 in June).
Increased WUE by the native grasses reduces but does not eliminate the
effect of resource competition with Centaurea.

Acorus calamus L. (Sweetflag, Acoraceae) is a helophyte found in riparian

areas in temperate and subtropical regions of the world. We investigated
variation in shoot density, rhizome biomass, rhizome length, leaf scars and
leaf scars per unit length on rhizomes in eleven population patches of A.
calamus in southeastern Ohio with respect to edaphic variables. Soil samples were collected from the inside, periphery and areas adjacent to populations and subsequently analyzed for moisture, organic matter, pH, texture, Ca, Mg, Al and P content. Multivariate Analysis of Variance (MANOVA) indicated no significant variability (l 5 0.29, P . 0.05) in soil
variables within a population patch. However, significant (l 5 0.00006, P
, 0.001) differences were observed among population patches for all the
soil variables measured. MANOVA also indicated significant (l 5 0.45, P
, 0.001) differences among population patches for all the biotic variables
measured. Redundancy analysis indicated that edaphic variables explained
71.5 % of the variation in biotic variables. Monte Carlo simulations indicated significance of the overall model (F 5 0., P . 0.01). A soil organic
matter gradient defined the first RDA axis while the second axis was defined by a soil Ca and P content gradient. Rhizome length, rhizome biomass, total number of leaf scars and leaf scars per unit length are positively
correlated to Ca and P but negatively correlated to the organic matter content. Shoot density is positively correlated with silt and nitrogen content
but negatively correlated with organic matter content. Patterns of rhizome
morphology and shoot density in A. calamus populations in southeastern
Ohio indicate the life history patterns of a stress tolerator. The rhizome in
A. calamus functions as a storage and foraging organ by means of sequestering biomass and varying rhizome length and shoot density respectively
in sites with varying edaphic conditions.
PAN, SHUFEN,1,2 HANQIN TIAN,1,2 JIYUAN LIU,2 JERRY MELILLO,3
MINGLIANG LIU1,2 and XIANGZHENG DENG.2 1 The University of
Kansas, Lawrence, KS 66045; 2 Institute of Geographic Science and Natural Resource Research, Chinese Academy of Science, Beijing, 100101,
China; 3 The Ecosystem Center, Marine Biological Laboratory, Woods
Hole, MA 02543. Linking ecosystem and economic models for integrated ecosystem assessment of western China.
In this research we develop a system modeling framework for integrated
ecosystem assessment at a regional scale. This system modeling framework
is a close coupling of three existing models including the terrestrial ecosystem model, the land use model and an economic decision model. We
apply the system modeling framework for the integrated ecosystem assessment of Western China, a sub-global assessment of the Millennium
Ecosystem Assessment Project (MA). We investigate how changing human
impacts (e.g. population growth, urbanization, industrialization and landuse change) and changing natural processes (e.g. climate variability and
change) affect the sustainability of ecosystems and economic productivity
in the region. The multi-scale assessment approach has been taken in this
research. The county-level modeling component of the research is designed

PANKEY, J. R.,* R. W. LEE and R. A. BLACK. Washington State University, Pullman, WA. Phenologic and physiologic response of native
bunchgrasses to Centaurea invasion.

PAPAIK, MICHAEL J.,1,2,* CHARLES D. CANHAM,1 KERRY D.

WOODS3 and ERIKA F. LATTY.4 1 Institute of Ecosystem Studies, Millbrook, NY; 2 University of Massachusetts, Amherst, MA; 3 Benningon College, Bennington, VT; 4 University of Wisconsin, Madison, WI. Interactions between beech bark disease and wind disturbance with consequences to stand structure and composition.
Over the past century introduced pests and pathogens have caused widespread declines of a number of tree species in forests of eastern North
America, including dominant species such as beech (Fagus grandifolia).
Most studies have focused on changes to forest composition and structure
as a direct result of the mortality caused by the pest or pathogen. Our
studies of the resistance of trees to windthrow in forests of northern New
York and the upper peninsula of Michigan suggest the hypothesis that the
resistance of beech trees to windthrow is strongly dependent on the presence of beech bark disease (BBD). The disease is absent from forests of
northern Michigan, and in those forests, beech and sugar maple (Acer saccharum) have identical susceptibility to windthrow, while in northern New
York, where BBD has been present for 50 years, beech trees have much
higher susceptibility to windthrow. Within the New York population, there
is a significant negative effect on resistance to windthrow of the level of
BBD on individual trees. We tested potential consequences of this effect
on long-term structure and composition in these forests using a computer
simulation model, SORTIE. In the absence of BBD, beech is the competitive dominant of the 9 major tree species in this region. In simulations in
which the hypothesized impact of BBD on the resistance of beech to wind
was included, no competitive dominant emerged, species such as hemlock
(Tsuga canadensis) and yellow birch (Betula alleghaniensis) increased dramatically in abundance, and the overall diversity of tree species increased.
Our results highlight the effects that host-specific pathogens of dominant
species can have on long-term competitive dynamics and species coexistence in forests.

Abstracts

259

PARADISE, CHRIS J.* and NICOLE P. HARLAN. Davidson College,

Davidson, NC, USA. Relationships among habitat size, dissolved oxygen, insects, and microbes in artificial treehole communities.
In this ongoing study we are examining interactions among abiotic factors
and the diversity of insects and microbes in treeholes, aquatic detritus-based
communities. We employed mesocosms (2 depths crossed with 3 diameters)
to test the hypotheses that volume, surface area, and dissolved oxygen (DO)
affect insect and microbial diversity, and the latter factors affect one another. We sampled habitats for insects, dissolved oxygen, and bacteria. DO
should be lower in deep containers with small surface area, which we did
find. Large treeholes with a large opening have high DO, and should support high larval densities. Our containers did not show the latter; containers
with small diameter and shallow depth had high larval densities, and low
DO was correlated with high densities. Densities of common species were
positively correlated with one another and with species richness. At the
peak of species richness, an interaction between surface area and depth was
revealed: some, but not all, shallow treatments had more species than corresponding deep treatments. Results to date indicate that surface area, volume, and resulting DO levels help to shape communities in treeholes, possibly by affecting where females oviposit. Further results from the 2003
season will be presented, as will results from microbial sampling. We predict that as insects consume microbes, microbial diversity increases, which
further increases litter breakdown and facilitates insect diversity.
PARDO, LINDA H.,1,* STEVEN G. MCNULTY2 and JOHNNY L.
BOGGS.2 1 USDA Forest Service, lpardo@fs.fed.us, Burlington, VT;
2
USDA Forest Servce, Raleigh, NC. Effects of N deposition on high
elevation forests in the northeastern US: foliar d15N patterns.
Recent studies have demonstrated that natural abundance of 15N can be a
useful tool for assessing N saturation, because as nitrification and nitrate
loss increase, d15N of foliage and soil also increases. In this study, we
measured foliar d15N at eleven high elevation spruce-fir stands along a N
deposition gradient from Maine to New York in 1987-88 and from seven
paired northern hardwood and spruce-fir stands in 1999. In 1999, foliar
d15N in the spruce-fir stands ranged from 25.2 to 20.7 while foliar C:
N ranged from 52 to 30 with increasing N deposition from Maine to NY.
Foliar d15N in the hardwood stands was on average 1 higher than the
spruce-fir foliage, and ranged from 24.2 to 20.5 with increasing deposition. Hardwood foliar C:N ranged from 28 to 16 along the gradient.
No changes were observed in foliar d15N between 1987 and 1999, however
foliar %N increased and foliar C:N decreased at all sites except the site
receiving the lowest deposition (Howland Forest in Maine). Additional
measurements included soil C:N and base cations, net mineralization and
nitrification potential, basal area, and seedling regeneration. Foliar d15N was
correlated most strongly with N deposition, and was also positively correlated with net nitrification potential and negatively correlated with soil
C:N ratio. The spruce-fir foliar d15N values at the upper end of the deposition gradient approach values observed at a site (Mt Ascutney, VT) where
foliar d15N went from 23.5 to +0.5 as the site reached N saturation. This
study allowed us to evaluate ecosystem response to different levels of N
deposition.
PARHAM, JAMES E.* University of Nebraska, Lincoln, NE,
jparham2@unl.edu. Predicting instream habitat and reach occupancy
for native Hawaiian stream fishes.
The interaction between physical landscape attributes and the ecology of
native Hawaiian fishes was examined to develop a model to predict potential occupancy and community membership within different stream reaches.
The model considered the life history traits of the fishes and quantified
barriers to migration, distance inland, habitat suitability, and biotic interactions among species to determine potential community membership within a stream reach. To determine predicted species membership within a
reach of a stream, geomorphic and biotic factors were combined for each
species, and then the extent of instream distribution was calculated for each
10m cell within the stream network. To test the predictions of the model,
predictions were compared to results of stream surveys. Between 20 and
30 observation of fish species were made in approximately 200 m of stream
to determine current reach occupancy. The surveys were carried out for 16

260

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reaches in eight different streams. The streams included in the analysis

were: Hakalau, Kolekole, and Wailoa on the Island of Hawaii; Waimea,
Waiahole, and Kahana on the Island of Oahu; and Hanakapiai, Waiahuakua
on the Island of Kauai. Overall, the model correctly predicted reach occupancy at an 89% success rate. All of the false positive errors occurred
in streams on the highly developed Island of Oahu and were likely the
result of degraded habitat quality or low fish populations as opposed to the
model incorrectly classifying habitat suitability. The results highlight the
need for management techniques that depict instream habitat as a function
of natural geomorphic processes within a stream. Additionally, assessment
of the expected species assemblage needs consideration before determining
the extent of deviation of stream conditions from the natural state. By
applying a single standard to all streams, the chances of observing anthropogenic disturbances are small and due to the wide range in natural variation.
PARK, RICHARD A.1,* and JONATHAN S. CLOUGH.2 1 Eco Modeling,
dickpark@cableone.net, Diamondhead, MS, USA; 2 Warren Pinnacle Consulting Inc., jsc@warrenpinnacle.com, Warren, VT, USA. Modeling the
fate and effects of pollutants in estuarine ecosystems with AQUATOX.
AQUATOX is a process-based, time-varying, fate and effects simulation
model that integrates aquatic ecology, chemical dynamics, bioaccumulation, and ecotoxicology. AQUATOX can be used to predict not only the
environmental fate of nutrients, pesticides, and other chemicals in aquatic
ecosystems, but also their simultaneous direct and indirect effects on organisms. AQUATOX provides Latin hypercube uncertainty analysis for any
and all loadings and chemical and biotic parameters. Results are given in
tabular and graphical forms, including concentrations, rates, mass balances,
and probabilistic risk graphs. The model has been peer reviewed and released for use by the U.S. Environmental Protection Agency. It has been
validated for a variety of environments including lakes, reservoirs, small
streams, and rivers. Recently an estuarine version was developed and tested. Application to Galveston Bay TX exemplifies the effects of eutrophication, turbidity, detrital input from wetlands, and toxic organic chemicals
on water quality and commercial fisheries, and bioaccumulation of toxicants in commercial fish and sea gulls.
PARKER, SOPHIE S.* and JOSHUA P. SCHIMEL. University of California, Santa Barbara, CA, USA. Nitrogen losses from native perennial
and exotic annual grasslands in California.
Given that the introduction of even a single plant species can have large
effects on nitrogen cycling in an ecosystem, the invasion of native perennial
grassland communities by exotic species of annual grasses from Europe
raises many questions regarding the processing and retention of nitrogen
in these systems. To examine the effects of grassland species change on
nitrogen cycling, we employed the use of experimental grassland plots with
homogeneous initial soils. Plots were seeded with a mix of either native
perennial grasses (Nassella pulchra, Bromus carinatus, and Elymus glaucus) or nonnative annuals (Bromus hordeaceus, Bromus madritensis, and
Hordeum murinum). Using tension lysimeters, we quantified leaching losses of nitrate, ammonium, and dissolved organic nitrogen. Rates of mineralization, nitrification, and denitrification were measured to examine the
relative importance of these different microbial processes in contributing
to nitrogen loss from each experimental grassland type. Comparisons of
microbial process rates were also made in naturally occurring annual and
perennial grasslands. Exotic annual grass stands may be somewhat more
leaky with regard to nitrogen than stands of native perennial grasses. Seasonal trends in microbial process rates that correspond with soil moisture
indicate that rates of nitrification and denitrification, as well as leaching
losses, may be dependent on the amount and timing of rainfall. In dry
years, low soil moisture may be more important than species composition
in determining nitrogen losses from grasslands, while differences in plant
phenology and microbial process rates may drive nitrogen loss in wetter
years.
PARMENTER, ROBERT R.* Department of Biology, MSC03 2020, 1
University of New Mexico, Albuquerque, NM, USA. Vegetation recovery
following fire in a desert grassland in New Mexico, USA.
The post-fire recovery of dominant plant species in a desert grassland on
the Sevilleta Long Term Ecological Research site in central New Mexico

was measured from 1992 through 2001. Pre- and post-burn data were collected on individual, permanently mapped plants. Annual regrowth data for
the first 5 years were used to develop species-specific linear models to
predict time requirements for complete fire recovery. Woody plants were
measured again after 9 years to evaluate the model predictions. As expected, plant species displayed a wide range of tolerance to and recovery
from fire. Mortality rates were highest in cacti (particularly chollas), and
were lowest in grasses, yuccas, and certain shrub species. Regrowth rates
across grass species ranged from 3 months to an estimated 6-7 years. Yucca
recovered in a single year, and exhibited large increases in numbers of
stump sprouts. Shrubs displayed widely varying regrowth rates among species, ranging from 1 to 13 years. Most species flowered and produced seeds
within 2 years of the fire. Herbivory by black-tailed jackrabbits on several
of the shrub species was particularly heavy in the first year following the
fire, but declined to near zero in subsequent years. Overall, this grassland
community is expected to show complete recovery from this particular fire
event within 15 years. This time interval is consistent with estimates of
natural fire return times in desert grasslands of New Mexico and Arizona.
PARMESAN, CAMILLE. University of Texas - Austin, Austin. Invasions
as a consequence of climate-mediated range shifts.
Observed changes in natural systems, largely over the past century, indicate
a clear global climate change signal. Even in the face of apparently dominating forces, such as direct, human-driven habitat destruction and alteration, this climate fingerprint implicates global climate change as a new
and important driving force on wild plants and animals. Analyses of climate
and of biotic change indicate that on a global scale, climate change has
affected about half of the wild species studied (Parmesan & Yohe, 2003).
One of the most prominent responses has been poleward and upward shifts
in species ranges, apparently tracking climatic shifts. In many parts of the
world, this process has led to introductions of species into areas where they
were previously absent, leading, in the short term, to local increases in
species richness. Long-term effects are not obvious, as the outcome of
interactions among novel and ancestral species are often difficult to predict.
The evidence for this process occuring during recent climate change will
be reviewed, and the implications discussed.
PARRY HECHT, BROOKE1 and KRISTIINA A. VOGT.2 1 Yale University, School of Forestry and Environmental Studies, New Haven, CT; 2 University of Washington, College of Forest Resources, Seattle, Washington.
The edge paradox: Investigating the impacts of multiple and novel
disturbances on forest ecosystem thresholds.
Edges have been considered potentially sensitive locations for detecting
ecosystem response to anthropogenic and natural disturbances. The current
paradigm is that plant responses to stress are amplified at ecotones, and
that edges such as the treeline will be among the first to respond to climate
change. However, previous edge studies have not been designed to examine
ecosystem level resistance and resilience characteristics in response to multiple disturbances. For example, it is not clear how the function of the
forest limit at the boreal-arctic ecotone changes when there is an overlay
of human land use on climatic disturbances. We assessed the resistance
characteristics of plant communities at the boreal-arctic forest limit in landscapes with existing and long-term legacies of human land use. It was
hypothesized that the existence of multiple disturbances and land use legacies at the forest limit would decrease the resistance of the edge to disturbance, such that it would be less able to sustain its acquisition of limiting
nutrients. It was also hypothesized that the accumulation of disturbance
factors in the landscape could shift the location of functional thresholds
through space. Research plots were established at the Betula pubescens
forest limit and at 50m and 100m in elevation below the forest limit in
Iceland. Plots were manipulated by the addition of sugar, with the aim of
immobilizing soil nitrogen. Site response to the sugar disturbance was
tracked by measuring foliar carbon and nitrogen, relative foliar chlorophyll
content, leaf area, and leaf weight. The results of this research suggest that
structural edges can have varying functional roles in the landscape, that
functional thresholds are not necessarily associated with the structural edge,
and that patterns associated with the treeline zone can change or disappear
as legacies of disturbance accumulate in an ecosystem.

PARRY, DYLAN1,* and GEORGE H. BOETTNER.2 1 State University of

New York, College of Environmental Science and Forestry, Syracuse, NY,
USA; 2 University of Massachusetts, Amherst, MA, USA. Vanishing native parasitoids: Competitive exclusion by an exotic biological control
agent?
The non-target effects of parasitoids released for biological control have
increasingly drawn attention from ecologists. A classic case is that of
Compsilura concinnata, a tachinid fly introduced to control gypsy moth in
1907. This old world species is now known to attack .200 species of
native Lepidoptera in North America and has been implicated in the decline
of several species of giant silk moth. The effect of C. concinnata on native
parasitoids is not known, even though it is often the only species reared
from several different native hosts in New England. To investigate whether
C. concinnata has displaced native parasitoids, we focused on the North
American analog of C. concinnata, the highly polyphagous tachinid Lespesia frenchii. Our study was conducted in areas of New England where
comprehensive data on parasitoid diversity was collected for more than 100
native Lepidoptera shortly (1915-1929) after the release of C. concinnata.
This enabled us to compare present day parasitoid diversity with historical
data from the same region. We utilized two complimentary approaches.
First, we deployed experimental populations of several forest lepidopteran
hosts and retrieved sentinel larvae to assess parasitism and species diversity. Secondly, we quantified the presence or absence of L. frenchii in 15
museum collections from 1920-2000 in three states where the initial arrival
of C. concinnata was well documented. We did not recover L. frenchii
from any experimentally reared or wild collected host species in New England, including those where L. frenchii was the dominant parasitoid prior
to 1930. Museum collection data corroborated the results from our field
studies. The rate at which L. frenchii acquisitions were added to entomological collections dropped sharply after the introduction of C. concinnata.
Although the mechanism that allows C. concinnata to be competitively
superior is unclear, both interference and exploitative competition are likely.
PARSHALL, TIM E.,1 CHARLES W. ANDERSON,2 JIM J. GALLAGHER,2 G. GREEN,1 MIKE J. KLUG1 and G. P. ROBERTSON.1 1 Michigan
State University, Kellogg Biological Station, Hickory Corners, MI; 2 Michigan State University, East Lansing, MI. Long-term ecosystem education
through the Kellogg Biological Stations K12 Partnership for Science
Literacy.
The Kellogg Biological Stations Partnership for Science Literacy is a collaboration between K12 science teachers in southwestern Michigan and
research scientists and educators from Michigan State University. All participants are involved in the partnership over the course of three years with
the understanding that educational reform is a long-term endeavor. Seventy
teachers from 15 rural school districts attend workshops and summer institutes and meet regularly with advanced ecology Ph.D. students with the
goal of improving science classroom instruction and teacher understanding
of science content. The science is rooted in ecosystem ecology, based directly on five research areas studied by Long Term Ecological Research
sites: primary productivity, decomposition, populations and trophic levels,
nutrient cycling, and disturbances. This ecosystem framework is used to
connect science concepts across disciplines and to link science teaching to
ongoing scientific research. After two years, our approach has facilitated
the most change in teacher understanding of science content and classroom
teaching practice. In our first set of week-long summer institutes, teachers
engaged in scientific study of local ecosystems and developed individual
research projects in order to acquire a firsthand understanding of science
inquiry. The ecological study and research project helped teachers develop
instructional units to implement in their classrooms during the coming
schoolyear. Most teachers implemented their instructional units, oftentimes
with critical assistance from ecology Ph.D. students. The schoolyear workshops serve to introduce and reinforce ecosystem concepts and further explore inquiry-based science, assessment, and components of teaching for
understanding. Ongoing communication and assessment among project participants has led to several insights and changes to improve the effectiveness of the workshops, including encouraging small learning communities,
modeling effective science instruction, and paying attention to the diverse
backgrounds of teachers.

Abstracts

261

PARTON, WILLIAM J.,1,* STEVE DELGROSSO,1 DENNIS S. OJIMA,1

ARVIN R. MOSIER2 and JACK MORGAN.2 1 Colorado State University,
Fort Collins, CO, USA; 2 USDA-ARS, Fort Collins, CO, USA. Simulated
Nitrogen Constraints on Elevated CO2 Responses for Grasslands and
Forests.
There is increasing evidence that the initial positive increases in net plant
production (NPP) following step increases in atmospheric CO2 levels are
substantially reduced by soil nutrient limitations. This paper will use the
Daycent ecosystem model to simulate the short and long term impacts of
soil nutrient limitations on grassland and forest system responses to step
increases in atmospheric CO2 levels. The Daycent model has been tested
using observed data from grassland elevated CO2 experiments in Colorado,
California, and Kansas and the Oak Ridge and Duke Forest sites. Model
results mostly represent the observed ecosystem response to increasing atmospheric CO2 levels and suggest that the initial increase in NPP following
a step increase in atmospheric CO2 levels will be greatly reduced by nutrient limitations after 5 to 10 years for the forest systems, while the elevated CO2 impacts for grasslands are sustained for up to 100 years. The
positive impact of increasing atmospheric CO2 levels for grasslands are
largest for the short grass steppe and minimal for the annual grassland
system. Model results will contrast the short (1 to 5 year) and long term
(10 to 20 year) responses of grasslands and forests to elevated CO2 levels
and compare observed versus simulated ecosystem responses to experimental data from elevated CO2 and soil nutrient experiments.
PASCARELLA, JOHN B.,1,* STEPHEN L. BUCHMANN2 and ARTHUR
J. DONOVAN.2 1 Valdosta State University, Valdosta, GA, jbpascar@
valdosta.edu; 2 The Bee Works, Inc., 1870 West Prince Road Suite 16,
Tucson, AZ, steve@thebeeworks.com. Bee biodiversity in the southeastern U.S.
Are pollinators declining in Southeastern ecosystems? We sampled pollinator biodiversity, focusing on bees (Family Apoidea), in the upland forested ecosystems of the Southeastern United States. Sites sampled included
three sites in Northern Florida (St. Marks National Wildlife Refuge (STM),
Osceola National Forest (ONF), and Tall Timbers Research Station (TT))
and one site in Georgia (Wade Tract (WT)). WT and TT were sampled
biweekly (1/2 day each) from May 1998-May 2002 and ONF biweekly (1
full day)from July 2001 to July 2002. STM was sampled 3 times (3-4 day
trips) in 2001-2002 during the warmer months. The areas sampled represented a wide variety of environments, including xeric sandhills and mesic
flatwoods, old-growth longleaf and secondary forests of loblolly and slash
pine, and burned and unburned areas in each habitat. We sampled bees
using a variety of sampling methods (sweep netting on flowers, pan traps,
and trap nests) during the active months of the year (February-November).
From all four sites combined, 3171 specimens representing 170 species
(132 pollen-collecting and 38 parasitic bees) were collected. The majority
of specimens were caught using sweep nets. Total bee species richness was
104, 119, 62, and 52 species at WT, TT, ONF, and STM, respectively. The
majority of specimens and species were from the Apidae, Halictidae and
Megachilidae, with fewer numbers from the Colletidae, Andrenidae, and
Melittidae. Two temporal peaks in species richness were noted in the spring
(April) and fall (October). Regionally, TT and WT were most similar, with
STM being intermediate and ONF least similar. Both regional (5soils,
hydrology, vegetation) and historical (5landuse) effects influence patterns
of bee biodiversity in the Southeast.
PASCUAL, MERCEDES.* University of Michigan, pascual@umich.edu,
Ann Arbor, MI. Introduction: From fuzzy trajectories to new patterns.
There have been many earlier debates in ecology on the role of stochasticity: Andrewartha and Birch vs. Nicholson and Smith, Gleason vs. Clements, Chaos vs. Noise, etc. In these debates, however, random fluctuations
were seen as an alternative to determinism. Today, ecological modelling is
beyond this simple dichotomy and it is generally agreed that noise and
determinism interact. There is, however, no consensus on the consequences
of this interaction. At one extreme, noise has been assumed merely to blur
equilibria or deterministic attractors, while more interesting examples have
shown how noise provides the fluctuations responsible for the transient
transitions between attractors. Recent work highlights a much more inter-

262

Abstracts

esting outcome in which noise, via its interaction with nonlinearities, gives
rise to surprising dynamics. I introduce the session by briefly presenting
these different classes of interaction and the questions they raise on the
usefulness of deterministic skeletons. Examples are drawn from recent results on quasicycles in predator-prey systems and on the irregular interannual variability of historical cholera patterns.
PASSMORE, HEATHER A.* and WILLIAM J. PLATT. Louisiana State
University, Baton Rouge, LA. Disturbance interactions: Effects of canopy removal and fine fuels on fire characteristics and vegetation response.
Interacting natural disturbances, especially in frequently disturbed systems,
may alter plant communities. Increased fuel loads and openness of the
canopy after hurricanes have been postulated to increase fire probability
and intensity in Southeastern habitats, potentially converting forests into
pine savannas. We predict that if fire effects are modified by a prior wind
disturbance such as a hurricane, these effects should be most evident along
slopes between upland savannas and downslope hardwood forests. In this
study, we experimentally manipulated canopy cover and fuel loads to explore whether hurricanes alter the effects normally expected from fires in
the ecotone between frequently burned second growth savanna and downslope hardwood forests. We measured characteristics of fires and responses
of vegetation. Canopy cover was reduced by removal of approximately 10
canopy trees per plot. We manipulated fuel type by adding longleaf pine,
mixed pine, and deciduous fine fuels to different 1m2 vegetation subplots.
Reduced canopy cover resulted in increased light levels at both 0.65m and
1.5m compared to undisturbed plots. Canopy removal resulted in higher
maximum temperatures in fires. Maximum fire temperatures also differed
with fuel types. Mean maximum fire temperatures were lower for hardwood
fuels and controls than both pine fuel types. There were also significant
interactive effects between canopy removal and fuel additions on maximum
fire temperatures. Canopy removal and pine fuel type treatments resulted
in decreased mean stem density in canopy removal plots compared to hardwood fuels and controls. Fuel additions resulted in fires with higher maximum temperatures and reduced plant survival and density compared to
control plots. In this study, prior canopy disturbance modified the effects
of subsequent fires and also influenced understory vegetation. Repeated
interactions between these frequent, natural disturbances may influence the
composition and structure of ecotones between Southeastern pine savannas
and downslope forests.
PATAKI, DIANE E. University of Utah, Salt Lake City, UT. Mechanisms
of cottonwood decline in a riparian ecosystem along the Colorado River.
Riparian ecosystems are a critical but rapidly degrading habitat in the arid
West. Populus fremontii (Fremont cottonwood) was once a dominant species in desert riparian forests that has been increasingly replaced by the
exotic invasive Tamarix ramosissima (saltcedar). In addition, reduced
flooding frequency and increased salinity have been implicated in the widespread decline of P. fremontii. To elucidate some of the multiple and interacting mechanisms of this decline, I examined several aspects of ecosystem function in a control stand of P. fremontii along the Colorado River
in Moab, Utah, USA, as well a disturbed stand characterized by high
groundwater salinity and invasion of T. ramosissima. Water isotope data
showed that P. fremontii and T. ramosissma utilized groundwater at both
sites. Sap flux data showed that P. fremontii at the saline site experienced
large reductions in afternoon stomatal conductance relative to the control.
Thus, average daily stand transpiration was 4.8 mm d-1 at the saline site in
comparison to 9.8 mm d-1 at the control. Despite the fact that P. fremontii
and T. ramosissma shared the same water source, gas exchange and sap
flux data scaled to canopy transpiration indicated that T. ramosissima constituted a negligible portion of canopy water use due its small contribution
to total leaf area. However, tissue N data indicated that T. ramosissima was
out-competing P. fremontii for nitrogen, likely due to a larger concentration
of roots in the upper soil horizons. Coupled with physiological stress
caused by groundwater salinity, perturbations to nutrient uptake may be
contributing to the decline of P. fremontii during invasion of T. ramosissima.

PATTEN, BERNARD C.,1,* STUART R. BORRETT,1 STUART J. WHIPPLE,2 ROBERT R. CHRISTIAN3 and CASSONDRA R. THOMAS.4 1 Institute of Ecology, University of Georgia, Athens, GA; 2 Skidaway Institute
of Oceanography, 10 Ocean Science Circle, Savannah, GA; 3 Biology Department, East Carolina University, Greenville, NC; 4 Department of Environmental Sciences, University of Virginia, Charlottesville, VA. Discrete-time dynamic environ analysis of indirect effects in ecological
networks: Basic considerations.
Environ analysis is a set of input-output methods to analyze the environments within systems (termed environs) of the systems component parts.
Analyses of internal pathways, throughflows (summed node flows), and
node storages provide core methodologies, from which additional analyses
for utility and control are derived. Existing methods are restricted to timeinvariant (stationary) systems at steady state. Needed are methods for more
realistic cases, like food webs and biogeochemical cycles, which are neither
stationary nor at steady state due to time variation of internal structure and
function, or inputs. Methods for time-continuous dynamic analyses are
cumbersome, and discrete-time approaches have yet to be attempted. This
paper explores a discrete approach for the forward-time (output-environ)
version of throughflow analysis; methods for throughflow input environs,
and storage input and output environs, follow parallel development. Three
cases are considered: (1) intensive analysis of nonstationary systems with
constant input, and extensive analyses of (2) stationary and (3) nonstationary systems with time-varying input. For Case 1 we address how to couple
discrete steps in the time series, and investigate how dominant indirect
effects typically observed for stationary systems are altered by discretetime successions of flow-generating matrices. For Cases 2 and 3, we also
consider how to couple discrete steps in the time series, we determine what
fractions of extended pathway networks are engaged during each time period, and we explore the effects of time-series truncation on indirect-effects
dominance. We also consider alternative scenarios with respect to the correspondence between time and pathway length, and we show that as time
increases with increasing length, the relative role of indirect effects declines. In all cases, however, indirect effects still dominate. We illustrate
our results with a discrete-time model (16 seasons, over 4 years) of nitrogen
loading in the Neuse River Estuary, North Carolina.
PAVAO-ZUCKERMAN, MITCHELL A.* and DAVID C. COLEMAN. Institute of Ecology, University of Geogia, Athens, GA, USA. Soil quality
indicators of ecosystem health for urban places.
Urban ecology answers the need to understand the environmental implications of global trends in urbanization. The health of an ecosystem is
related to its ability to sustain functions and provide services through time.
In terrestrial ecosystems, soil quality is directly related to ecosystem health.
Several indicators of soil health have been proposed which take into account various physical, chemical and biological properties of soils. This
paper will introduce the concepts of ecosystem health and soil quality.
Additionally, it will discuss the use of indicators to assess ecosystem health
in urban ecosystems, illustrated by the application of soil quality indicators
to the study of an urban ecosystem (Asheville, NC). This study implements
the urban gradient paradigm, comparing the soils of forested plots along a
transect running from downtown Asheville, NC to the Pisgah National
Forest to the southwest. Soils were sampled seasonally, and analyzed for
a suite of physicochemical (e.g. bulk density, moisture, organic matter),
biological (e.g. microbial and nematode populations), and process data (e.g.
decomposition, mineralization). These data will highlight some of the physical, chemical, and biotic interactions possible in urban soil ecosystems.
This study indicates that the ecosystem health of forests in Asheville, NC,
is impacted by urbanization with respect to soil quality and nutrient cycling.
Moreover, this study demonstrates that indicators of soil quality for ecosystem health can be applied to urban ecosystems. Indicators of soil quality
for ecosystem health have the potential for a widespread application for
ecological planning, monitoring, and restoration in urban places.
PAWLOWSKI, CHRISTOPHER W.* and AUDREY L. MAYER. US EPA
National Risk Management Research Laboratory, Cincinnati, OH, USA.
Resilience of ecosystems to disturbances.
Resilience, in an ecological context, is one of several terms that characterize the response of an ecosystem to disturbance. Other such terms include

persistence, resistance and stability. Two definitions of resilience have become prominent in the literature, both of which derive from the dynamic
systems theory concept of stability, and both of which concern important
aspects of ecosystem management. One refers to the speed of return of a
system to its prior, undisturbed steady state. The other refers to the magnitude of disturbance an ecosystem can absorb before it falls into the basin
of attraction of a fundamentally different steady state. We argue that these
conceptualizations of resilience do not admit the direct study of disturbance
effects on ecosystems because they use state displacement as a proxy for
disturbance. Here we suggest a theoretical measure of resilience that uses
the concept of invariance to determine, in terms of disturbance size directly,
how much disturbance an ecosystem can tolerate. Presentation of these
resilience concepts is made in the context of a simple model of lake eutrophication.
PEARLSTINE, LEONARD G.,1,* FRANK J. MAZZOTTI1 and DONALD
L. DEANGELIS.2 1 University of Florida, Ft Lauderdale, Florida; 2 United
States Geological Survey, Miami, Florida. Spatial decision support systems for Everglades restoration and management.
Restoration of the Florida Everglades habitat is characterized by conflicting
goals, difficult issues, scientific uncertainty, and complex interrelationships.
Decision support systems provide support methodologies that help authorities involved in ecological restoration sort out spatial explicit (1) knowledge acquisition and representation, (2) goals and issues identification and
conflict resolution, (3) alternatives evaluation, and (4) group negotiation
support. Spatial decision support software and many existing spatial data
layers from modeled output including vegetation classifications, wildlife
models, and hydrology models have been brought together for regional
habitat evaluation in south Florida. Other necessary layers such as socioeconomic impacts are not well developed and need attention. Uncertainty
in the modeled outputs is explicitly presented to the user in a manner that
increases the decision-makers understanding of the range of responses and
provides a mechanism for deciding where additional information is needed.
Development of decision support systems has proceeded in an iterative
process that operates within the context, needs, and objectives of the resource agencies, planners, and managers targeted as users. The collaboration of key agency individuals, helps to ensure that this effort will be
focused on the needs and objectives of on-the-ground natural resource planners and managers.
PECHMANN, JOSEPH H.K.,1,* DAVID E. SCOTT,2 J. WHITFIELD GIBBONS,2 JUDITH L. GREENE,2 BRIAN S. METTS2 and PHILIP M. DIXON.3 1 University of New Orleans, New Orleans, LA; 2 Savannah River
Ecology Laboratory, Aiken, SC; 3 Iowa State University, Ames, IA. Updated amphibian population trends at Rainbow Bay, South Carolina:
Natural and human influences.
Evaluating the extent and causes of declines in amphibian populations requires long-term census data. We have counted amphibians migrating to
and from Rainbow Bay, SC daily since September 1978 using a drift fence
with pitfall traps. Numbers of breeding adults and metamorphosed juveniles
of five focal species varied among years by several orders of magnitude.
Breeding population sizes of Ambystoma tigrinum, Pseudacris nigrita, and
P. ornata declined to near zero over the study. Breeding female A. opacum
increased from zero to .1000 during the period, whereas A. talpoideum
exhibited no overall trend. Population declines were related to recurrent
intense droughts, especially those associated with strong La Nina events,
which reduced or eliminated breeding activity and juvenile recruitment.
Although we have viewed drought effects as natural fluctuations, there are
suggestions that global warming may have increased drought frequency
and severity. Drought and fire suppression allowed trees to invade the pond
basin over the study, which may have reduced hydroperiods. The effects
of canopy closure on temperature, food resources, and other factors may
have made the habitat less suitable for Pseudacris. Ambystoma opacum
was less affected by droughts than the other Ambystoma because of its life
history. Predation by the increased numbers of A. opacum larvae may have
reduced juvenile recruitment of other species. Population trends at Rainbow
Bay have many causes which are difficult to distinguish because many are
temporally confounded.

Abstracts

263

PECK, AMANDA E.* and CRAIG R. ALLEN. South Carolina Cooperative Fish and Wildlife Research Unit, Clemson University, Clemson, SC,
USA. Change in relative abundance of small mammal species through
50 years of forest restoration.

PEDERSEN, AMY B.* and JANIS ANTONOVICS. University of Virginia, abp3a@virginia.edu, Charlottesville, VA. Removal of intestinal nematodes in wild populations of Peromyscus leucopus: Effects on behavior
and survival.

Vegetation change through succession can impact the small mammal community composition of an area. The Savannah River Site (SRS) has been
an important geographic area for the study of biodiversity and the effects
of human-induced landscape manipulation since its acquisition in 1951. We
examined how small mammal species composition has changed in habitat
types, after acquisition (1951) to current maturing forest (2002). Specifically, we examined if small mammal abundance differed among the periods
1950s, 1960s and 1990s. Next, we determined if the variance in abundance
differed among those same periods. Lastly, we determined if abundance of
any particular species changed more than expected based on change in
habitat area. Small mammals were more abundant in the 1960s than in the
1950s or 1990s (p50.07). The most abundant species in the 1950s and
1960s were associated with old-fields (Peromyscus polionotus and Sigmodon hispidus), a habitat that declined from 20% to 0.07% of the SRS over
the past 50 years. The 1990s were dominated by the forest associated species Peromyscus gossypinus, Cryptotis parva and Neotoma floridana. The
only two species captured within the forested habitats during all time periods were Peromyscus gossypinus and Ochrotomys nuttali. Variance in
small mammal abundance differed among time periods and was highest in
the 1960s (p50.06). Relative abundance, in terms of occurrence in particular habitats relative to total abundance, of cotton mice increased in all
forested habitats except upland hardwoods, where they decreased in relative
abundance by 23%. Relative abundance of golden mice only increased in
the planted pine habitat (by 45%) and decreased in the upland and lowland
hardwood habitats (by 21 and 13%, respectively). Despite an increase in
abundance and variance in abundance in the decade following acquisition,
in the 1990s small mammal communities had changed little as compared
to 40 years earlier.

Intestinal parasites can directly affect their hosts by causing mortality or

sterility, or indirectly by changing behavior, increasing susceptibility to
predation or modifying competitive interactions. Recent studies have shown
that these effects can have substantial impacts on host population mortality
and recruitment. Here we investigate how intestinal parasites can directly
affect host survival and fecundity, and may also change activity patterns
and behavior. In wild populations of the white-footed mouse (Peromyscus
leucopus) at the Mountain Lake Biological Station, we removed intestinal
nematodes using Ivermectin from individuals and estimated the effects on
survival, fecundity and activity patterns. We followed treated and control
mice throughout the summer and measured survival, fecundity, intestinal
parasite intensity and diversity, and changes in behavior. Intestinal parasites
were shown to have a significant effect on activity patterns such as horizontal and vertical movement in the white-footed mouse. In addition, Ivermectin treated mice had lower intestinal nematode prevalence, diversity
and intensity. These results may have important implications for the impact
of intestinal nematodes on wild mouse population dynamics.

PECOT, STEPHEN D.,1,* ROBERT J. MITCHELL,1 BRIAN J. PALIK,2

ROBERT H. JONES3 and PAUL P. MOU.4 1 Joseph W. Jones Ecological
Research Center, stephen.pecot@jonesctr.org, robert.mitchell@jonesctr.org,
Newton, GA, USA; 2 USDA Forest Service, North Central Research Station, Grand Rapids, MN, USA; 3 Virginia Polytechnic Institute and State
University, Blacksburg, VA, USA; 4 University of North Carolina, Greensboro, NC, USA. Overstory retention in longleaf pine forests: resource
effects and regeneration responses.
Recent interest in variable overstory retention silviculture has resulted from
a desire to mimic the complex structure resulting from natural disturbances.
However, overstory retention benefits are tempered by increased competition between the overstory and seedlings. To investigate these competitive
interactions in longleaf pine ecosystems we used treatment plots 2 ha in
size (with three replications) that were harvested to similar residual basal
areas using three removal techniques: singletree, small group (;0.08 ha),
and large group (;0.18 ha) selection. Uncut control plots were also established. We sampled resources (light, soil water, and nutrients) as well as
seedling growth and survival throughout the range of overstory density in
each plot. Annual percent light transmittance varied greatly (35 to 80% of
the maximum). Light reaching the understory was closely related to overstory abundance (r250.76) and was predicted well using hemispherical
photographs (r250.77). Available nitrogen (measured with ion exchange
resin membranes) greatly increased only in open areas where the understory
was controlled (r250.59). Pine root production compensated for reduction
in non-pine roots if understory vegetation was removed (P,0.001); however, in large gaps, non-pine roots quickly filled root gaps if the understory
was left intact (P50.015). Seedling growth was closely related to overstory
density (r250.55). Large increases in resource availability in open areas
(.65% gap fraction) suggest that grouped removal patterns facilitate greater seedling growth through higher light and N availability (though at a cost
through increased belowground competition from nonpine roots) as well
as less light captured by the overstory. These data argue that manipulation
of the spatial distribution of overstory trees may allow for land managers
to retain the benefits of overstory retention while allowing for regeneration
of longleaf pine.

264

Abstracts

PEDERSEN, BRIAN S.* and SARAH L. PEARS.* Dickinson College,

Carlisle, PA, USA. Teaching simulation modeling from the ground up.
Simulation models, mathematical representations of real-world processes
implemented on computers, are increasingly being included in undergraduate ecology and environmental science courses. Working with simulation
models allows students to (1) further develop and integrate their understanding of biological, chemical, and physical processes as they apply to
ecological and environmental problems; (2) learn by conducting realistic,
long-term "experiments" that would not otherwise be feasible in a typical
course; and (3) gain direct experience with the merits and limitations of a
scientific tool widely used to address complex challenges, such as global
change. Undergraduate ecology and environmental science courses that include simulation modeling commonly rely on pre-developed models or on
software tools designed for model development. This is understandable
given that the students in these courses often have limited training in mathematics and computer programming. However, students who work with
pre-developed models or modeling tools may not gain a deep understanding
of how models work. At worst, students may experience a model as a
"black box" and not grasp the relationship between the model and the real
world. We (a teacher and an undergraduate student) describe a simulation
modeling course for undergraduates. There are no mathematics or computer
programming prerequisites. Students learn to model using only the basic
operations available in a standard spreadsheet program. Models are constructed around the finite difference equation. Students initially work with
and modify pre-developed models, and then go on to develop their own
models starting from a blank spreadsheet. Past student models include simulations of the fate of a bioaccumulating pollutant in a population subject
to harvest, the Earths energy budget, and the dynamics of a migrating
herbivore interacting with its food supply. The course has been taught at
two liberal arts colleges. We invite discussion of the effectiveness of alternative methods for teaching simulation modeling.
PEDERSON, NEIL,* GORDON JACOBY and ED COOK. Tree-Ring Laboratory, Lamont-Doherty Earth Observatory, Columbia University, Palisades, NY. Recent growth increases of northern range margin trees:
Disturbance, climatic, and site interactions.
Climate change has increased high-latitude treeline growth in Northern
Hemisphere forests over the last century. An important question to ask is
"have eastern US mid-latitude trees experienced similar changes?" These
trees provide fiber products, aesthetics for ecotourism and comprise one of
the largest aboveground global carbon pools. Growth of northern rangemargin trees, a species treeline, is a good place to address this question.
Here we focus on several populations of Carya glabra, Chamaecyparis
thyoides, Pinus rigida, Quercus prinus, and Q. rubra. At least 20 trees of
each population were cored along a northerly transect in the Hudson River
Valley of NY and NJ. Sampling was limited mostly to forests with little

recent disturbance. Growth rings were crossdated, measured and combined

to create two annual chronology types, average raw and relative ring
widths. Four species had consistent recent growth increases across populations, C. thyoides, P. rigida, Q. prinus, and Q. rubra. Populations with
recent increased growth tended to be positively correlated to temperature.
Site characteristics along with proximity to a northern range-margin seemed
important for temperature sensitivity, although this has yet to be quantified.
All C. thyoides populations had increased growth rates over the last 40
years. Two Q. prinus populations had increasing raw ring widths after being
in the canopy for more than 100 years, contrary to typical ring width trends.
Nearly one third of the rings more one standard deviation above the postcanopy accession mean have occurred since 1982. Both populations had
an average DBH greater than 0.5 m and many individuals 280 to .425
years old. Wider rings around such large trees indicate a substantial increase in biomass and carbon uptake. Age was not a limiting factor in
growth. Our initial research indicates that climate, perhaps interacting with
site factors, and competition has spurred biomass production.
PEEK, MICHAEL S.,1,* JOSH LEFFLER,1 CAROLYN Y. IVANS,1,2 RON
J. RYEL1 and MARTYN M. CALDWELL.1 1 Department of Forest, Range
and Wildlife Sciences and The Ecology Center, Utah State University, Logan, UT; 2 Department of Biological Sciences, Eastern Kentucky University, Richmond, KY. Fine root distribution and life-span in three cooccurring Great Basin life forms.
Plant investment below-ground is considerable, particularly in arid environments as evidenced by high root:shoot ratios. Low water and nutrient
availability contributes to the large investment in root structures in these
systems. Fine roots play a major role in acquiring these essential resources,
yet a paucity of information exists on the distribution and longevity of fine
roots. Minirhizotrons provide a valuable means of non-destructively examining fine roots over time. We examined fine roots in three co-occuring
Great Basin plant species of contrasting life form; the perennial shrub Artemisia tridentata, the perennial grass, Agropyron desertorum and the invasive annual grass Bromus tectorum. Fine root length, diameter and lifespan were followed for each species from a depth of 5 cm to 120 cm
during the 2001 and 2002 growing seasons. All species exhibited flexibility
in fine root length distribution with depth. In 2001, a year of reduced winter
precipitation relative to 2002, fine roots were found deeper in the soil
profile. In 2001, Bromus had 75% of total root length reach a depth of 45
cm, whereas in 2002, 75% of the root total length only reached a depth of
30 cm. Agropyron and Artemisia showed similar trends with 75 % of total
root length reaching a greater depth in 2001 (43 cm and 50 cm, respectively), than in 2002 (35 and 37 cm, respectively). Mean fine root diameter
(mm) was not significantly different between the two perennials in either
year (; 0.26), but fine roots were significantly smaller for the annual
(0.2160.004, 2001; 0.2360.004, 2002). Surprisingly, mean diameter did
not positively correlate with fine root life-span. Artemisia, with the largest
fine root diameters, had had shorter life spans in both years than the two
grasses. Interestingly, there was apparent persistence of fine roots after seed
set and shoot senescence of the annual, Bromus tectorum. Flexible rooting
distributions combined with persistent fine roots may facilitate water and
nutrient acquisition in these nutrient-poor environments.
PEET, ROBERT K.1 and RICHARD P. DUNCAN.2 1 University of North
Carolina, peet@unc.edu, Chapel Hill, NC, USA; 2 Lincoln University, duncanr@lincoln.ac.nz, Canterbury, New Zealand. Constructing templates
for restoration of longleaf pine ecosystems based on groundcover biodiversity.
Vegetation restoration projects present practitioners with the problem of
determining appropriate goals or targets when attempting to restore the
structure, composition, and biodiversity of the original vegetation on highly
modified sites. Longleaf pine ecosystems are especially challenging in that
examples of natural vegetation have become rare, remnant patches are
quickly degraded by fire suppression, local diversity is often high, and there
is substantial spatial turnover compared to other North American ecosystems. When remnants of natural vegetation persist elsewhere, one approach
is to model the relationship between the composition and structure of the
remnant vegetation and the physical and spatial characteristics of the sites
on which it persists. The model can then be used as a template to predict

the original structure and composition of vegetation on sites where the

original plant community is gone, but the physical characteristics of the
site can be measured. We present a template for reconstructing the original
longleaf pine vegetation of the fall-line sandhills, southeastern United
States, using vegetation, soil, and spatial position data from 188 plots located in remnant patches of natural vegetation throughout the region. The
plots clustered into eight vegetation series that were further resolved into
19 communities. Linear discriminant function analysis correctly predicted
the plant community at a site from the physical characteristics of the site
with a 64% success rate. We modelled the relationship between species
number and plot area in each community using species numbers recorded
in a series of nested quadrats in each plot. Finally, we combined data on
predicted community type and species number to generate lists of species
likely to be found in areas targeted for restoration. We illustrate this approach by comparing actual with predicted fire-maintained pineland vegetation on several sites near Aiken, SC.
PEGORARO, EMILIANO, RAMESH MURTHY, GREG A. BARRONGAFFORD, KATIE GRIEVE and JOOST VANHAREN. Biosphere 2 Center, Columbia University, Oracle, AZ, USA. Effect of CO2 and VPD on
isoprene emissions during drought in a Populus deltoides ecosystem.
From July to December 2002 we studied the effect of CO2 and vapour
pressure deficit (VPD) on isoprene emission from a cottonwood forest during a drought experiment in the Intensive Forestry Mesocosm at the Biosphere 2 Center. We measured leaf and ecosystem isoprene emission rates.
Photosynthesis, stomatal conductance, VPD, and soil water content were
also monitored. Difference in isoprene emission rates between elevated CO2
and ambient CO2 grown trees decreased as the drought progressed. Although isoprene emission rates decreased when water stress was severe,
they were considerably less sensitive to drought than photosynthesis and
stomatal conductance. As a result, the percentage of carbon lost as isoprene
increased as they became more water stressed reaching carbon losses of 11
%. Results also showed that during the drought isoprene biosynthesis used
recently fixed carbon, and this decreased when it was limited. We also
observed that the inhibitory effect of elevated CO2 on isoprene emission
was compensated by the decrease in intercellular CO2 concentration (Ci)
caused by the high VPD treatment. Results show the strong negative relationship between isoprene emission rate and Ci in all CO2 treatments. The
existence of a soil sink for isoprene was also investigated. Biosphere 2
Centre is a unique facility for the studying the response of entire ecosystems to selected environmental variables.
PELLATT, MARLOW G.,1,2,* ANITA HOLTHAM3 and DARREN BOS.1,2
1
Parks Canada, Western Canada Service Centre, 300-300 West Georgia
Street, Vancouver, British Columbia, Canada; 2 Simon Fraser University,
School of Resource and Environmental Management, Burnaby, British Columbia, Canada; 3 Queens University, Department of Biological Sciences,
Kingston, Ontario, Canada. Paleoecology of salmon lakes on the west
coast of Canada.
Parks Canada has a mandate to maintain the ecological integrity of terrestrial and marine ecosystems in representative regions within Canada and
its national waters. In order to understand the ecological integrity of any
given ecosystem it is essential to understand the natural range of variability
in which the system exists as well as the impact people and climate change
have had on it. This study is a multiproxy investigation that uses paleoecological methods to reconstruct environmental conditions for the last
;300 years in three lakes and their watersheds on the west coast of Vancouver Island. Pollen, diatoms, cladocera, sediment, and stable isotope
analyses of 210PB dated sediments from lakes in or near Pacific Rim National Park Reserve of Canada reveal changes in forest structure, limnological conditions, and possibly salmon populations. We also present data
that investigates differences in sedimentary d15N amongst a suite of salmon
lakes from British Columbia and relate these differences to the terrestrial
sedimentary component and other environmental variables. The results of
this research program contribute to our understanding of the role of marine
derived nutrients in regard to lake productivity along the west coast of
North America. This research indicates that while salmon derived nutrients
may be of key importance in juvenile salmonid development in some lakes,
this may not be the case in all systems, especially those in which flushing

Abstracts

265

rates are high. In these systems diatom communities appear to be more

sensitive to fluctuations in salmon populations than stable isotope methods,
provided that other changes in trophic status are minor. The role of this
type of research in ecosystem management is important to Parks Canada
and will assist in understanding the complexities of land sea interactions
and the interplay between terrestrial and marine protected areas.
PENDALL, ELISE,1,* JENNIFER Y. KING2 and ARVIN R. MOSIER.3
University of Wyoming, Laramie, WY; 2 University of Minnesota, St.
Paul, MN; 3 USDA-ARS, Fort Collins, CO. Dynamics of soil organic
matter turnover under elevated CO2.
1

Some grassland ecosystems have been shown to store carbon as a result of

exposure to elevated CO2, but dynamics of soil organic matter (SOM) turnover rates are expected to vary with time and contribute variability to net
ecosystem production estimates. A 5-year-long elevated CO2 experiment
was conducted on the shortgrass steppe in Colorado using open-top chambers (OTCs). Above- and below-ground biomass increased by 15 to 35%
under doubled atmospheric CO2 concentrations. Decomposition rates also
increased, as indicated by stable isotope partitioning of soil respiration
rates. We collected soil samples from the OTC experiment after 1, 2, and
3 years of exposure to elevated CO2 to investigate the early response of
SOM pool sizes and turnover rates. Long-term laboratory incubations (up
to 1 year) were used to investigate changes in active and slow pool sizes
and turnover rates, and stable C isotopes were measured to evaluate changes in turnover rates of recently fixed carbon. After just one year, we found
significant increases in active and slow pool sizes and turnover rates of
both SOM pools. However, the increases diminished somewhat over the
next two years. The turnover rate of new active pool SOM (determined by
isotope labeling) was significantly lower under elevated than ambient CO2,
suggesting that increasing litter C:N ratios under elevated CO2 will eventually slow decomposition rates, and potentially limit N availability. Microbially respired CO2 had significantly lower d13C values than did bulk
soil C, owing to an increase in C3 biomass on all treatments. The use of
fossil fuel CO2 in the elevated CO2 treatment added an additional 13C depleted tracer to the ecosystem. After 3 years of exposure to elevated CO2,
d13C of bulk soil C decreased by 2 permil, while d13C of microbial C
decreased by 10 permil. These findings suggest that net ecosystem C storage is likely to change over time, possibly increasing with continued exposure to elevated CO2.
PENDLETON, ROSEMARY L.,1 BURTON K. PENDLETON,1,* JEFFREY R. JOHANSEN2 and LARRY L. ST. CLAIR.3 1 Rocky Mountain
Research Station, USDA Forest Service, 333 Broadway SE, Suite 115,
Albuquerque, New Mexico, USA; 2 Department of Biology, John Carroll
University, University Heights, Ohio, USA; 3 Department of Integrative
Biology, Brigham Young University, Provo, Utah, USA. Soil microorganisms affect survival and growth of shrubs grown in competition with
cheatgrass.
Invasive annual grasses such as cheatgrass (Bromus tectorum) create changes in soil microorganism communities and severely limit shrub establishment. We examined the effects of biological crust-forming algae and arbuscular mycorrhizal fungi on growth and survival of five western shrub
species with and without competition from cheatgrass. Germlings of Ephedra viridis (EPVI), Coleogyne ramosissima (CORA), Artemisia filifolia
(ARFI), Chrysothamnus nauseosus(CHNA), and Artemisia nova (ARNO)
were planted into pots with and without fertilizer additions. We used four
inoculation treatments; algal crust inoculum of the genus Schizothrix, arbuscular mycorrhizal inoculum of spores extracted from soil collected from
a mixed shrub location, dual inoculation, and a non-inoculated control. Half
of the low fertility pots were planted with one seed of cheatgrass. Shrub
survival declined as soil fertility increased. Few shrubs were able to survive
in competition with cheatgrass in fertilized growth medium. Algal inoculation increased shrub survival. Soil fertility level and competition with
cheatgrass significantly affected growth of all five shrub species. At high
fertility, the faster-growing shrubs grew best with no microorganism additions. The slower-growing CORA grew best in the presence of mycorrhizae and EPVI in the presence of the algae. Under low nutrient conditions, all shrub species responded positively to the addition of mycorrhizal
inoculum. CHNA and EPVI responded positively to the algal inoculum.

266

Abstracts

The addition of mycorrhizal inoculum intensified competition with cheatgrass, reducing shrub shoot biomass over that of the control treatment in
all species except ARFI. However, shoot growth in cheatgrass was reduced
to an even greater extent in all shrubs except ARNO. Algal inoculation
appeared to beneficially affect the growth of ARFI and EPVI at the expense
of cheatgrass. Fire or other disturbance that results in a nutrient flush can
put native species at a competitive disadvantage with cheatgrass. Our findings suggest that soil microorganisms can, to some extent, mitigate this
disadvantage.
PENG, CHANGHUI,1,2,* XIAOLU ZHOU,2 QINGLAI DANG2 and JIAXIN CHEN.2 1 Institute of Atmospheric Sciences, South Dakota School of
Mines & Technology, Rapid City, SD, USA; 2 Faculty of Forestry and the
Forest Environment, Lakehead University, Thunder Bay, Ontario, Canada.
Quantifying carbon sequestration of the Lake Abitibi Model Forest in
Ontario.
Boreal forest ecosystems play a significant role in the global carbon cycles
and are sensitive to global climate change, especially to the changes in
temperature and precipitation over the next century. In this study, we use
a process-based forest growth and carbon dynamic model of TRIPLEX to
simulate carbon dynamics in boreal forest ecosystems of the Lake Abitibi
Model Forest (LAMF) in northern Ontario, Canada. The simulations are
performed not only for forest structure (growth and composition) but also
for ecosystem function (including productivity, carbon, water, and nitrogen
cycles) in order to investigate the dynamics of carbon stocks and fluxes,
which are dependent on the annual change of temperature, precipitation,
soil texture and moisture, and nitrogen dynamics. The model input data
sets include climate conditions, soil initial carbon, tree mortality, and tree
form; The model outputs provide about 50 variables including each part of
biomass (stem, foliage, coarse and fine root), soil carbon (slow and passive
pool), and growth and yield (tree density, height, DBH, and volume). More
than 40,000 stands are simulated from 1990 to 2000. The input and output
of these stand polygons are processed using GIS technology to integrate
different spatial themes for the LAFM. Our results suggest that total net C
sequestrations through forest growth (NPP) were estimated at about 1.35,
1.36, and 1.39 Mt C for 1990, 1995 and 2000, respectively. The total
biomass carbon stocks were 35.0, 36.8 and 40.1 Mt C for the years of
1990, 1995, and 2000. The net balances of carbon sequestrations were
between 0.39 and 0.41 Mt C without the harvest removal, and the net
biome production (NBP: remainder of carbon balance after harvesting) was
approximately 0.27 Mt C/year during l990-2000, suggesting that the LAMF
forest ecosystem was acting as a carbon sink (about 0.4 Mt C) in 1990s.
PENNINGS, STEVEN C.,1,* MARK D. BERTNESS,2 PATRICK J.
EWANCHUK,2 ELIZABETH R. SELIG2 and LETISE T. HOUSER.2 1 Department of Biology and Biochemistry, University of Houston, Houston,
TX, USA; 2 Department of Ecology and Evolutionary Biology, Brown University, Providence, RI, USA. Temporal and geographic variation in
positive and negative interactions among salt marsh plants.
A pressing problem for ecologists is determining whether our understanding of communities, often developed from work at fine scales, can explain
processes across broader scales. Here, we explore whether models of positive interactions developed in southern New England can be applied at
geographic scales. Salt marsh plants may interact positively by ameliorating
harsh physical conditions such as salinity stress. Because salinities are higher in low- versus high-latitude marshes, and in dry versus wet years, we
tested the hypotheses that positive interactions would be increasingly important at low latitudes and in dry years. We conducted parallel experiments
in Maine, Rhode Island, Georgia and Alabama. Multiple species of salt
marsh plants were transplanted into control plots or plots with neighboring
vegetation removed, in multiple marsh zones, at two sites/state. The experiments were repeated over 3-4 years. Our hypotheses were supported at
the regional scale. Within New England, positive interactions were rare in
Maine but common in Rhode Island, and were more important in dry than
wet years. At the geographic scale, our hypotheses were not supported.
Positive interactions were not more important in southern (Georgia, Alabama) than northern (Maine, Rhode Island) sites. Rather, most interactions
in southern sites were competitive. The most likely explanation for our
results is that northern floras were dominated by salt-sensitive species that

were likely to be facilitated by neighbors, whereas southern floras were

dominated by salt-tolerant species that were unlikely to benefit substantially
from neighbor-amelioration of salinities. Intraspecific variation in salt-tolerance may also have contributed to a reduced occurrence of facilitation at
southern sites. These results illustrate the difficulties inherent in extrapolating results from even well understood systems to different geographic
locations or scales. Intraspecific adaptations and community composition
will often vary geographically, complicating our efforts to construct geographically robust generalities about community structure and processes.
PERFECTO, IVETTE,1,* RUSSELL GREENBERG2 and GUILLERMO
IBARRA-NUNEZ.3 1 University of Michigan, Ann Arbor, Michigan;
2
Smithsonian Migratory Bird Center, Washington, DC; 3 El Colegio de la
Frontera Sur, Tapachula, Mexico. Food web structure of the coffee agroforests: does biodiversity makes a difference?
The coffee agroecosystem in Northern Latin America is characterized by
a high floristic diversity. These agroforests serve as habitat for a high diversity and density of associated fauna, including many generalist predators. Does this diversity contributes to the reduction of insect herbivores,
and therefore, to the productivity and sustainability of the agroecosystem?
In our studies of the Mexican agroforests we examined the trophic relationships of bird, spiders, ants and herbivores. Through a series of exclosure experiments in coffee farms with different levels of shade and floristic
diversity, we have identified the significant trophic interactions in this very
diverse tropical agroecosystem. We will discuss the results from these experiments as well as the implications of these complex direct and indirect
interactions on the productivity and sustainability of the coffee agroecosystem.
PERKINS, ALISON E.H.,* CAROL A. BREWER and JOHN P. TWIGGS.
University of Montana, Missoula, MT, USA. What are your kids watching? Developing evaluation criteria for nature and science films and
videos.
Television provides an unparalleled opportunity for the public to learn
about science in a non-school environment. Recently cable channels have
expanded into niche markets in nature and science programs, once only
available on public television. Moreover, a considerable secondary market
exists for nature and science videos via general distribution and direct sales.
Although a large audience obtains science and nature "educational content"
via these media, no consistent and credible system exists to evaluate the
ecological content and educational value. Therefore, we developed criteria
to assess the educational value of science and nature films and videos using
the Guidelines for Excellence (NAAEE 1996) for environmental education
materials, and in consultation with educators and scientists familiar with
the constraints of film and television production. Criteria included: scientific context, factual correctness and freedom from misconceptions, unbiased interpretations, exploration/presentation of nature of science "issues,"
presentation of scientists, and structure for interpretation breaks. We tested
the system at an annual wildlife film festival held in Missoula, Montana,
which assigns awards in categories including scientific content, educational
value, presentation of animal behavior, and conservation message. Approximately 25% of the 200 films entered are presented to the public as award
winners; therefore we were able to evaluate productions that were specifically targeted as viewer-chosen education opportunities. A panel of ecologist judges rated festival films using our criteria to assess merit. Ratings,
along with explanations, were translated into the A, B, C, D, and F grading
system to be easily recognized by educators. Because producers operate in
a market-driven system, we designed the final rating summary to offer
educational strategies for films or videos with poor presentation of concepts
but exceptionally well produced. We anticipate this system can be used to
promote collaboration between producers and ecologists to ensure productions with both excellent science and aesthetic appeal.
PETERS, HALTON A.,1,2,* HAROLD A. MOONEY1 and CHRISTOPHER
B. FIELD.2 1 Department of Biological Sciences, Stanford University, Stanford, CA; 2 Carnegie Institution of Washington, Department of Global Ecology, Stanford, CA. The response of feeding patterns to climatic changes: Generalist herbivores in an annual-dominated California grassland.
This study explores the influence of future climates on the feeding patterns
of generalist herbivores in California annual grassland. Any climate-change

induced modification of herbivore feeding tendencies may alter competitive

outcomes in plant communities, resulting in shifts in plant species composition. Such modifications include changes in the amount of plant material ingested or in the relative proportions of food plant species consumed.
Partly because most studies on plant-herbivore interactions in response to
changing climates have used single plant species feeding trials in which
host switching was not possible, few data are available on changes in herbivore preferences in response to changing climates. In this study, generalist herbivores (land slugs) were fed leaf material from naturally occurring
plant communities that had been grown under one of the following climate
change treatments: 1) elevated atmospheric CO2; 2) increased nitrogen deposition; 3) increased precipitation; or 4) increased temperature. Herbivore
consumption levels were altered by climate change treatments, with implications for plant and herbivore population sizes in future climates.
PETERSEN, JOHN E.,1,* LINDSEY SCHROMEN-WAWRIN1 and W. MICHAEL KEMP.2 1 Oberlin College, Oberlin, OH; 2 University of Maryland,
Cambridge, MD. Water exchange, nutrients and "top-down" vs. "bottom-up" limitation of plankton: Mesocosm and modeling studies.
The magnitude and quality of biological, material and energetic exchange
are recognized as key factors governing the dynamics of coastal ecosystems. Additionally, ecologists invoke two distinct mechanisms to explain
organism abundance: "bottom-up" control by resource availability and "topdown" control by organism removal (i.e. herbivory, predation and other
removal mechanisms). The situation is further complicated by the fact that
organisms at different trophic levels often have very different generation
periods (e.g. in our system producers reproduce in 1 d, and primary
consumers in 10 d). We used mesocosm experiments and developed a simulation model to explore how water exchange rate and nutrient concentration govern trophic interactions to control the abundance of phytoplankton
and zooplankton. Specifically, experiments were conducted in 1 m3 mesocosms with 0%, 10% and 30% d-1 water exchange using both high and
low nutrient influent waters. We found that with high nutrient influent,
phytoplankton abundance and primary productivity increased with increasing exchange while zooplankton abundance declined. In contrast, with lownutrient influent waters, zooplankton, phytoplankton and primary productivity all increased with increasing exchange. Modeling studies corroborate
the hypothesis that a transition in the effects of exchange occurs under
these two scenarios: in the high nutrient experiment primary producers
were controlled from the top-down by zooplankton grazing and zooplankton abundance was, in turn, controlled from by the rate at which these
organisms washed out of the mesocosm. In contrast, in the low nutrient
experiment phytoplankton were controlled from the bottom-up by availability of nutrients, and zooplankton were controlled from the bottom-up
by phytoplankton availability. The complexity of possible interactions even
in this relatively simple system points towards challenges and opportunities
for interpreting dynamics in natural coastal ecosystems and the potential
value of mesocosms and simulation models for interpreting such dynamics.
PHILPOTT, STACY M.* University of Michigan, Ann Arbor, MI. A behaviorally-modified trophic cascade: Parasitic flies affect ants and ant
prey in coffee agroecosystems.
Trophic cascades exist in numerous terrestrial systems, including many
systems with ants as the top predator. In particular, behavioral modifications
of herbivores may be important in mediating trophic cascades. Although
most studies of trophic cascades focus on predator-herbivore-plant links,
trophic cascades could be applied to almost any level of trophic interactions. I consider interactions between the parasitic phorid fly, Pseudacteon
sp. (Diptera: Phoridae), its ant host, Azteca instabilis (Hymenoptera: Formicidae), and the herbivore, Spodoptera frugiperda (Lepidoptera: Noctuidae) as a potential species-level trophic cascade in coffee agroecosystems.
I specifically tested the hypotheses that 1) Phorid flies modify the behavior
of their ant hosts reducing ant recruitment (cooperative foraging behavior)
and 2) Phorid flies, via their negative influences on ants, indirectly positively affect ant prey. I examined ant recruitment by counting ants at baits
every 2 min for 30 min on trees with or without phorids. To assess phorid
influences on ant prey, I introduced lepidopteran larvae onto coffee plants
and recorded ant affects on larvae either with or without phorids. Phorids
restricted ant recruitment (foraging behavior) by halving ant numbers at

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267

tuna baits (ANOVA, P , 0.043, F . 4.331, df 5 1). With phorid flies,

some larvae escaped ant attack altogether (Chi-square test, P 5 0.004, df
5 1), and it took ants on average more than twice as long to carry away
larvae (ANOVA, P 5 0.033, F 5 4.720, df 5 1) or to force larvae off
plants (ANOVA, P 5 0.006, F 5 8.092, df 5 1). I thus conclude there is
a behaviorally-modified species-level trophic cascade in the coffee agroecosystem where phorids limit ants, and indirectly benefit ant prey. Additionally, phorid flies may have further important effects in ant and herbivore
communities as well as for coffee production.
PIPPEN, JEFFREY S.,1,* WILLIAM H. SCHLESINGER,1 MATTHEW D.
WALLENSTEIN,1 DEBRA M. KLEPEIS2 and BRUCE E. MAHALL.2
1
Nicholas School of the Environment and Earth Sciences, Duke University,
Durham, NC, USA; 2 Department of Ecology, Evolutionary and Marine
Biology, University of California, Santa Barbara, CA, USA. Community
composition and photosynthetic rate of photoautotrophs under quartz
pebbles in the southern Mojave Desert, California.
We used 16s rDNA sequences to identify novel species of cyanobacteria
beneath translucent quartz pebbles in the desert pavement on an alluvial
piedmont of the Coxcomb Mountains in the southern Mojave Desert, California, USA. Transmission of light, as measured with an integrating sphere,
was about 0.08% beneath the thickest pieces of quartz (25 mm) harboring
these hypolithic autotrophs. The photosynthetic rate ranged from 0.1 to 1.0
micromol m-2 sec-1 in the linear range of its response to light (PAR of 050 micromol m-2 sec-1), over which the apparent quantum use efficiency
was 0.019. Light-saturated rates of 1.7 to 2.7 micromol m-2 sec-1 were
recorded at light intensities of 200 to 400 micromol m-2 sec-1. The hypolithic community has an upper thermal tolerance of . 90 C in laboratory
conditions. The quartz pebbles confer a modest greenhouse effect that may
be important for photosynthetic activity during cool, wet wintertime periods
that prevail in the Mojave Desert.
PITHER, JASON,* MICHELLE COOK and LONNIE W. AARSSEN. Department of Biology, Queens University, Kingston, Ontario, Canada. Patterns of taxonomic diversity among freshwater lakes: Sensitivity to
scale, productivity gradients, and regional geographic influences.
Examining regional patterns of diversity within a spatially explicit framework adds insight to several current lines of inquiry in ecology and biogeography: local versus regional influences on site diversity; ecological
versus evolutionary influences on site diversity; and the relationship between productivity and diversity. We explored patterns of taxonomic diversity using a dataset of 239 north-eastern U.S. freshwater lakes, sampled
as part of the Environmental Monitoring and Assessment Program. The
dataset is ideal for this research, owing in particular to its broad coverage
and sampling design. Using matrices of environmental conditions (e.g. productivity, pH), species tolerances (niches), spatial co-ordinates, and an
index of the lakes environmental uniqueness, we are examining the spatial
details of diversity of diatom taxa in the contexts of the above lines of
inquiry. Preliminary results have revealed that, consistent with recent published work, dissimilarity in species composition among lakes increases
along the productivity gradient, and correspondingly, the proportion of the
species pool represented by lakes decreases with increasing productivity.
This pattern holds after accounting for geographical proximity. Water pH,
however, explains regional patterns of alpha and beta diversity better than
productivity for diatoms. A scale-sensitivity analysis of the patterns revealed that most trends in richness become apparent only at the coarsest
scales, while others rear their head at within-ecoregion scales. As expected,
patterns are also sensitive to the taxon: zooplankton and fish exhibit different scale-sensitivities. We developed an analytical approach to test for
scale-sensitivities that uses generalized linear mixed models and randomization procedures, and suggest this as an appropriate framework for future
analyses of geographical patterns of diversity.
EIRO, GERVASIO,1,* JOSE M. PARUELO,1 ESTEBAN G. JOBBAPIN
GY,1 MARTIN OESTERHELD1 and ROBERT B. JACKSON.2 1 pineiro@ifeva.edu.ar, University of Buenos Aires, Cap. Fed., Buenos Aires, Argentina; 2 Duke University, Durham, North Carolina. Long term grazing
impacts on carbon and nitrogen soil particle size fractions in South
American grasslands.
We evaluated the effects of grazing on grasslands of South America using
CENTURY, a process based biogeochemical model and field experiments.

268

Abstracts

Our simulations suggested that grazing by domestic herbivores accelerate

nitrogen cycling and increases nitrogen losses, mainly due to increases in
nitrogen volatilization and leaching from urine and feces patches. The lower nitrogen availability under grazing limited soil organic matter formation
and resulted in a mean reduction of 22% in soil organic carbon (SOC)
content after 400 years of grazing (since introduction of cattle by European
people in the region to the present). The model suggested that these losses
should be higher, and hence detectable, in the more labile SOC pools (33%)
and less (5%) in the large passive SOC pools. To test this hypothesis, we
selected 7 grazing-exclosure sites in the Rio de la Plata grasslands of Uruguay. We sampled soil and roots and measured carbon and nitrogen contents at six depths in two different soil size fractions: the particulate organic
matter (POM) of rapid turnover and the mineral associated organic matter
(MAOM) of low turnover. Unlike other Northern-hemisphere experiments
(with increased nitrogen depositions form industry), our results show that
for the first 30 cm grazing reduces total SOC and alters vertical distribution
of POM, increasing the more labile SOC fractions towards the surface.
Vertical changes of POM can be associated with the higher root contents
in the first 5 cm of depth under grazing.
PLATT, BILL, SUSAN CARR and MATT REILLY. Using concepts for
old-growth pine stands to enhance biodiversity in longleaf pine savanna
groundcover.
Old-growth longleaf pine stands contain variable sized openings without
trees. Many second-growth stands contain dense pines with few openings.
Restoration logging (group selection of patches of trees) that generates
open spaces in such stands could enhance ground cover, but ground surface
disruption potentially could produce negative effects and increased invasion
by exotic species. We studied effects of restoration logging on groundcover
in eastern Louisiana pine savannas. In each of 8 drainages in fertile loess
plains savannas at Camp Whispering Pines, we randomly established three
permanent plots: 1) restoration logged five years earlier, 2) open with no
overstory trees for .30 years, and 3) no overstory trees removed. We
sampled numbers of species of three life forms (graminoids, shrubs, forbs)
in nested plots from 0.01m2 to 100m2. Abundance classes (common, intermediate, infrequent) were assigned to species based on relative occurrences
in plots. MANOVA analyses indicated that significant treatment effects
(overstory trees present or absent; overstory tree removed recently or decades previously) were attributable to forbs; intermediate and infrequent
forbs were more abundant when overstory trees were absent, especially if
absent for decades. At Abita Springs Preserve, we randomly established
nine plots in pine flatwoods, five with overstory pine removal and four
with no pine removal. We sampled numbers of species in nested permanent
subplots from 0.01m2 to 1000m2 before and for three years after logging.
Repeated measures MANOVA indicated significant effects of overstory
removal; effects were attributable to increased numbers of graminoids, primarily sedges. These two studies suggest that ecologically sensitive restoration logging can enhance groundcover biodiversity; less common species increase in abundance, but different life forms may respond in different
habitats. Moreover three groups increased biodiversity at both sites: species
with a seed pool, species immigrating from surrounding areas of pine savanna, and species invading from different habitats.
PLOG, LAURA B.,* JOHN F. WEISHAMPEL, JAMES D. ROTH and
LLEWELLYN M. EHRHART. University of Central Florida, Orlando, FL.
Nutrient flow from sea turtle nests to dune vegetation.
Sea turtles are important biological transporters, bringing nutrients and energy from distant foraging grounds to nesting beaches through egg deposition. Because nesting beaches are typically nutrient-poor, an increase in
nitrogen (N) concentrations could be critical for maintaining dune vegetation and promoting stabilization from erosion. The 40.5 km stretch of beach
between Melbourne Beach and Wabasso Beach, Florida, supports the
worlds third largest nesting population of loggerhead turtles (Caretta caretta) and the largest nesting population of green turtles (Chelonia mydas)
in the continental United States. Nest locations of both species exhibit
similar clinal patterns, ranging from approximately 100 nests per km in the
northern part of the beach to over 1000 nests per km in the southern portion
of the beach. To evaluate the contribution of nutrients from sea turtle eggs
to dune vegetation, we measured stable-nitrogen isotope ratios 15N/14N) of

turtle eggs and other marine sources (e.g., drift algae), which are typically
enriched in 15N over terrestrial sources, and compared these to stable-nitrogen isotope ratios of dominant dune plants. Plant samples were collected
along a gradient of nesting density parallel to the shore and at three locations on the dune (foredune, middune, and top of dune). Eggs of carnivorous loggerhead turtles were enriched in 15N by 5.0o/oo over eggs of herbivorous green turtles. Isotopic signatures of sea oats (Uniola paniculata)
were significantly higher in the area of high sea turtle nest density. Within
this area, sea oats at the foredune were enriched in 15N by 3.1o/oo over
plants at the top of the dune. Thus, sea turtle nesting may represent an
important biogeochemical link between marine and terrestrial environments.
POLLEY, WAYNE,1,* BRIAN J. WILSEY2 and JUSTIN D. DERNER.3
USDA-Agricultural Research Service, Temple, Texas; 2 Iowa State University, Ames, Iowa; 3 USDA-Agricultural Research Service, Cheyenne,
Wyoming. Plant species diversity in native and restored tallgrass prairies: Patterns and controls.
1

One goal of ecological restoration is to restore diversity of native vegetation, but mechanisms responsible for diversity in targeted communities often are poorly understood. We measured diversity (Simpsons index, 1/D)
of plant species and functional groups of species in replicated 0.5-m2 plots
within paired restored and relict tallgrass prairies at three locations in central Texas. To identify mechanisms responsible for diversity differences
between prairie types, we decomposed diversity into richness (S) and evenness (E; relative biomass) and into spatial (alpha, beta, gamma) components
and assessed relationships between species and functional group richness
in small plots. Species S was greater in native than in restored prairies at
the within-plot (alpha), among-plot (beta), and prairie (gamma) scales. Proportional contribution of alpha and beta S to gamma S did not differ between prairie types, indicating that prairies did not differ in proportional
scaling of S across space. Species diversity was greater in native than in
restored prairies at alpha and gamma scales. The beta component of diversity, however, contributed proportionally more to gamma diversity in
native than in restored prairies, implying that the importance of E to diversity changed with spatial scale. Indeed, although small-scale or plot E
did not differ between prairie types, E at the prairie scale was greater in
native than in restored grasslands. Plot-scale richness and evenness of functional groups (defined based on growth form and phenology) both were
greater in native than in restored prairies. Because ecological function
should differ more consistently between functional groups than between
species, the greater functional group richness and evenness observed in
small plots in native grasslands implies that niche differentiation contributes to the high diversity of native tallgrass prairie.

different life history strategies. However, the reasons for the absence of N.
dombeyi in forest canopy remain unclear. The importance of disturbance
in influencing patterns of Nothofagus forest regeneration suggests again
that a model emphasizing the response of individuals to a fluctuating abiotic/biotic environment is appropriate for understanding the dynamics and
coexistence of these forests. The replacement pattern after disturbance and
developmental history of stands studied were mediated by the significance
of life history differences. Interactions between natural disturbances and
sites lead to effects influencing composition and forest dynamics.
POLSENBERG, JOHANNA F.,* JOHN MCMANUS and FELIMON
GAYANILO. National Center for Caribbean Coral Reef Research
(NCORE), Rosenstiel School of Marine and Atmospheric Science
(RSMAS), University of Miami, Miami, FL. Interdisciplinary simulations of coral reef systems for science, education and management:
Results from the Florida Keys.
The establishment of a global set of comparative, long-term, interdisciplinary studies of coral reefs and reef-dependent people has been widely accepted throughout the coral reef community as an important approach to
understanding the complexity of interactions involved in coral reefs and
their management. It has similarly been widely recognized that decisionmaking for reef management is extremely complex, and that every effort
must be made to make relevant information readily accessible for managers
in a spatially-explicit Geographic Information System (GIS) format, supplemented with scenario-testing models. To initiate this process, we, along
with groups from Mexico, Belize, the Philippines, and Australia, have begun to analyze key reef processes using data from multi-scale remote sensing and interdisciplinary fieldwork on hydrodynamics, geology, ecology,
fisheries, human ecology and economics related to coral reef resilience and
management. We have developed initial GIS models for the Florida Keys
reef tract to integrate existing knowledge, identify information gaps, and
guide the gathering of further data. In order to test various interventions
or investigate a range of potential naturally-induced changes to a reef, these
framework models will be augmented with multi-layer, agent-based scenario-testing simulation models toward the creation of Dynamic Decision
Support Systems (DDSS). We will use a Component Assembly System
with the JavaBean as the component standard. A component assembly approach facilitates the adaptation of DDSSs to new reef systems without the
need for additional programming. We hope that these new computational,
simulation and data analysis methods and tools to model physical, ecological and social aspects of coral reefs will also contribute to improved environmental management approaches for all ecosystems.
PONOMARENKO, ELENA1,* and SERGUEI PONOMARENKO.2
ArchEco 5A-601 Laurier Ave.W., Ottawa, Ontario, Canada; 2 NatureServe-Canada, c/o Parks Canada Agency, Ecological Integrity Branch 25
Eddy Street, K1A 0M5, Hull, Quebec, Canada. Beavers as a trigger of
peat bog development.

POLLMANN, WILLIAM.1,2,* 1 University of Colorado, Boulder, Colorado,

U.S.A.; 2 Munster University, Munster, Germany. Disturbance history
and the coexistence of species in mixed floodplain forests, south-central
Chile.
Disturbance history and stand dynamics were investigated in two Chilean
Nothofagus forest stands affected by catastrophic floods, Andean Range,
south-central Chile, in order to explore regeneration patterns of various
Nothofagus species and examine the perspective of both coarse and finescale gap dynamics. Population and stand structure, and tree locations were
used to identify cohorts of trees that were initiated by flood events. This,
along with information on life history traits and spatial patterning, was
used to reconstruct the patterns of establishment of four dominant canopy
trees (Nothofagus alpina, N. dombeyi, N. macrocarpa, and N. pumilio) in
response to flooding disturbance. Intense floods formed large gaps and left
homogeneous surface deposits that became dominated by even-aged Nothofagus; however, stand history reconstruction analysis showed that smallscale gap formation may occasionally generate uneven-aged stands. Thus,
disturbance was critical to the maintenance of Nothofagus dispersion because it was the source of spatial heterogeneity in the physical environment
to which species responded differentially. Results confirm that dominance
of deciduous Nothofagus trees (e.g. N. alpina and N. macrocarpa) in forest
canopies appears to be locally associated with topographic depressions subject to cold air drainage. In the absence of the evergreen small-leaved N.
dombeyi canopy tree, deciduous Nothofagus appear to coexist by way of

In summer 2001, the driest year in decades, we had a rare opportunity to

dig through a hanging bog/fen in a mountain valley in Fundy National
Park, New Brunswick, Canada. The soil section crossed a number of peat
layers corresponding to several distinctive vegetation communities. A wetter and nutrient richer Grey Alder (Alnus rugosa) community at the bottom
was replaced by nutrient poorer and drier community dominated by Cinnamon Fern (Osmunda cinnamomea), and sphagnum mosses (Sphagnum
spp.). Above this layer, wetter and richer conditions reoccurred and a Grey
Alder community was restored. This community was replaced in turn by
a similar fern-sphagnum community that graded into an even poorer Picea
mariana / Chamaedaphne caliculata / Sphagnum spp. community that can
be now observed on the surface. If this stratigraphy was identified using a
peat borer or auger, it would be logical to interpret such a sequence of
layers as evidence of climate change that distorted a normal succession
from rich swamp to poor bog. That was also our first interpretation - that
there was a prolonged pluvial period between two drier periods. However,
a thorough field investigation of plant remnants revealed some problems
with this conclusion. In the middle of both Grey Alder layers we found
Red Pine logs with very wide annual rings, which would not be possible
for trees growing in wetland conditions. After additional searching we
found numerous alder branches with beaver cut marks in both alder layers

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269

that indicate they were burried beaver dams, and concluded that very ancient beaver activity was the principle driver for wetland creation and its
successional changes. The first dam was constructed across a fast mountain
brook about 7,600660 years BP and evidently caused swamping. The second is dated 4,860650 years BP.
PONOMARENKO, SERGUEI1,* and DONALD MCLENNAN.2
1
NatureServe-Canada, c/o Parks Canada Agency, Ecological Integrity
Branch 25 Eddy Street, K1A 0M5, Hull, Quebec, Canada; 2 Parks Canada
Agency, Ecological Integrity Branch, 25 Eddy Street, K1A 0M5, Hull,
Quebec, Canada. Applying the Canadian National Vegetation Classification to ecosystem classification and inventory updating in Canadian
national parks.
Parks Canada is presently updating ecosystem inventories and will be linking them to Canadian National Vegetation Classification (CNVC) approaches to provide a common national and international basis for ecosystem and
vegetation community classification. The CNVC is associated with the International Vegetation Classification (IVC) (Grossman et al. 1998) initiative, developed by NatureServe and the Ecological Society of America.
National parks represent the largest land base under federal jurisdiction,
and are relatively evenly distributed across Canada. Applying CNVC approaches for updating national park ecosystem inventories provides a standard basis for vegetation/ecosystem classification across the system of national parks, and provides an opportunity to introduce the CNVC to our
conservation partners across the country. A major update of ecosystem
inventories is required in many of Canadas national parks to develop a
long-term database for important conservation management functions. An
important requirement of updated ecological maps for national parks is that
mapped entities be correlated with similar inventories conducted by park
neighbours, regionally and internationally. The major challenges are to apply the CNVC vegetation units in the development of a site-based ecosystem classification, and to link to classifications presently used by provincial
and territorial neighbours. We have initiated a pilot project in Bruce Peninsula and Auyuittuq National Parks, where we have conducted vegetation
classification and ecosystem mapping that correlates with CFEC and CNVC
programs, and with ecological units common to neighbouring jurisdictions.
First year classification and mapping results will be presented and discussed.
POORTER, LOURENS.1,2,* 1 Forest Ecology and Forest Management
Group, Wageningen University, Wageningen, The Netherlands, The Netherlands; 2 Bolfor Project, Santa Cruz, Bolivia, Bolivia. Resprouting ability
of 50 rainforest tree species differing in adult stature and shade tolerance.
Interspecific variation in sapling survival is an important determinant of
the performance, shade tolerance, and dynamics of rain forest tree species.
As much as 25% of the understorey saplings are damaged annually by
falling debris. The ability of species to resprout allows them to survive in
the understorey, or regenerate successfully in treefall gaps. I tested the
hypothesis that the resprouting ability is closely linked to the shade tolerance and the adult stature of tree species by clipping the stem of saplings
of 50 rainforest tree species, leaving unclipped saplings as control. After 2
months, cut plants had a slightly higher mortality (2.6%) compared to control plants (0.5%). All cut plants that died were pioneers. Ninety percent
of the cut plants sprouted within two month, 6.5 percent formed buds, and
3.5% showed no response. The ability to resprout was not related to the
adult stature or shade tolerance of the species. Cut plants realized faster
height growth rates than control plants, indicating that stored reserves allowed for a compensatory growth response. The height growth rate of the
control plants was positively correlated to the light demand of the species.
In contrast, the height growth rate of the cut plants was positively correlated
to the adult stature of the species, indicating that large-statured species are
adapted to disturbance, and resprout quickly to continue their snail-race for
the canopy.
PORDER, STEPHEN,1,* PETER M. VITOUSEK,1 ADINA PAYTAN2 and
THOMAS BULLEN.2 1 Dept. of Biological Sciences, Stanford University,
Stanford, CA; 2 Dept. of Geological and Environmental Sciences, Stanford
Univeristy, Stanford, CA. Landscape development and ecosystem function: Results from the Hawaiian Islands.
Most landscapes are shaped by erosion and deposition, but the effects of
these processes on ecosystem function are not well understood. This limits

270

Abstracts

our ability to elucidate ecological and biogeochemical processes at a landscape scale. We examined the ecosystem effects of erosion along two toposequences in the Kohala Mountains, Hawaii, analyzing Sr and Ca isotopes and N, P and cation concentrations in vegetation. By comparing the
results to a well-studied, minimally eroded chronosequence, we conclude
that some, but not all, ecosystem properties along the slope were rejuvenated by erosion. For example, on 150 ky uneroded surfaces in the Kohala
mountains, 80% of Sr in vegetation is obtained from atmospheric sources,
while on slopes 80% of plant Sr is derived from local bedrock, similar
to the percentage on a 300 year old, uneroded site. This new supply of
rock-derived cations correlates with consistent changes in leaf Ca, but not
N or P. Sr to Ca ratios, which are positively correlated with Sr and Ca
isotopes along the chronosequence, are negatively correlated with isotopic
measurements from the toposequences. d44Ca correlates positively with Sr
isotopes along the chronosequence and on one slope, but not the other. We
conclude that the effects of erosion on ecosystem function are more complex than rejuvenation, and that eroding surfaces may be functionally different than stable surfaces of any age.
POSADA, JUAN M.,* STEPHEN S. MULKEY and KAORU KITAJIMA.
University of Florida, posada@botany.ufl.edu, Gainesville, Florida. Distributions of leaf carbon gain in the crowns of three tropical tree species.
In order to scale photosynthesis from leaves to whole trees it is necessary
to understand how light availability changes within tree crowns, and how
leaves respond to these changes. Leaves can acclimate to different light
environments by modifying the stoichiometry of the components of the
photosynthetic apparatus, but leaves can also influence their light environment by changing their orientation. Thus, there is a complex interaction
between the light environment, leaf physiology, and leaf orientation. The
objective of this study was to describe part of this interaction by measuring
light directly incident on leaves, and their corresponding photosynthetic
light responses. We selected leaves that covered a broad range of light
microenvironments within the crowns of three tropical canopy tree species.
Light was measured every two seconds for 10-17 days, using small GaAsP
photocells that were placed on the leaf surfaces. We measured photosynthetic light response curves in situ, at the end of the light measurement
period. The results indicate that all leaves, regardless of their position,
received significantly less light than available above the forest, and that
light was of low intensity most of the time. Consequently, the frequency
distributions of estimated leaf carbon gain peaked at low light intensity.
The peaks of the distributions were close to the point were leaves can have
the highest absolute carbon gain for a minimum light, i.e. the tangent of
the curve through the origin. We conclude by discussing the implications
of these results for the scaling of photosynthesis from leaves to whole
plants.
POST, DAVID M.* Yale University, New Haven, CT. Re-integrating body
size into the food chain-length discourse.
It has been clear since Eltons seminal work on animal ecology in 1927
that body size is central to the structure and function of food webs. Body
size is fundamental to energy flow and energetic efficiencies, species diversity, population densities, and nutrient cycling. Body size may also be
important for understanding variation in and limits upon food-chain length,
through which body size would have additional indirect effects upon community structure and ecosystem function. The vast majority of predatorprey interactions are size structured, such that predators are larger than
their prey or parasites are smaller than their hosts (although there are well
and long recognized exceptions such as social predators and venomous
organisms). The effects of body size on food-chain length should be greatest in food webs where predator-prey interactions are size structured. Yet,
with a few exceptions (e.g., Pimms design constraint hypothesis), explicit
consideration of body size has been conspicuously absent from the discussion of food-chain length. Here I present a theoretical framework for integrating body size into the food-chain length discourse. I investigate the
general mathematical theory, explore the heuristic value of the theory for
linking variation in food-chain length to environmental gradients such as
resource availability and ecosystem size, and discuss how the theory can
be evaluated and applied empirically within the context of size-structured
food webs.

POTEET, MARY F.* The University of Texas at Austin, Austin, TX. Sensitivity of parasite transmission to changes in host density following
disturbance.
Anthropogenic disturbance is widely implicated in altered patterns of parasitism in wildlife. Few empirical studies evaluate this claim and even
fewer assess the causal relationship between disturbance and parasitism.
The lack of data is not surprising. Many disturbances are unplanned and
unreplicated, making evaluation of disturbance-induced changes in hostparasite systems difficult at best. I study a 1-parasite, 3-host system in small
streams in the Cascade Mountains. Clear-cut logging is replicated across
watersheds in this area and thus provides an ideal setting to assess the
effects of disturbance on complex host-parasite systems. In previous work,
I show strong host-specific effects of logging on parasite prevalence and
intensity. Here, I study the aspect of parasite-host interactions that will be
most sensitive to disturbance: parasite transmission between hosts. Models
predict that parasite dynamics are sensitive to the form of transmission.
Mass-action models predict a single threshold host density and one stable
equilibrium whereas models that assume nonlinear transmission exhibit
more complex behaviors that include multiple stable states, limit cycles,
and in some cases no host threshold density. Quantifying the form of transmission is essential to build realistic models of parasite-host dynamics and
to predict the direction and strength of response of parasites to disturbance.
I conducted a series of field experiments to test how transmission of each
parasite life-history stage responds to disturbance-caused changes in the
density of donor (infected) and target (susceptible) hosts. I show that parasite transmission is not a simple mass action process but it is non-linear
and context dependent in that limits to transmission differentially depend
upon reproduction and recruitment based on relative densities of donor and
target hosts. The change in host densities in response to logging alters the
functional response of transmission and forces the parasite-host system into
an alternate, disturbance-induced state.
POTTS, DANIEL L.,1,* TRAVIS E. HUXMAN,1 BRIAN J. ENQUIST,1
JAKE F. WELTZIN2 and DAVID G. WILLIAMS.3 1 University of Arizona,
Tucson, AZ; 2 University of Tennessee, Knoxville, TN; 3 University of Wyoming, Laramie, WY. Multivariate analysis of ecosystem response to
precipitation pulses.
We currently do not have a framework by which to evaluate how wholeecosystems respond to environmental variation and perturbations either
over short or long time scales. Here we assess and quantify complex system
dynamics by utilizing multivariate techniques. During June 2002, we measured a suite of physiological and ecosystem measurements during 5 diurnal periods for two weeks following a simulated 25mm rainfall pulse for
replicated plots of either native (Heteropogon contortus) or nonnative (Eragrostis lehmanniana) warm-season bunchgrasses on two distinct soil
types. PCA of process variables reveals that independent of species or soil
type, treatments followed a similar, nonlinear trajectory with measurement
periods forming distinct clusters in multivariate space. PCA axis 1 and 2
combined to explain .70% of variation in treatments. We conclude that
bunchgrass species and soil type has influence over the magnitude of ecosystem processes, but does not change the fundamental trajectory of ecosystem metabolism through time in response to a rainfall pulse. This multivariate approach provides a unique lens through which to view and characterize interactions between ecosystem properties and responses to shortterm fluctuations in resource availability.
POULTER, BENJAMIN.* Duke University, Nicholas School of the Environment and Earth Sciences, bp4@duke.edu, Durham, North Carolina.
Lagged responses to sea level rise by estuarine forests on the Albemarle
Peninsula, North Carolina.
Over the next 100 years, sea level rise is forecast to inundate more than
2000 km2 of North Carolinas coastal and estuarine forests. Much of this
land is concentrated around the Albemarle Peninsula where loblolly pine
forests extend down to the shoreline. These forests are vulnerable to projected sea level rise due to a combination of low elevation, exceptionally
high rates of recent sea level rise (due, in part, to regional subsidence),
little to no slope on which forests might migrate upland, and intensive
human land-use on potential inland (refuge) areas. Field observations show

a patchy distribution of forest retreat and marsh transgression inland. Preliminary evidence suggests that the heterogeneity in forest retreat may be
correlated with the interaction of sea level rise with disturbance, such as
fire, hurricanes, and land-use (e.g. timber harvest). We are investigating the
potential for lagged responses of forest communities to sea level rise caused
by feedbacks delaying the invasion of marsh species, and the role of disturbance as disrupting these feedbacks. Using a combination of historical
reconstruction of the pattern of forest communities and field observations
of the status and structure of forest communities we are testing the hypothesis that forest retreat occurs in punctuated events driven primarily by
the interaction of sea level rise with stochastic disturbance events. The
potential for lagged, stochastic responses of estuarine forests to the gradual
effects of sea level rise will have important implications for modeling future forest migration.
POWELL, THOMAS,1,* ROSVEL BRACHO,2 ROSS HINKLE3 and
BERT DRAKE.1 1 Smithsonian Environmental Research Center, Edgewater,
MD; 2 National Research Council, Kennedy Space Center, FL; 3 Dynamac
Corporation, Kennedy Space Center, FL. Comparison of closed-path and
open-path eddy covariance systems over a Florida scrub oak ecosystem.
Eddy covariance using a closed-path infrared gas analyzer (IRGA) has been
a useful tool for making composite measurements of ecosystem gas exchange. Recently an open-path IRGA has become available as an alternative to the closed-path IRGA to circumvent concerns over flux loss associated with sampling tubes and sensor response. From November 2002
through February 2003, measurements comparing the two different eddy
covariance systems were made above a scrub oak ecosystems in central
Florida. System one was comprised of a Gill R3 anemometer and a LICOR closed-path IRGA (LI-6262). System two was comprised of a Campbell Scientific CSAT3 anemometer and a LI-COR open-path IRGA (LI7500). Wind statistics and mean maximum half-hour fluxes for daytime net
ecosystem exchange (NEE) and sensible heat were all within 2% between
the two systems. However, latent energy (LE) for the open-path system was
35% greater (R2 5 0.89) than the closed-path system. The open-path system also achieved a greater level of energy closure, a problem chronic to
eddy covariance measurements, when compared to an independent measurement of net radiation. When the two IRGAs where collocated on the
CSAT3 anemometer, there was less than a 1% difference (R2 5 0.99) in
NEE, but the difference in LE was still considerable at 18% (R2 5 0.90).
These results provide a benchmark for comparing eddy covariance measurements from multiple sites by enabling us to ensure that any differences
in ecosystem fluxes are attributable to ecological differences and not to
instrumentation.
POWER, ALISON G.* and CHARLES E. MITCHELL. Cornell University,
Ithaca, NY, USA. Host diversity and pathogen spillover in plant communities.
The barley yellow dwarf viruses (BYDVs) have strong effects on the interactions between plant species in mixed host plant communities and can
significantly modify plant community attributes through the process of
pathogen spillover from highly susceptible to less susceptible hosts. The
aphid-transmitted BYDVs can infect over 150 species of wild grasses, but
susceptibility to the virus varies significantly among hosts. We used the
PAV strain of BYDV in field experiments in which we manipulated virus
incidence and host community diversity. The wild oat Avena fatua is a
highly susceptible host of BYDV whose performance can be severely affected by infection with the virus. In these experiments, BYDV reduced
the abundance of A. fatua in mixed host communities, but the presence of
A. fatua increased virus prevalence both across the plant community as a
whole and in more resistant hosts. This process of pathogen spillover from
A. fatua decreased the abundances of less susceptible hosts, decreased host
community evenness, and increased dominance. These results suggest that
plant viruses may have significant and complex impacts on plant communities.

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271

PRATER, MARGARET R.,1,* JOHN A. ARNONE2 and EVAN H. DELUCIA.1 1 University of Illinois Urbana-Champaign, Urbana, IL; 2 Desert
Research Institute, Reno, NV. Conversion of sagebrush to non-native
communities following fire: Impacts of ecosystem carbon and water
exchange.
Invasion of non-native annuals across the American West is causing a widespread transition from perennial sagebrush communities to fire-prone annual grasslands. To determine how this invasion may be disrupting ecosystem function, carbon and water fluxes were quantified during the spring
and summer in three paired sagebrush and adjacent post-fire invaded communities in the northern Great Basin using a 1-m3 gas exchange chamber.
Temporal and spatial patterns of carbon uptake and evapotranspiration were
altered in post-fire communities and were correlated with changes in leaf
area and phenology in each community. Intact native sagebrush communities maintained positive carbon balance (mean CO2 uptake ranged between 4.3 and 0.4 mmol m-2 s-1) throughout the growing season, driven by
significantly greater (P,0.05) C uptake by shrubs. Not unexpectedly, intershrub spaces contributed little to overall C gain in the sagebrush community
but decreases in soil evaporation caused declines in overall communitylevel evapotranspiration. Carbon uptake in the post-fire communities was
determined by the dominant species in each of the three communities.
Perennial bunchgrasses had relatively high rates of C uptake (mean maximum CO2 uptake of 3.3 mmol m-2 s-1) but cheatgrass and mustard communities had lower rates and became net sources of carbon (mean maximum CO2 uptake declined to 20.5 mmol m-2 s-1) with the onset of the
summer drought. Seasonal patterns of water use in post-fire communities
differed from sagebrush communities, and disrupted evapotranspiration patterns led to lower surface soil moisture content and increased soil temperatures in the cheatgrass-dominated community. With altered patterns of
carbon and water use, conversion of native sagebrush to post-fire invasive
communities may disrupt surface-atmosphere exchange and degrade the
carbon storage capacity of these systems.
PRATT, MARNEY C.* Duke University, Durham, NC. Why is the bryozoan Membranipora membranacea such an effective invasive species?
Certain species have life histories characteristics that make them particularly effective invaders. The marine bryozoan Membranipora membranacea was first noticed in New Hampshire in 1987. This species became the
dominant epiphyte on laminarian kelps in the Gulf of Maine within 2 years,
and has had huge effects on kelp forest communities. Kelps encrusted
heavily with M. membranacea tend to get damaged and break more easily,
and massive defoliation of kelp beds has been reported as a result. What
makes M. membranacea such an effective invader? This species has a high
reproductive output, can disperse long distances, is good at colonizing new
substrates, and can grow very rapidly. All of these characters can be related
to hydrodynamics in some way, which is why it is important to understand
how water motion affects different aspects of the life history of M. membranacea. For example, the direction and magnitude of currents largely
determine dispersal patterns, local water flow characteristics may help determine settlement choices, and water flow influences food acquisition
which in turn affects growth, survival, and reproductive output. My research has focused on the effect of water velocity on food capture success,
growth, and survival of M. membranacea. I have found that M. membranacea has a higher ingestion rate, grows faster, and survives longer than the
other species I have tested at all of the velocities examined. But the question remains why? I tested some potential reasons why M. membranacea
captures food so effectively including how the location, arrangement, and
size of the feeding zooids affect overall colony feeding success.
PRATT, R BRANDON,1 FRANK W. EWERS,2 MICHAEL C. LAWSON,1
ANNA L. JACOBSEN1 and STEPHEN D. DAVIS.1 1 Pepperdine University, Malibu, California; 2 Michigan State University, East Lansing, Michigan. Freezing limits the distribution of Malosma laurina in the chaparral of southern California.
We examined the freezing response of two co-occurring evergreen chaparral shrubs, Malosma laurina and Rhus ovata. M. laurina leaves die at
temperatures #26oC whereas R. ovata leaves do not. The death may be
caused by direct damage to cells or from desiccation following freezing-

272

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induced cavitation of stem xylem. We tested three hypotheses: 1) freezinginduced cavitation causes the damage seen in M. laurina; 2) M. laurina
exhibits cavitation at higher temperatures than R. ovata; and 3) M. laurina
seedlings are more frost sensitive than R. ovata seedlings. To test our hypotheses we simulated natural freeze-thaw cycles in a freezing chamber on
branches $1 m in length followed by measurements of the temperature at
50% cell death (LT50s) of leaves and the percentage loss of hydraulic
conductivity (PLC) of stems. Additionally, we measured dark-adapted fluorescence (Fv/Fm) of seedlings in the field during freezing events. The LT50
of leaves was 210oC60.23 for R. ovata and 26oC60.17 for M. laurina.
Stems of both species displayed high PLC post-freezing to #26oC
(PLC592%62.6 for R. ovata and 90%64.2 for M. laurina) compared to
unfrozen control stems (PLC558%65.2 for R. ovata and 45%67.6 for M.
laurina). M. laurina seedlings displayed freezing damage at 24oC (Fv/
Fm50.44160.074; n512) compared to unfrozen seedlings at a control site
(Fv/Fm50.82660.004; n512) and all seedlings were killed by a 26oC
freeze. No freezing injury was observed in R. ovata seedlings. We conclude
that M. laurina is co-limited by direct freezing damage to leaves and high
PLC of stem xylem at temperatures #26oC. R. ovata leaves are more
tolerant of freezing than M. laurina and can survive at high PLC in the
field (native PLC in November584%63.9). M. laurina seedlings are more
vulnerable death from freezing explaining their absence from colder microsites where R. ovata often dominates.
PREISSER, EVAN L.* and DONALD R. STRONG. Center for Population
Biology, UC Davis, Davis, CA, USA. Climate affects predator control
of herbivore outbreaks.
Plant vigor promoted by natural enemies of herbivores provides a powerful
demonstration of food web dynamics. While climate affects both herbivores
and plants, we know little about its role in regulating tritrophic interactions.
We demonstrate that variation in rainfall linked to the El Nino/Southern
Oscillation (ENSO) cycle, through its effect on soil moisture, impacts predators (the entomopathogenic nematode Heterorhabditis marelatus) more
strongly than either herbivores (root-feeding larvae of the ghost moth Hepialus californicus) or their host plants (the bush lupine Lupinus arboreus).
ENSO effects thus modulate this powerful trophic cascade. We performed
a two-year field experiment on lupine bushes, with a two-factor crossed
design: predator (nematode added) or no predator, versus water (supplemental water) or no water. Watering simulated the soil moisture conditions
of wet summers that occur irregularly due to heavy winter/spring rains
produced by ENSO climatic events. The predator treatment tested whether
the nematode was capable of inducing a trophic cascade, while the water
treatment tested whether rainfall directly affected the species in the cascade;
the interaction of the two effects tested whether the indirect effects in the
cascade are influenced by ENSO variation. During the first year, the predator decreased caterpillar abundance and indirectly increased lupine growth
& seed set, while watering did not affect either species. During the second
year, a natural outbreak of ghost moths increased herbivore densities 16fold from the previous year. When confronted with this herbivore outbreak,
predatory nematodes were capable of protecting lupines throughout the
year only in the wet ENSO soils. The predator/water treatment decreased
herbivore abundance in plant roots by 42%, and increased lupine growth
by 54% and seed set by 44%. This suggests that the periodic large-scale
lupine die-offs linked to underground herbivory, and the nematode-ghost
moth-lupine cascade, are affected by climatic variation.
PRENGER, JOSEPH1 and WILLIAM F. DEBUSK.1,2 1 Soil and Water Science Department, University of Florida, Gainesville, Florida, USA; 2 Ecology and Environment, Inc., Pensacola, Florida, USA. Changes in soil microbial activity related to military training and forestry activities.
Soil microbial respiration and enzyme activities are indicators of organic
matter decomposition rates and nutrient (C, N, P) cycling and may be
useful in monitoring soil quality and ecosystem integrity. Military training
activity and forest management practices often cause disruption of soil
structure and increased oxidative or erosional losses of soil organic matter.
We have examined microbial activity and nutrient cycling in forested and
formerly forested ecosystems in the Ft. Benning Military Installation in
west-central Georgia. Upland pine and hardwood forests on the installation
are subject to disturbance both from mechanized training activity and forest

thinning, removal, or management. The primary impacts from training are

related to erosion and sedimentation, while tree removal and soil disturbance from harvesting equipment are the main influences from forestry.
Soil samples were obtained from transects in areas of low, moderate and
severe disturbance, and areas of transition from planted pine to oak forest.
Differences in total carbon, water extractable carbon, and soil respiration
were observed among the various levels of impact. Correlations with enzyme activities will be used to understand the effects on microbial metabolism and nutrient cycling.
PRICE, CHARLES A.,* WILL R. TURNER, MICHAEL L. ROSENZWEIG and BRIAN J. ENQUIST. Department of Ecology and Evolutionary Biology, University of Arizona, Tucson, Az. Constancy in community
measures through time in a tropical forest.
A central goal of ecology is the identification of patterns and processes
responsible for generating patterns of diversity across varying scales and
landscapes. There have been two general approaches, which appear to offer
the ability to provide a mechanistic and quantitative framework for ecology.
The first approach has focused on the mathematical properties of speciesarea relationships. Species-area relationships have been examined at scales
ranging from within small biotas to among continents and the biotic and
abiotic factors thought to influence species-area patterns are numerous. A
second approach has focused on elucidating the fundamental role of organism size and abundance. Body size has been observed to be strongly
correlated with a suite of physiological and life history characteristics such
as: population size, abundance, generation time, growth rate, home range
size and metabolic rate. Surprisingly, few studies have explored how size
and area are fundamentally linked; namely, how organismal size, abundance and area combine to influence patterns of diversity across differing
spatial scales or though time. Here we explore how tree density, aggregation and size in the form of basal stem diameter measures influence the
scaling of diversity in a 12 ha, dry tropical forest plot through time. Despite
significant turnover, shifts in species proportional abundance, shifts in centroid position, and marked aggregation; species-area, individual-area, species-individual and body-size abundance relationships remain constant following twenty years of change. These results highlight several invariant
community-level scaling relationships. Further, our results point to links
between allometric (or body mass) scaling rules and species-area scaling
rules. Together, these patterns show that despite local change in species
dominance and composition, ecological communities display highly robust
patterns that are revealed by scaling across size and area.
PRICE, JENNIFER E.,* ROBERT HAMILTON, ELIZABETH RAVIT and
PETER J. MORIN. Invasion success and the impact of microorganisms
on a resident community.
We conducted an experiment monitoring the invasion success of nine protozoan species into established communities of protozoans and bacteria.
We also studied their effect on the resident community. Three invading
species represented each of three trophic levels: bacterivores, omnivores,
and predators. All of the invaders successfully invaded the resident community, although one species, the predator Didinium nasutum, was rapidly
declining by the conclusion of the experiment. Invader success, measured
by total biomass of the invading species, was not related to the trophic
level of the invader. Predators, however, had the greatest effect on the
resident community. By the end of the experiment, only predators had
significantly affected any of the resident species in comparison with the
control community, which contained no invaders. There was also more
variation in the way that predators affected the resident community. Each
predator eliminated at least one resident species, but the species driven to
extinction differed with the identity of the predator. In contrast, final densities of resident species were similar in treatments with bacterivorous and
omnivorous invaders. The biomass of an invading species was not related
to its effect on the resident community. Stentor coeruleus, a predator, had
consistently low biomass throughout the experiment, but had the strongest
effect upon the resident community. Therefore, we conclude that the biomass of a species was a poor predictor of its effect upon other species in
this community.

PRIHODKO, LARA,* A. SCOTT DENNING and MELVILLE NICHOLLS. Colorado State University, Fort Collins, CO. Climate models and
landscape heterogeneity: Can we see the forest without the trees?
Ecological processes and land surface atmosphere interactions occur at spatial scales much finer than is commonly represented in global and regional
atmospheric circulation models. Restrictions on the representation of land
surface processes in these models are typically driven by computational
limitations and by the spatial scale of available input data. Mismatches in
scale are commonly addressed through functional groupings of vegetation
types and averaging of surface properties and soil characteristics. Remotely
sensed data is often relied upon to characterize those surface processes and
conditions which cannot be easily measured on the surface, either spatially
or temporally, at the necessary scales. What are the consequences of disregarding fine scale landscape heterogeneity for regional and global simulations of land surface-atmosphere interactions and boundary layer processes? This study reports on experiments using a spatially explicit coupled
land-surface atmosphere model, the Simple Biosphere Model (SiB2) coupled to the Regional Atmospheric Modeling System (RAMS). The coupled
model was parameterized using AVHRR data, the STATSGO soil database,
a land cover type map and NCEP reanalysis data for a domain in the upper
Midwest United States. Simulations were run with varying land surface
representations and used to explore how point and regional fluxes of carbon
dioxide, latent and sensible heat, and atmospheric properties, depend on
the resolution with which the land surface is represented. Differences in
simulated fluxes and scalar fields were observed between resolutions however domain average fluxes appear to smooth this variability.
PRINGLE, CATHERINE,1,* ELIZABETH BLOOD,2 LUCINDA JOHNSON,3 CORY BERISH,4 JOHN FELDT,5 JOE DEVIVO6 and JAQUELINE
FARLEY.1 1 Institute of Ecology, University of Georgia, Athens, GA, USA;
2
Joseph Jones Ecological Research Center, Newton, GA, USA; 3 Natural
Resources Research Institute, University of Minnesota, Duluth, MN, USA;
4
U.S. Environmental Protection Agency, Atlanta, GA, USA; 5 Southeast
River Forecast Center, Atlanta, GA, USA; 6 National Park Service, Atlanta,
GA, USA. Thirsty landscapes: Drought and the vulnerability of aquatic resources in protected areas in southeastern USA.
Human population growth combined with drought and regional climate
change are threatening the biological integrity of many remaining natural
areas in the world. It is imperative that efforts to protect these natural areas
incorporate a regional hydrologic perspective. Protected areas (both public
and private) are vulnerable to hydrologic alterations well outside of their
boundaries. While this issue has received some attention in arid regions, it
is now emerging as a major concern in wetter regions of the world. As an
example, the southeastern U.S. (50 inches of annual rainfall) harbors globally-significant biodiversity but is experiencing some of the most rapid
population growth in North America. Multi-year regional droughts occur
approximately every 20-25 years and during these periods there is even
greater human demand for surface and groundwater, which can affect protected areas in a variety of ways. Effects range from decreased base flows
in rivers and increased incidence of fires resulting from groundwater withdrawals - to extirpation of aquatic biota and changes in biogeochemical
cycling in rivers fragmented by new dams and reservoirs. Here we use the
recently created Southeastern Ecological Framework (SEF) as a basis to
examine the vulnerability of aquatic resources in both public and private
protected areas in the southeastern U.S. The SEF Project is a GIS-based
analysis that identifies ecologically significant areas and connectivity in
eight southeastern states. We use the SEF as a tool to identify protected
areas where freshwater resources are vulnerable to regional drought based
on overall geography and location with respect to river drainages and underlying aquifers. Our analysis is designed to aid land managers, involved
in the stewardship of protected areas in the southeastern U.S., in making
science-based management decisions which require a regional hydrologic
perspective.
PRYSBY, MICHELLE D.* Great Smoky Mountains Institute at Tremont,
michelle@gsmit.org, Townsend, TN. Citizen scientists contribute to education and research in Great Smoky Mountains National Park.
Projects underway in Great Smoky Mountains National Park demonstrate
that citizens can play an important role in the monitoring and inventory of

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273

natural resources. Great Smoky Mountains Institute at Tremont (GSMIT),

Appalachian Highlands Science Learning Center, and the non-profit Discover Life in America work in conjunction with the Great Smoky Mountains National Park Resource Management and Science and Resource Education Divisions to involve citizens in research in a way that optimizes
both the educational and scientific benefits to all involved. Citizen scientists
from age 13 through adults are collecting data for studies on salamanders,
pollinators, snails, beetles, moths, and more. Many of these projects are
part of the All Taxa Biodiversity Inventory, an effort to document distributional and ecological data for every species in the park. Citizen involvement ranges from data collection alone to independent research by students
that engages them in the full scientific process. Citizen science is a useful
research tool, making long-term monitoring and large-scale inventories
more feasible. Citizen science also is an important tool for education, providing a way to offer visitors a real connection to park resources. Some
students have chosen careers in science because of their research experiences, and teacher participants have incorporated more inquiry into their
curricula.
PSUTY, NORBERT P.* Rutgers, The State University of New Jersey, New
Brunswick, New Jersey. Global Climate Change and Mobility of Coastal Dunes.
Some consequences of global climate change show a forcing of sea-level
rise and consequent shoreline displacement, incorporating a shift of the
water-land boundary through encroachment of the sea and accompanying
sediment loss. Although seldom considered in the general treatment of
shoreline response to these changes, the coastal foredune will undergo concomitant modifications as sediment transfers are generated and sediment
budgets are altered. Increased exposure of the foredune to storm events
will lead to more frequent scarping, reduction of foredune volume, increased opportunity for blowout development, increased opportunity for
mobilization and inland transport of sediment, and increased opportunity
for inland migration of dunal forms. Scenarios of enhanced foredune mobility under conditions of rising sea level and diminishing coastal sediment
budgets suggest a sequential change in the foredune morphology that fosters dissection of the coherent foredune and support of active sand transport
into hinterland dune fields and a potential reactivation of transgressive dune
forms. This stage in the sequence contrasts foredune deterioration with
enhancement of inland dune development. Past episodes of dune field mobilization and stabilization can be linked to global climate changes as well
as human interventions. Human mismanagement can exacerbate the rates
of change and affect the magnitude of inland transgression. Attempts at
mitigation of dunal response to global climate change should strive to restore the natural rates of change in this dynamic system rather than force
stability on a system in disequilibrium.
PYKE, CHRISTOPHER R.* National Center for Ecological Analysis and
Synthesis, Santa Barbara, CA. Biased reserve networks alter climate
change impacts.
Human activities are altering the geographic distribution of habitat while
simultaneously changing climate patterns. This creates dynamic landscapes
where species are squeezed into smaller ranges with increasing limitations
on available habitat. Biological reserves are set aside to promote species
survival; however, reserves are rarely designed explicitly to proportionally
represent the full range of environments inhabited by target organisms.
Often networks of biological reserves protect only a relatively small, potentially biased sample of regional conditions. This problem may result in
counter-intuitive distributions of remaining environmental conditions under
combinations of climate and land-use change. This study explored these
implications for reserve networks in California. The study combined a
broad-scale analysis of climatic conditions available to wildlife across the
state with a detailed analysis of changes in hydrologic regimes for vernal
pool wetlands in the Central Valley of California. Models of regional climate change and land-use change were combined with geographic data
about the distribution of land management to estimate the availability of
climatic conditions to species under a series of scenarios for the years 2040
and 2100. The results indicate substantial bias in the distribution of climatic
conditions in existing biological reserves for most species. Lands managed
for combinations of resource extraction and biodiversity value (so-called

274

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working lands) can make up the difference and, in combination with reserves, provide adequate climatic representation. In the Central Valley, climate change alone could shift the distribution of hydrologic conditions
toward longer, more frequent periods of inundation. However, existing reserves are biased toward the drier portions of the region and if habitat
outside reserves is lost, the surviving wetlands will experience drier conditions even under wetter climate scenarios. These findings indicate that
establishing reserve networks that proportionally represent current climate
can contribute to a predictable and more manageable response to climate
change.
QI, SHI,1 GE SUN,2 STEVE MCNULTY2 and JENNIFER MOORE.2 1 Beijing Forestry University, Beijing, Beijing, P.R.China; 2 SGCP, USDA Forest Service, Raleigh, NC. Modeling the Effects of Land Use and Climate
Change on Water Yield in a Costal Watershed of North Carolina.
Continued population increases will place increased demands on forests to
supply water for commercial and residential use. In addition to increases
population demands, land use and climate change will alter the availability
of water resources. We used a landuse and climatesensitive hydrological
model called PRMS to study the potential change in water yield across a
large coastal plain watershed (Trent River) in North Carolina. The model
was first calibrated and then validated using historical long-term hydrologic
data collected within the watershed. We examined the sensitivity of streamflow to elevated air temperature and variable precipitation patterns. The
results show that the water yield decreases with the increases in air temperature and decreases in precipitation. We then compared predicted water
yield using historic land use and several land use change scenarios. Compared to climate change, landuse change may have more pronounced effects
on water yield. When forest cover changes into agriculture land or urban
use, both total water yield and peakflow rates increase. The results and
implications of this research for regional water planning will be presented.
QUINTANA-ASCENCIO, PEDRO F.,* ERIC S. MENGES, CARL W.
WEEKLEY and OROU G. GAOUE. Archbold Biological Station, Venus,
Florida, USA. Demographic variation with fire in an endemic mint.
We report analyses of the demography of the narrowly endemic, endangered Florida scrub mint Dicerandra frutescens based on data collected
from 1988-2000. Data include survival, growth, and estimated fecundity
of thousands of plants in 7 populations, as well as 10 seed bank/germination experiments. Demographic patterns are driven by fire. Finite rates of
increase (lambda) calculated from 79 stage-structured transition matrices
are highest shortly after fire and decline (fit with an inverse function) sharply through 10 years postfire. The break-even value of l 51 is passed
quickly, in about 6 years postfire, suggesting that older populations are
already facing decline. Population decline is probably related to rapid shrub
growth in the habitat of D. frutescens. In long-unburned sites, finite rates
of increase were negative (l , 1) but they were no lower than rates found
in sites 9-10 years postfire, suggesting some degree of persistence of D.
frutescens in long-unburned sites. Firelanes, road edges, and other disturbed
areas also provide habitats for D. frutescens. In abandoned firelanes and
an irrigated suburban site, finite rates of increase were often . one. However, such sites may not offer stable environments for species persistence.
RACHEL, GARY* and RONALD HENDRICK. University of Georgia,
gcrachel@uga.edu, Athens, Georgia. Assessment of the mycorrhizal
community by soil DNA sequencing and its correlation with soil N and
P fractions.
We assessed the distribution of extra-matrical hyphae within soil horizons
of the three major types of mycorrhizae (arbuscular, ericoid, and ectomycorrhizae). We also determined the distribution of soil N and P fractions
within these same soil horizons to assess whether a correlation exists between these fractions and the hyphal distribution. The study was carried
out within forested stands of the Coweeta Hydrological Laboratory, in Otto,
North Carolina. The stands were chosen for their occupation by plant hosts
of each of the three mycorrhizal types. Mycorrhizal distribution assessments were carried out using DNA based techniques. Total soil DNA was
extracted using five different techniques to determine which protocol provided the highest amount of total DNA and greatest diversity of fungal

DNA. A combination of spectrophotometer, Picogreen, and ARDRA assays

were utilized to determine best protocol. For extraction of mycorrhizal
fungal DNA from the soil, commercial kits utilizing bead beating followed
by GITC/silica gel purification proved the most efficient, providing almost
twice the DNA content and greater banding in the ARDRA assay, indicating a higher diversity of amplified fragments, than the other extraction
methods. The total soil DNA extracts were then PCR amplified using several sets of specific primers. The products were cloned, sequenced, and
analyzed for their phylogenic relationships with sequences of known mycorrhizal species. The distribution of the mycorrhizal fungi were then compared to various soil N and P fractions. The fractionation procedure provides measures of the inorganic, amino and sugar N concentrations, as well
as several organic and inorganic P pools. Preliminary results indicate a
correlation between mycorrhizal fungal distribution and soil N and P fractions. Ectomycorrhizal fungi were detected in all soil horizons, with several
species ubiquitous throughout the soil profile, while other species were
more discreetly localized. Arbuscular mycorrhizal fungi were detected in
predominately A horizons, with some species extracted from O horizon
soil. Ericoid mycorrhizal fungi were the predominant mycorrhizal fungi in
the O horizon, and were extracted from A and B horizon soil as well.
RAICH, JAMES W.* Iowa State University, Ames, IA, U.S.A. Anabolic
and catabolic contributions to total soil respiration.
The results of numerous studies indicate that soil respiration rates are likely
to increase in response to increases in atmospheric CO2 and to increases
in temperature. However, the effects of higher soil respiration rates on soil
organic matter (SOM) pools are difficult to ascertain because soil respiration is a net flux that reflects changes in SOM decomposition and root
respiration rates, both of which may respond to environmental changes. I
review published data and present new data that was collected to distinguish belowground autotrophic (root) and heterotrophic (microbial) contributions to the total soil CO2 efflux. Isotopic data, in particular, have shown
promise of allowing these two main CO2-producing processes to be individually quantified, in sites that have recently been subject to a change in
vegetation cover. What evidence exists, though, for the remaining 99% of
the worlds land surface? I suggest that the classic distinction between
autotrophic and heterotrophic sources of CO2 in soils needs to be modified
to better understand belowground C-cycling processes, and propose a conceptual model that is consistent with existing data, and which allows for
the estimation of anabolic and catabolic contributions to the total soil-CO2
efflux. Such an approach would greatly enhance our capacity to interpret
the results of experimental studies, and to evaluate the potential responses
of terrestrial ecosystems to changing environmental conditions.
RAIMONDI, PETER T.,1,* DAVE LOHSE1 and CAROL BLANCHETTE.2
Department of Ecology and Evolutionary Biology, University of California, Long Marine Lab, Santa Cruz, California; 2 Marine Science Institute,
University of California, Santa Barbara, California. Unexpected dynamism in zonation and abundance revealed by long-term monitoring on
rocky shores.

A disproportionate number of the studies that have provided the foundations for current ecological theory have been manipulative in nature. While
the experimental approach has played an essential role in our investigations
of the processes that affect communities, it is increasingly clear that the
detection of an ecological process is not equivalent to a demonstration of
its importance. This is a much more difficult task, and our lack of success
in this regard has been the basis for potent criticism. Perhaps the biggest
impediment to our development of general rules for community organization is a lack of knowledge about the spatial and temporal variability of
natural communities. This type of information is needed to provide a background against which the results of short-term, manipulative studies can be
discussed. Here we report the results of perhaps the most spatially and
temporally comprehensive study done in the rocky intertidal system. We
have found extremely strong latitudinal patterns of community organization, as well as equally strong site-specific temporal dynamics. These results can be used to assess the relative importance of physical and biological forcing across spatial and temporal scales.

RAINS, KAI C.1,2,* and CAROLINE S. BLEDSOE.2 1 University of South

Florida, kcrains@ucdavis.edu, Tampa, FL; 2 University of California, Davis, CA. Acquisition of nitrogen from decomposing plant residues by
ectomycorrhizal bishop pine (Pinus muricata) and ericoid mycorrhizal
evergreen huckleberry (Vaccinium ovatum).
Ericoid mycorrhizas are generally considered more adept than ectomycorrhizas at acquiring N from organic sources. This may be particularly important in ecosystems where nutrient cycling occurs primarily in surface
organic-rich horizons. We contrasted the ability of an ericoid mycorrhizal
shrub and a co-occurring ectomycorrhizal tree to obtain N from decomposing plant residues. Plants were grown individually and in pairs, including all intraspecific and interspecific combinations, in soil collected from
the O/A horizon of a California mixed conifer forest. Pots were amended
with one of four 15N-enriched plant residues: bishop pine needles, bishop
pine roots, evergreen huckleberry foliage, or evergreen huckleberry roots.
Plants grown individually were harvested after 2, 5, and 12 months, while
those grown in pairs were harvested after 12 months. The quantity of N
obtained from all residue types increased for both species between 2 and
5 months. At both harvests, we found pines and huckleberries had accumulated greater amounts of N from pine root residues than from the other
residue types. After 5 months, pines had obtained more N than had huckleberries from decomposing pine needles and pine roots, while huckleberries had obtained more N than had pines from decomposing huckleberry
foliage. Both species obtained similar amounts of N from decomposing
huckleberry roots. Thus, during the initial stages of decomposition, ectomycorrhizal bishop pines, as opposed to ericoid mycorrhizal huckleberry,
may have greater access to the N found in the abundant pine litter layer in
these forests. The results from the 12 month harvest also will be presented.
RAJANIEMI, TARA K.* Indiana University, Bloomington, IN. Root foraging strategies of herbaceous plants: Trade-offs and size-dependence.
Plants may use various strategies to forage for spatially heterogeneous soil
resources, including high scale foraging (exploring maximum soil volume),
high precision foraging (concentrating root growth in resource-rich patches), and high rate foraging (reaching and/or depleting resource-rich patches
quickly). Differences in foraging strategies might increase or decrease coexistence. Trade-offs among foraging traits may allow competing species
to coexist. In contrast, if larger individuals forage more precisely or at a
higher rate, those individuals might pre-empt resource patches, contributing
to size asymmetric competition belowground. Size asymmetry is expected
to increase competitive exclusion. I used a greenhouse experiment to measure the scale, precision, and rate of root foraging in eight herbaceous
species over a range of individual sizes. No trade-offs among the species
were detected: the species with the greatest scale of foraging also had the
greatest precision and reached patches most quickly. Precision and rate of
foraging were size-dependent for some species: in two species (Bromus
inermis and Festuca rubra) larger individuals foraged more precisely; in
three species (B. inermis, Achillea millefolium, and Centaurea maculosa)
larger individuals reached patches more quickly; and in two species (B.
inermis and C. maculosa) larger individuals had greater root growth rates
within patches.
RANDALL, JESSE A.* and MICHAEL B. WALTERS.* Department of
Forestry, Michigan State University, East Lansing, Michigan. Deer and
sedge effects on vegetation dynamics in a deer altered landscape.
Great Lakes northern hardwood ecosystems with high deer densities have
undergone profound vegetation changes. These include tree recruitment
failure, decreased vertical structure complexity, and shifts in composition
to less palatable species such as sedge (e.g. Carex pensylvanica). Some of
these shifts may persist, even if deer are removed, potentially creating an
alternate stable vegetation state. We are quantifying the impacts of high
deer and sedge densities on vegetation dynamics with a field experiment
that includes vegetation removal treatments, that are possible restoration
methods (sedge removal, herbicide, herbicide and scarification, and controls), in deer exclosure and open areas. Here we present 2nd year results
of a 5 year study. Planted sugar maple seedling survival was significantly
greater in deer exclosures (81.6%) compared to open areas (54.2%). Sedge
removal resulted in 1- increased survival for planted white ash when com-

Abstracts

275

pared to controls in open areas, and, 2- compared to controls, large declines

(.30%) in survival of sugar maple seedlings and biomass of herbs ,25
cm tall, in open areas but not in exclosures. In 2001 Michigan experienced
a prolonged growing season drought. Areas with sedge removal had higher
soil moisture (9.9%), and lower visible damage to sugar maple seedlings
than areas with sedge (8.6%). Treatments thus far show no effects on N
mineralization rates, but standing pools of NH4+ in sedge removal areas
have increased. Preliminary results from a companion greenhouse experiment corroborate our field results as sedge closes its stomates at lower
predawn water potentials and transpires at a greater rate than sugar maple,
and sedge decreases soil NH4+ below that of sugar maple. To date the
restoration of vegetation does not appear to be effective in open areas, but
herb biomass has returned to control levels within exclosures.
RANDRIAMAHEFA, ALEXANDRINE, OETINO ONESIMUS and NATHAN MCGINNIS. Oakwood College, Huntsville, AL 35896, USA,
Huntsville, AL, USA. Practical experience leads minority to ecology.
Oakwood College has been the recipient of the Strategies for Ecology Education Development and Sustainability (SEEDS) grant for the past four
years. SEEDS is a joint program of the ESA and the United Negro College
Fund (UNCF) which addresses the lack of minority representation in the
ecological field. The thrust of the SEEDS program at Oakwood College is
two-fold. The first step is to enhance the existing curricula by developing
several outdoor laboratories. The second step is to raise ecological awareness on campus. Firstly, curriculum enhancement activities include the development of a nature trail (1.5 mile) and a campus pond which provide
several well-defined ecotones to explore. In addition, a dendrochronology
exhibit is completed and offers students the opportunity to examine the
climatic history as illustrated by a cross section of a 60 year- old tree.
These hands-on labs help to enlighten the students by providing more active learning experiences that will further assist them while they study
ecology, plant biology, and environmental courses. Secondly, the SEEDS
students along with the Ecology Club promote awareness of environmental
issues and career opportunities through different activities including, but
not limited to Earth Day, America Recycles Day, and field trips. The
SEEDS program at OC is gaining momentum and showing evidence of
success in motivating minority students to pursue Ecology related fields.
The average enrollment in General Ecology has more than doubled (7 to
15). Additionally, two students are finishing their graduate studies, while
two others are applying to pursue post-graduate degrees in the ecological
sciences. There has also been a rapid increase in non-biology major participation in the ecological awareness projects.
RASCHER, UWE. Biosphere 2 Center, Columbia University, P.O. Box 689,
Oracle, AZ, USA. Hyperspectral imaging serves as a powerful tool to
monitor the spatio-temporal dynamics of photosynthesis within a
drought-stressed tropical canopy.
Thus far, the spatio-temporal dynamics of photosynthetic light use efficiency of complex canopies cannot be quantified non-destructively by passive, remote sensing techniques. However, hyperspectral imaging techniques and indices derived thereof have the potential for large scale ecosystem monitoring and can be closely linked to photosynthetic electron
transport. The tropical rainforest within the Biosphere 2 Laboratory provides an ideal testbed for hyper-spectral instrumentation and can deliver
data from complex canopy structures. A new field portable hyperspectral
imaging device proofs a detectable change in the reflectance signature from
a complex tropical canopy, which was subjected to a mild, prolonged
drought. Ecosystem carbon balance responded to drought. The effects occurred within days and the changes were reversible within weeks. Reduced
gross photosynthetic production (GPP) was off-set by reduced soil respiration, leaving the 24 hour net ecosystem exchange almost unaffected. The
reduction of GPP reflects two underlying mechanisms: (i) an increased leaffall during the first days of the drought, which resulted in decreased foliage
coverage of the canopy and (ii) reduced maximum photosynthetic electron
transport rate, which was recorded on 50 different leaves using chlorophyll
fluorescence techniques. Both effects, were also detected within the hyperspectral reflectance images of the canopy. Moreover, the canopy was not
affected uniformly. Patches of the canopy responded more strongly than
others, which can be deducted from the hyperspectral images and also from

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simultaneously performed leaf level measurements. Indices, such as the

photosynthetic reflectance index (PRI), were used to quantify those spatiotemporal effects.
RASHLEIGH, BRENDA1,* and DAVID RANDALL.2 1 U.S. Environmental Protection Agency, Athens, Georgia; 2 City University of Hong Kong,
Kowloon, Hong Kong. Modeling the response of fish populations to
eutrophication.
Eutrophication resulting from nonpoint source pollution is one of the largest environmental problems in lakes and reservoirs around the world. Two
characteristics of eutrophication, decreased dissolved oxygen and increased
concentration of ammonia, are known to affect fishes, yet models of eutrophication rarely include fishes. We developed a model to quantify population-level effects of eutrophication on fish. The model uses oxygen, ammonia, pH, and temperature outputs from a water quality model as driving
variables and incorporates results from individual-level fish toxicology
studies of growth, survival, and reproduction conducted in the laboratory.
The model is applied to Lake Peipsi in Estonia/Russia. We focus on two
common, commercially-important fish species, smelt (prey) and pikeperch
(predator). Results indicate that the two fish species respond similarly to
eutrophication, the fish populations are most sensitive to changes in oxygen, and over time, ammonia and oxygen changes are synergistic in their
effects on populations. This model can be used to assess the response of
the fisheries to future water quality scenarios and restoration activities.
RAY, CHRIS.1,2,* 1 University of Colorado, cray@colorado.edu, Boulder,
CO; 2 University of Nevada, Reno, NV. Modeling plague in prairie communities: using sensitivity analysis to guide data collection and model
construction.
Sylvatic plague, the source of human bubonic plague, has invaded wildlife
communities across western North America. The plague bacterium, Yersinia pestis, is typically transmitted between mammals by fleas. The effective
virulence of Y. pestis varies among host species, and the efficiency of Y.
pestis transmission varies among vector species. No single host-vector association has been identified as the predominant reservoir for plague in
North America. Instead, the plague reservoir appears to involve several
host and vector species and frequent disruption of normal host-vector associations. In prairie communities, the most obvious impacts of plague
include infrequent, isolated human infections and frequent, extensive prairie dog die-offs. These die-offs may affect the ecological and epidemiological function of prairie communities. Many species may benefit from
the ecological services provided by prairie dog colonies, so the density of
many plague hosts and the contact rates between hosts and vectors may be
altered by prairie dog presence. Given these unknown and potentially complex relationships between hosts and vectors, no single model of plague
dynamics can yet be proposed for prairie communities. But I have used a
type of sensitivity analysisfamiliar from population viability analysis
to analyze the behavior of many potential plague models. This analysis
begins with a matrix model to project dynamics of infectious and noninfectious segments of several host and vector populations. Each matrix
element represents a transition rate governed by a (linear or nonlinear)
relationship between segments of the community (e.g., the rate at which
mouse fleas become infectious through contact with infectious prairie
dogs). Model predictions are sensitive to changes in each matrix element,
and this sensitivity can be used to suggest appropriate model structures and
data collection efforts. Appropriate models should predict observed patterns, and research priorities should focus on the more sensitive elements
of these models.
REAGEL, PETER F., MATTHEW D. GINZEL and LAWRENCE M.
HANKS. University of Illinois at Urbana-Champaign, Urbana, Illinois. Aggregation and mate location in the red milkweed beetle (Coleoptera:
Cerambycidea).
The red milkweed beetle, Tetraopes tetrophthalmus Forster, is a univoltine
cerambycid that feeds upon the leaves and flowers of common milkweed,
Asclepias syriaca L. Adult milkweed beetles prefer to aggregate on milkweeds that have multiple inflorescences of large size. Males actively
searched for females, often flying between host plants, and apparently do

not locate females from a distance using long-range pheromones or vision,

but rather land on milkweed stems arbitrarily, whether females are present
or not. Males remain for longer periods, and so tend to accumulate, on
milkweed stems that had female-biased sex ratios. We conclude that aggregation of T. tetrophthalmus is cued by host plant characteristics, but
dynamically influenced by the sex ratio of conspecifics present on individual stems.
REAL, LESLIE A.,1,* JENNIFER M. SNAMAN,1 TRACY LAMBERTJACK,2 LANCE WALLER2 and JAMES CHILDS.3 1 Department of Biology and the Center for Disease Ecology, Emory University, Atlanta, GA;
2
School of Public Health, Emory University, Atlanta, GA; 3 Centers for
Disease Control and Prevention, Atlanta, GA. Ecological phylogeography
of fox rabies in Ontario.
Often spatial analysis of genetic structure in phylogeographic studies are
not coupled to ecological variables and underlying spatial population dynamics. At the same time, epidemic models of disease spread often lack
detailed genetic information. Here we present an analysis of the spatial
organization of rabies virus genotypes associated with the epidemic expansion of fox rabies from the arctic region into western Ontario. We use the
spatial organization of 82 rabies isolates from red fox that have been previously sequenced by S.A. Nadin-Davis (1999) at the N and G genes of
this 12 KB single-stranded negative-sense RNA virus. The 82 isolates cluster into 21 distinguishable molecular variants. We applied modern geostatistical methods to determine the spatial location of distinct sequence clusters (by type), the geographic boundaries associated with these clusters
arrived at by categorical wombling, and correlations among disease sequence clusters and GIS derived environmental variables (e.g. extent of
rivers, human population corridors, vegetation type and cover, and presence
or absence of bays and/or lakes). We tested the hypothesis that spatial
pattern in sequence variation corresponds to and was generated by the
spatial pattern of epidemic expansion of fox rabies in Ontario. Genetic
distance and spatial distance are significantly correlated and spatial structure does correspond to underlying ecological variation. The rabies virus
appears to genetically differentiate and form distinct spatial paths of expansion where the direction of spread was influenced by the Ottawa River
to the north, the Georgian Bay and Lake Huron to the west, and Lake
Ontario to the south.
REBBECK, JOANNE,* ROBERT P. LONG and DANIEL A. YAUSSY.
USDA Forest Service, Northeastern Research Station, Delaware, OH, USA.
Topkill and mortality of maple saplings following a prescribed fire in
the Ohio Hills.
Since adoption of fire-suppression programs in the 1930s, the composition
of mixed oak forests throughout the eastern US has been shifting from oak
(Quercus spp.) and hickories (Carya spp.) to more-shade tolerant and fireintolerant species such as red (Acer rubrum L.) and sugar (A. saccharum
Marsh.) maples. The use of prescribed fire may improve oak regeneration.
Previous research with prescribed fires conducted in southeastern Ohio
showed that fire temperatures of 1508C or more caused 75% topkill of 36 cm d.b.h. (diameter at breast height) saplings and 55-65% of larger saplings (6.1-10 cm d.b.h.). Our objective was to estimate threshold bark temperatures during prescribed burns that elicit reduced vigor and mortality of
saplings and small trees. In Spring 2001, aluminum tags coated with a
series of temperature sensitive paints were secured on the lower stems of
red and sugar maple saplings in thinned or unthinned 20-hectare study areas
at three sites to estimate maximum bark temperature during a prescribed
fire (N5240 saplings). Fourteen months after a prescribed fire, 37.0% of
medium (3-6 cm d.b.h.), and 30.0% of large (6.1-10 cm d.b.h) saplings and
17% of small trees (10.1-14.2 cm d.b.h.), were topkilled but resprouted,
while 9.3, 8.6, and 5.7% of medium and large saplings, and small trees,
respectively, died and had not resprouted. Threshold bark temperatures for
topkilling ranged from 1008C for medium and large saplings to 1848C for
small trees. Mortality (no resprouting) was infrequent and commonly observed at bark temperatures of 400-4838C. Since achieving high intensity
fires may not be practical, a series of frequent, numerous lower intensity
spring burns (.1008C) which suppress repeated sprouting may be necessary to kill oak competitors such as maple.

REBEL, KARIN T.,1,* SUSAN J. RIHA,1 TIMOTHY J. FAHEY1 and

JOHN SEAMAN.2 1 Cornell University, Ithaca, NY; 2 Savannah River Ecology Lab, Aiken, SC. Simulated and measured tree water uptake using
tritium as a tracer.

Methods for determining in the field if tree species differ in where they
take up water in the soil profile are limited, as are methods to determine
root distribution for modeling purposes. In this study, tritium transpired by
trees periodically irrigated with tritium enriched water at a southern US
coastal plain site was measured and simulated. Tritium in transpired water
was measured by liquid scintillation analysis following collection in bags.
Water and tritium uptake were simulated using a multi-layer one-dimensional water balance model. Soil water movement and storage were simulated using a capacitance approach, soil water and tritium uptake were
simulated using an absorption approach. The potential demand for water
and tritium uptake was partitioned among soil layers based on relative root
density distribution, estimated using inverse modeling techniques. When
soil water content decreased in surface layers, the demand was transferred
between layers. Modeled and measured tritium activity in the transpiration
water were compared, as well as the measured differences between tree
species. The measured tritium activity in the transpiration water followed
the simulated tritium activity well. When tritium activity in the upper part
of the soil profile was high relative to the lower part of the profile, a large
range in the values of measured tritium activity in the transpiration water
was observed. This measured increase in range of the tritium activity cooccurred with an increase in simulated transpired tritium activity. Differences in measured tritium activity in the transpiration water was most apparent between understory and overstory trees, comparing the same species
as well as among species. These results imply that roots of individual trees
and tree species were taking up water in different parts of the soil profile.
On a stand basis the root distribution estimated using inverse modeling
techniques appeared to well represent water and tritium uptake.

REED, DAVID L.* Florida International University, Miami, Florida. Seed

germination and growth of four wetland tree species in response to
environmental factors in tree islands of northern Shark Slough, Everglades National Park.

Seedling dynamics are the initial filter in the establishment of tree species
in wetland forests. Patterns of seedling establishment of dominant species
can be attributed to key environmental variables including light and hydrology. The vegetative communities within Everglades tree islands known
as Bayheads and Bayhead Swamps are continuously inundated for parts of
the year and are delineated by heterogeneous canopies and microtopographic features, creating correlate variable light and hydrologic environments.
Hydrology is further affected by seasonal oscillations in water level in
Everglades National Park (ENP), allowing for temporary drainage of soils
during spring. This may cause seedling emergence to be a function of the
timing of seed germination. Likewise, microtopographic highs along the
elevation gradients within these communities provide temporary refugia
from rising water levels and prolonged drainage. Life history and physiology of four dominant wetland tree species: pond apple (Annona glabra),
cocoplum (Chrysobalanus icaco), sweetbay (Magnolia virginiana), and
wax myrtle (Myrica cerifera) with regard to variable shade and flood conditions were explored. Field experiments quantified elevation, hydrology,
and light environments, while monitoring recent germinant density and
growth in three tree islands of northern Shark Slough, ENP. The physiological and morphological responses of early juveniles to variable light and
flood conditions were assayed in shadehouse experiments, including three
light and three flood treatments. Preliminary data show seasonal trends in
seedling density and species diversity. A. glabra dominates the wetter sites
all year, and both sites during winter months. C. icaco appears ubiquitously,
but at lower densities, while M. virginiana and M. cerifera often appear at
higher elevations and in the spring and summer months when overall densities increase. Synthesis of these and site hydrologic data allows for prediction of early juvenile species composition and density, and may further
predict community response to long-term changes in hydropattern.

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277

REGAN, HELEN M.,1,* DAVID A. KEITH2 and MARK TOZER.2 1 San

Diego State University, hregan@sciences.sdsu.edu, San Diego, CA, USA;
2
New South Wales National Parks and Wildlife Service, Hurstville, NSW,
Australia. A population model of a perennial shrub threatened by disease and adverse fire regime.
Xanthorrhoea resinifera is a long-lived plant occurring on the south-east
coast of Australia. Like many perennial Australian shrubs, X. resinifera has
a complex relationship with fire. Fire reduces the chance of survival of
plants whilst triggering flowering and seed production. Populations of X.
resinifera are subject to a range of threatening processes. While Xanthorrhoea may survive bushfires, such survival is highly structured in relation
to plant size and fire intensity and frequency. Furthermore, Xanthorrhoea
species are susceptible to the root pathogen Phytophthora cinnamomi which
reduces survival of established plants. The long-term effects of these threats
is poorly understood. We constructed a stochastic, individual-based model
using 10 years of demographic data to investigate the effects of an altered
fire regime and disease on the long-term persistence of sub-populations of
X. resinifera. We included density-dependent processes with con-specific
neighbors as well as shading by neighboring Banksia plants. We demonstrate that the chance of long-term persistence depends on the timing of
fire and the extent of disease. We show that, even though fire does not
eradicate the pathogen, the increased seed production and recruitment following a fire may somewhat alleviate the increase in mortality due to disease. In this way, fire could be an important management tool for this
species.
REHEIS, MARITH C.,1,* ROBERT L. SANFORD,2 JAYNE BELNAP,1
RICHARD REYNOLDS1 and JASON NEFF.1 1 U.S. Geological Survey,
Earth Surface Processes Team, Denver, CO; 2 University of Denver, Denver,
CO. Silver linings: Fertility gains and losses through dust redistribution at local, regional, and global scales.
Wind erosion is commonly invoked as a chief cause of depletion of soil
nutrients in arid and semiarid lands, especially those disturbed by land-use
practices. Nevertheless, modern dust (both wetfall and dryfall deposition)
is a source of macro- and micro-nutrients for soils and vegetation globally,
and may be particularly important in organic-poor arid ecosystems. Dust
is ubiquitous; modeled deposition rates are at least 10-100 kg/ha/yr
throughout the continents and 100-1000 kg/ha/yr or more in the dry midlatitudes. Example estimates of nutrient inputs in semiarid regions are
P50.03-6.0, N50.5-9.0, and C5110-180 kg/ha/yr. In the southwestern
U.S., deposition rates of micro-nutrients are Cu55-90, Mn518-160, and
Mo50.1-0.7 g/ha/yr. Aeolian dust is incorporated into surface sediments
and redistributed downward into soil profiles by water infiltration. The dust
component of soils can be distinguished from parent sediment by a particle
size dominated by fine silt, different magnetite content, and different chemical composition. We focus on study sites having different grazing histories
in the Canyonlands area of Utah and from desert areas of southern Nevada
and California to (1) examine the composition and nutrient content of modern dust, (2) identify the dust component of soils, and (3) infer the contribution of dust to plant-nutrient availability, especially with respect to phosphorus. In general, modern dust samples have much more labile and biological P than soil samples. P contents, however, vary with season: in dust,
total P and labile P are higher in winter than in summer, whereas labile P
in soils increases dramatically in early summer. In A horizons and pothole
sediments of our study sites, wind-deposited concentrations of such plantcritical trace elements as Cu, Mn, Mo, and Zn may be 25-100% greater
than in some parent sediments or rocks. Conversely, wind erosion may
have reduced concentrations of some elements in grazed soils.
REICE, SETH R.1 and CHRISTY JO GERACI.2 1 University of North Carolina, Chapel Hill, NC; 2 Clemson University, Clemson, SC. Impact of
development on stream community structure and decomposition rates:
Streams in piedmont North Carolina.
The conversion of forested landscapes to urban and suburban land uses is
the major threat to stream community health and dynamics. We analyzed
the effect of urban/suburban development and loss of vegetated cover
through comparisons between two dramatically different streams. The
catchment of Bolin Creek has a moderate gradient of development, ranging

278

Abstracts

from 74.5% forests and fields (combined) upstream to 54.9% downstream

(ArcView GIS). Conversely, urban and suburban land uses increase downstream. Meeting of the Waters Creek (MOW) originates on the UNC-CH
campus and is nearly 98% urbanized upstream, and gets progressively more
vegetated (to 75%) as it flows off campus into the N.C. Botanical Garden.
We sampled both streams for macrobenthos and leaf litter decomposition
rates. The benthic communities changed in both streams, becoming less
species rich as the vegetated proportion of the catchment declined. Leaf
litter decomposition rates were not significantly different over the more
narrow range of development in Bolin Creek. In MOW leaf litter decomposition rates increased significantly with increasing vegetated cover. So,
urbanization degraded both biodiversity and decomposition processes. Species richness is a valid and reasonably sensitive instrument for ecological
assessment. Leaf litter decomposition rates give a good estimate of the
function of the stream ecosystem and should be included in the assessment
of stream ecosystem health.
REILEY, DAVID K.,1,* DAVE D. BRESHEARS,2 PAUL H. ZEDLER,1
MIKE H. EBINGER2 and CLIF W. MEYER.2 1 University of WisconsinMadison, Madison, WI, USA; 2 Los Alamos National Laboratory, Los Alamos, NM, USA. Soil carbon patterns in a semiarid pinyon-juniper
woodland.
Pinyon-juniper-grassland complexes cover 60 million acres of the American west. An increase in tree and shrub components has been noted worldwide in the last century along the grassland-shrubland-woodland continuum. This woody encroachment may contribute to the missing sink of carbon noted in various evaluations of global carbon cycles. In Northern New
Mexico, pinyon-juniper (PJ) woodland encroachment into grasslands creates patchy landscapes with sparse herbaceous cover between tree clusters.
To evaluate soil carbon storage we measured total carbon % in a PJ stand
in Los Alamos, New Mexico. A total of 180 cores and 1071 samples of 5
or 10 cm depth increments were obtained and analyzed. We found that soil
carbon was greater on a percentage basis under tree canopies than grass
patches or bare soil in intercanopy areas and, as we generally expected,
was greater in the upper few centimeters of soil than at depth. We hypothesize the observed pattern is due to litter inputs into the upper few centimeters of soil where trees provide greater inputs, grasses somewhat less,
and bare soils less still. However, surface soils often have bulk densities
that are low compared to samples taken at depth. Comparisons using %
carbon may be overstating the actual differences in the carbon stored within
soils. We used a local bulk density evaluation to estimate carbon storage
on a mass/volume basis. Despite the lighter soils in the highest carbon
containing fractions (canopy and grass soils at the surface), our analysis of
soil carbon on a mass/volume basis lead to substantially the same conclusions. Our results suggest that management of PJ for carbon sequestration
will require careful consideration of patch dynamics and the interplay with
potential disturbances that may be common at different stages of woodland
development such as erosion or crown fire.
REINHARDT, CARRIE H.* and SUSAN M. GALATOWITSCH. University of Minnesota, St. Paul, MN, USA. Patterns in rhizome carbohydrate
fluctuation in the invasive Phalaris arundinacea: implications for wetland restorations.
The presence of invasive species in wetland restoration sites often preempts
native vegetation reestablishment. Phalaris arundinacea is an invasive perennial grass especially problematic for wetland restorations because it often dominates drained sites and persists after reflooding. Although Glyphosate application is a popular control method for Phalaris, success with
this method is routinely limited, possibly due to a failure to time application
with the phenology of Phalaris. Because Glyphosate is translocated to rhizomes with carbohydrates, characterizing patterns in seasonal rhizome carbohydrate fluctuations should identify the period of maximum herbicide
translocation to rhizomes. A three-year field study was conducted to determine changes in rhizome carbohydrate levels of Phalaris using nearinfrared spectrometry. Rhizome carbohydrate levels generally followed a
trend of depletion during early season vegetative growth, and accumulation
during late season reserve acquisition. The maximum rate of carbohydrate
translocation to rhizomes varied from year to year, suggesting that phenologic cues, rather than calendar date, must be used to predict carbohydrate

movement. Results from this research suggest that predicting carbohydrate

fluctuation is feasible and timing herbicide application with respect to phenology will do much to increase herbicide efficacy for Phalaris in wetland
restorations, thereby increasing the potential for ecosystem recovery in
these systems.
REINHART, KURT O.,1,* ALISSA PACKER,2 KEITH CLAY,2 WIM H.
VAN DER PUTTEN3 and RAGAN M. CALLAWAY.1 1 University of Montana, reinhart@selway.umt.edu, Missoula, MT; 2 Indiana University, Bloomington, IN; 3 Netherlands Institute of Ecology, Heteren, The Netherlands.
Introduction to plant-microbe feedback in plant invasions and invasive
non-native trees escaping from soil pathogens.
Soil biota are known to affect natural plant communities and may also
affect the impact of invasive non-native species. We tested the sensitivity
of invasive trees (Acer negundo, Acer platanoides, and Prunus serotina)
to the resident soil biota in their native and non-native ranges and related
this to the spatial aggregation of conspecifics in individual ranges. Overall,
there was a general negative effect of the soil biota on seedling survival/
growth of all three species within their native ranges; however, A. negundo,
near the fringe of its natural distribution, was not affected by soil biota. In
the non-native range, the soil biota had a positive effect on the growth of
P. serotina seedlings. Moreover, A. platanoides seedlings within their nonnative range were positively effected by the soil biota from heterospecifics
but negatively effected by the soil biota from conspecifics. Acer negundo,
within its non-native range, was not affected by the soil biota associated
with either conspecifics or heterospecifics. The negative effects of the soil
biota within the native ranges correspond with sparse aggregations of conspecifics. In contrast, the soil biota within the non-native ranges tended to
facilitate the growth of seedlings and corresponded with dense aggregations
of conspecifics. Our results suggest that negative soil feedbacks in native
ranges may help to maintain forest diversity while the positive soil feedbacks in the non-native ranges may help to facilitate invasion by releasing
invasive trees from belowground enemies.
REITER, MICHAEL A.1,* and GEORGE R. PARSONS.2 1 Delaware State
University, Dover, DE; 2 University of Delaware, Newark, DE. An interdisciplinary "four-tier" modeling methodology for resource management: An example from the St. Jones River watershed, Delaware.
An integrated, interdisciplinary conceptual model was developed for the
St. Jones River watershed, Delaware for use as a management and research
tool. The model utilizes the "two-tier" approach of the Environmental Cooperative Science Center methodology (drivers-stressors, stressors-valued
ecosystem components) based upon the VEC modeling structure of Harwell
and Gentile, and expands upon it to include a third (valued ecosystem
components-services) and fourth (services-drivers) tier. The four-tier model
has the potential to more clearly link human activities not only to significant ecological impacts within a habitat, but to possible social and economic impacts on human services including feedbacks upon the initiating
drivers. The four-tier model retains the "ranked matrix" assembly methodology utilized for the two-tier model, but introduces steps in later tiers
to limit the potential number of links to a manageable level. The present
four-tier model for the St. Jones River watershed in central Delaware suggests that climate, development, and agriculture are likely to be the most
important drivers in the watershed, that aquatic habitats are likely to receive
significant amounts of stress, and that development, habitat restoration/
mitigation, and land-use regulation are drivers with a high potential to be
affected by feedbacks from activities in the watershed.
NKOVA
, ELISKA,1,* JOHN GRIECO,2 NICOLE ACHEE,2 SAREJMA
RAH JOHNSON,1 PETR MACEK,1 KEVIN POPE3 and PENNY MASUOKA.2 1 University of California Davis, Davis, CA; 2 USUHS, Bethesda,
MD; 3 Geo Eco Arc Research, Aquasco, MD. Freshwater Community
Interactions and Malaria Persistence in Tropical Wetlands.
Presence and abundance of mosquito larvae in aquatic habitats reflect the
oviposition preference of females as well as the survival capability of larvae. These are regulated by a variety of ecosystem processes operating and
interacting at several organizational levels and spatial/temporal scales. Presence of water, food source, and protection are the key variables. Aquatic

plants (both micro- and macrophytes) provide protection from predators

and, together with other resident vegetation, contribute detritus that supports the bacterial community, which, in turn, serves as food for larvae. A
change in any component in this complex structure may have a substantial
impact on the mosquito population and can even lead to a replacement of
one species with another. Of the three species of anopheline mosquitoes
responsible for malaria transmission in the country of Belize, Central
America, two have their larval stages occurring in freshwater marshes.
Sparse Eleocharis/cyanobacterial mat-dominated marshes provide favorable habitat for Anopheles albimanus while tall dense macropytes (TDM)
such as cattails are typical habitat for Anopheles vestitipennis. We are testing the following hypotheses: (1): Phosphorus (P) enriched runoff from
agricultural lands, namely pasture and sugar cane, and human settlements
causes an expansion of TDM vegetation in wetlands of northern Belize;
(2): TDM marshes provide more productive habitat for Anopheles vestitipennis than for An. albimanus. Since An. vestitipennis is a more efficient
malaria vector than An. albimanus, such development would result in higher malaria risk in human settlements in proximity to the impacted marshes.
RELYEA, RICK A.* Department of Biological Sciences, University of
Pittsburgh, Pittsburgh, PA, USA. Controlling pests and killing amphibians.
Pesticides are used throughout the world to control undesirable pests and
thereby improve the health and well being of humans. For most pesticides,
we know very little about the effect on non-target species, especially on
amphibians. In this study, I assemble aquatic communities (25 species) in
a mesocosm experiment to determine how two insecticides (carbaryl and
malathion) and two herbicides (2,4D and Roundup) affect the species richness and productivity of the communities. I found that carbaryl and malathion had indirect positive effect on amphibians by killing off many of
the amphibian predators. In contrast, the herbicide 2,4D had minimal impacts while Roundup rapidly killed 95-100% of the amphibians. The results
make it clear that pesticides can have both direct and indirect effects and
that certain pesticides have the potential to decimate amphibian communities and lead to amphibian declines.
RENNE, IAN J.,1,* BIANCA G. RIOS,2 JEFFREY S. FEHMI2,3 and BENJAMIN F. TRACY.1 1 Department of Crop Sciences, University of Illinois,
Urbana, IL; 2 Dept. of Natural Resources and Env. Sciences, University of
Illinois, Urbana, IL; 3 U.S. Army ERDC-CERL, Champaign, IL. Allelopathic potential of an invasive forage grass on native grassland plants.
Tall fescue (Festuca arundinacea Schreb.), a highly competitive European
grass that invades US grasslands, is reportedly allelopathic to many agronomic plants. However, its ability to inhibit the germination or growth of
native prairie plants is unknown. In three factorial glasshouse experiments,
we tested the potential allelopathic effects of endophyte-infected (E+) and
uninfected (E-) tall fescue on native grasses and forbs from midwestern
tallgrass prairies. Relative to a water control, at least one extract made from
ground seed, or ground shoot-root tissue of E+ or E- tall fescue reduced
the germination of 10 of 11 species in petri dishes. In addition, the emergence of two native grasses in potting soil was lower when sown with E+
and E- tall fescue seedlings than when sown with seeds of conspecifics or
tall fescue. However, when seeds of 13 prairie species were sown in sterilized, field-collected soil and given water or one of the four tall fescue
extracts daily, seedling emergence was lower in one extract relative to
water for only one species, and subsequent height growth did not differ
among treatments for any species. We conclude that if tall fescue is allelopathic, its inhibitory effects on the germination and seedling growth of
native prairie plants are limited, irrespective of endophyte infection. On the
other hand, the apparent inability of these plants to detect tall fescue in
field soil could hinder prairie restoration efforts if germination near this
strong competitor confers negative fitness consequences. We propose that
lack of chemical recognition may be common among resident and recently
introduced nonindigenous plants because of temporally limited ecological
interactions, and offer a view that challenges the existing allelopathy paradigm. Lastly, we suggest that tall fescue removal will have immediate
benefits to the establishment of native grassland plants.

Abstracts

279

REYNOLDS, BARBARA C.1 and MARK D. HUNTER.2 1 University of

North Carolina at Asheville, Asheville, NC; 2 University of Georgia, Athens, GA. Nitrogen and carbon inputs from canopy herbivores:Comparing outbreak and endemic levels of herbivory.
Frass inputs from canopy herbivores were collected and measured in 20
plots on each of three sites along an elevation gradient at Coweeta Hydrologic Laboratory (USFS) in western North Carolina, USA. 24 hr collections
were made 2-3 times/month throughout the growing season for three years.
Frass from individual plots/date was analyzed for % total nitrogen and %
total carbon, and the C/N ratio was determined for each sample. Average
values for these three categories, plus the weight of frass, were calculated
for each date and extrapolated to gha-124h-1. Frass deposition generally
showed two pronounced peaks on the mid and high elevation sites, coinciding with leaf flush early in the growing season, and a second peak late
in the growing season. At the low elevation site, the most prominent peak
was in the mid to later part of the growing season, and varied with the
year. Frass inputs during a short, chronic, outbreak of sawflies (Periclista
sp.) on the high elevation site were almost twice those measured at other
times or sites. Nitrogen deposition followed the same pattern as frass deposition. Frass C/N generally increased over time for all elevations, except
the low elevation in 1996, but showed considerable variation among collections. Nitrogen content in frass from the high elevation outbreak led to
significantly higher amounts of stream NO 3 compared to previous, nonoutbreak, years or to the control stream.
RHEIS, HAROLD F.1,2,3,* 1 Harold F. Reheis, Environmental Protection Division, harold_reheis@mail.dnr.state.ga.us, Atlanta, GA, USA; 2 ; 3 . The
value and role of sound science in the formulation of Georgians holistic
approach to comprehensive water managment.
Water quantity resources regulation varies greatly from state to state, especially in the Southeast, since there is no federal mandate in this arena.
The State of Georgia began water quantity regulation in 1972. Sound science has played a major role in this vital aspect of water resources management: recognition of problems to be solved; communicating the problems to water users and legislators; building an alliance to support solving
the problems; getting funding to improve the science; and legislation to
authorize needed regulatory changes; and finally the development and implementation of a water management plan to solve the problem, resulting
in sustainable water use. The case of Georgias Flint River Basin, with its
significant municipal, industrial, and agricultural water use and a unique
ecosystem to protect, will be described.
RHOADS, ANNE G. Brown University, Providence, RI, USA. The effect
of abandoned pastures on the composition and structure of the northern hardwood forest of New Hampshire.
In 1870, at the peak of agriculture in the White Mountains of New Hampshire, more than 25% of the land was cleared for pasturing in the western
portion of the region. As farms were abandoned from the late 18009s to
early 19009s, forests returned. Whereas natural disturbances exert a pulse
of disturbance followed by a recovery period, agriculture in New Hampshire brought a sudden, dramatic change, lasted 70-100+ years, and occurred at a landscape level. This landuse is, therefore, hypothesized to have
had significant impacts on the composition and structure of the community
affected. In this study, fifteen abandoned pastures were found in the western
portion of the White Mountain region. Six, 20X20 m plots were established. Within each plot, the dbh of each tree .2 cm dbh was measured
and the density and basal area of each species were determined. Trees were
assigned to one of four size classes (2-10 cm, 10-20 cm, 20-30 cm and .
30 cm dbh). Similar data were collected from seven abandoned woodlots
near the pasture sites to understand current tree species composition in
relatively undisturbed forests. We found significant differences between
pasture and woodlot compositions. Pastures were dominated by conifers,
more shade-intolerant species, and wind-dispersed species (Abies balsamea
L., Populus spp., Picea rubens Sarg., Acer rubrum L.) in terms of basal
area and density. Woodlots were dominated by late-successional, shadetolerant species (Fagus grandifolia Ehrh., Acer saccharum Marsh., Tsuga
canadenisis, and Betula alleghaniensis Michx. F.). These striking differences between pastures and relatively undisturbed woodlots begin to sug-

280

Abstracts

gest that the northern hardwood forest is not resilient to this novel, widespread, and long-lasting disturbance.
RICCARDI, CYNTHIA L.* and BRIAN C. MCCARTHY. Landscape
quantification of fuel loads in central Appalachian mixed-oak forests.
Fuel loads are diverse in their physical attributes, and complex in structure
and origin of their components. As the source of all forest fire behavior
and fire effects, fuel loads must be characterized and quantified before any
calculation of fire potential can be made. The objective of our study was
to quantify the fuel loads before and after mechanical thinning and prescribed fire in three mixed-oak forests in southeastern Ohio. In each forest
a control (CU), thin (TU), thin-burn (TBU), and burn (BU) unit were established in 2000. Thinning and prescribed surface fire were applied in
winter 2000 and spring 2001, respectively. The Brown planar intersect
method was used along seventy-two 20 m transects (n5864) to measure
forest floor depth and 1-, 10-, 100-, and 1000-hour (CWD) woody fuels.
For the three forests, pretreatment mean (6SE) litter (Oi and Oe) and duff
(Oa) mass were 7.9 (60.3), 2.9 (60.19), 3.1 (60.11) and 33.1 (61.9), 5.4
(60.14) and 12.7 (60.49) MT ha-1, respectively. Pretreatment mean minimum fuel loads were 0.32 (60.01), 1.31 (60.06), and 1.47 (60.13) while,
mean maximum loads were 0.38 (60.01), 2.04 (60.08), and 1.99 (60.11)
MT ha-1, respectively. Pretreatment CWD volumes were 42.7 (60.02),
22.9 (60.02) and 40.8 (60.05) m3 ha-1. Litter and duff were significantly
decreased in TBU and BU compared to CU and TU (P,0.05). Overall,
fire did not reduce the increased woody fuels produced by the thinning but
thinning and fire did tend to decrease 1000-hr fuels. Finally, our results
indicate a high level of heterogeneity of fuel loads across the mixed-oak
forest landscape of southeastern OH.
RICHARDS, CHRISTINA L., LISA A. DONOVAN and JAMES H. HAMRICK. University of Georgia, Department of Plant Biology, 2502 Miller
Plant Sciences, Athensa, GA, USA. Genetic diversity and fine scale population structure of the salt marsh perennial Borrichia frutescens.
We used enzyme electrophoresis to evaluate fine-scale population genetic
structure of Borrichia frutescens L., a coastal salt marsh perennial in the
southeastern U.S. We hypothesized that the extreme environmental gradients in this habitat would lead to high genetic structure compared to other
outcrossing perennial species and to genetic differences among plants in
different microhabitats along the gradients. We also hypothesized that at
very fine spatial scales, extensive ramet production would cause subpopulations to be dominated by a few large clones. To test these hypotheses,
we sampled individuals from populations representing the full distribution
of this species in the marshes of Sapelo Island, GA. At 5 sites, leaf tissue
was collected from 96 individuals at 1 m intervals on grids spanning the
environmental and height gradients. Fifty-eight percent of the 17 loci examined were polymorphic, with 41.2% polymorphic within populations.
Our estimate of genetic structure for Borrichia (GST 5 0.139) was slightly
higher than other outcrossing, long-lived perennial species (GST 5 0.094,
Hamrick and Godt 1996), indicating only moderate barriers to gene flow.
There was no evidence of genetic differentiation among microhabitats within populations. Within populations, average GST among microhabitats was
0.042. Across populations, genetic diversity (He) was equivalent for the
three microhabitats (average He 5 0.089). Clonal diversity varied but populations often contained many multilocus genotypes, indicating that both
sexual reproduction and recruitment from seeds are important factors maintaining diversity. In conclusion, barriers to gene flow appear to maintain
slightly more differences among populations than would be expected for
an outcrossing perennial species. However, the environmental gradient does
not seem to have a strong influence on the fine-scale distribution of genetic
diversity at these loci.
RICHARDS, SHANE A.,1,* KATHRYN L. PATTERSON,2 MERCEDES
PASCUAL3 and JAMES W. PORTER.4 1 Biological Sciences, University
of Calgary, Calgary, AB, Canada; 2 Marine Sciences, University of Georgia, Athens, GA; 3 Ecology and Environmental Biology, University of
Michigan, Ann Arbor, MI; 4 Institute of Ecology, University of Georgia,
Athens, GA. Predicting Elkhorn Coral dynamics throughout the Florida Keys.
Since 1996 a long-term reef monitoring program has been established
throughout the Florida Keys with the aim of identifying the status and

trends in the benthic community. Initial analyses of the data have shown
significant sustained declines in coral cover and distribution, and an increase in disease prevalence for most coral species. We present a simple
mathematical model based on the metapopulation approach that describes
population dynamics of the Elkhorn Coral (Acropora palmata), which is a
species that has recently experienced pronounced declines in the region.
Predictions from this simple model are consistent with the field data and
the model provides an explanation for both the mean and the variance
observed in the data. Specifically, it suggests that local extinction rates
peaked recently; however, the rate of local coral recruitment is continuing
to decline. These trends are found to be consistent along the entire Florida
Keys. Using these estimated rates, the model predicts further declines in
abundance and continued local extinctions under a variety of environmental
scenarios.
RICHARDSON, CURTIS J.,1,* RYAN S. KING2 and SONG S. QIAN.3
Duke University, Durham, NC; 2 Smithsonian, Edgewater, MD; 3 Cadmus,
Chape Hill, NC. A bayesian estimaton of phosphorus thresholds in the
Everglades.

The Florida Everglades, an internationally designated wetlands of importance, is currently undergoing a significant shift in its native flora and fauna
due to excessive loadings of total phosphorus (TP) from agricultural runoff
and Lake Okeechobee outflow. However, the surface-water TP threshold
for sustaining the Everglades is unknown. Here, we present long-term experimental research and show that a surface water geometric mean TP
concentration of 15 ug/L maintains an ecological balance in algal, macrophyte, and macroinvertebrate populations as well as slough community
structure. Our results from a 6 year phosphorus (P) dosing experiment in
the Everglades suggest that the 10 ug/L P threshold often given for average
annual concentrations in oligotrophic waters (natural pristine lakes) is not
appropriate for defining oligotrophic wetland status in the Everglades due
to differences in ecosystem structure (periphyton versus phytoplankton)
seasonal water depth affects on P concentrations, and natural nutrient gradients (i.e., higher exterior nutrient concentrations in wetlands gradate to
lower interior nutrient levels) that exist in wetlands versus lakes. Importantly, the ecological responses to TP in the Everglades are highly predictable, but estimates of uncertainty must be utilized to accurately define
threshold responses, which change with seasons and water depths. Our
Bayesian change point methodology used to address ecological imbalance
along nutrient gradients is applicable to other aquatic ecosystems.
RICHARDSON, SARAH J.* and ROB B. ALLEN. Landcare Research,
PO Box 69, Lincoln, Christchurch, 8152, New Zealand. Seeding patterns
by mountain beech forest over 38 years in New Zealand.
Irregular seed production may have evolved to escape predation yet the
mechanisms that underpin such a reproductive strategy are poorly understood. The simplest explanation is that production of seed is limited by
resources: trees should produce more seed in favourable years and large
seeding events should be more frequent in productive environments. The
significance of temperature for seeding has been recognised across a range
of taxa in New Zealand and elsewhere, yet it remains unclear whether
temperature is simply a proxy for carbon availability or whether temperature represents a cue to expend stored reserves thus enhancing seed production in warm years. Here we present results from a time series analysis
of seed production data collected along an elevational gradient over 38
years in a monospecific mountain beech forest in South Island, New Zealand. We use elevation as a resource gradient and examine how within and
among-year variation in seed production and seed viability is affected by
resource availability. Mean seed production and viability over 38 years
were both negatively related to elevation, but there was marked year-toyear variation in these relationships. We use mixed models to test whether
the frequency of seeding increases with resource availability; whether climate exerts a greater influence in resource-poor environments; and whether
the importance of lag effects from previous seeding are affected by resource
availability.
RICKETTS, TAYLOR H.* World Wildlife Fund, Washington, DC. Analytical approaches and issues for the Millennium Assessment.
The Millennium Assessment (MA) is a global, multi-scale effort to assess
the variety of services provided by ecosystems to humanity. This enormous

geographic and thematic scope is a central strength of the MA, but it also
presents significant analytical challenges. Here I describe the overall analytical approach we have developed to achieve the MA9s goals, emphasizing several of the most pervasive issues encountered along the way. The
analytical approach comprises nine major tasks: (i) categorizing ecosystems, (ii) identifying types of ecosystem services, (iii) identifying indicators
of those ecosystems and services, (iv) identifying proximate and primary
drivers of change, (v) assessing the current status and recent trends, (vi)
evaluating impacts of changes on human well-being, (vii) developing scenarios for the future, (viii) evaluating response options, and (ix) analyzing
and communicating uncertainty. One pervasive issue involves balancing
analysis with synthesis. Although simpler to analyze individual systems
and services, assessing bundles of services from interacting ecosystems is
central to the MA and will help illuminate trade-offs inherent in many landuse decisions. Another major issue is one of spatial and temporal scale.
Ecosystems operate at a variety of scales, provide different services at
different scales, and will be assessed using data collected over various
spatial and temporal scales. Beyond the Assessment reports themselves,
devising effective ways of addressing these and other issues is one of the
major potential contributions of the MA.
RIEDER, JULIE P.* and EDWARD W. EVANS. Utah State University,
Logan, UT. Patterns of yellow starthistle (Centaurea solstitialis) emergence, establishment, and fecundity: The influence of vegetation cover.
Microhabitat characteristics strongly influence the fate and fecundity of
sessile organisms. The influence of cover type and class, two key microhabitat characteristics, on an individual9s success may be indicative of the
interplay among intraspecific competition, facilitation, and interspecific
competition. Yellow starthistle (YST; Centaurea solstitialis), a non-native
annual, exhibits a patchy distribution at our site in Weber County, Utah,
U.S.A. We studied the spatial associations between YST and its neighbors
to test for patterns consistent with and suggestive of competitive and facilitative effects for YST. During spring of 2001, more than two thousand
YST seedlings were marked across the site. Survivorship and flowerhead
production were recorded for each individual. Cover estimates for YST,
grass, forb, and bare ground were recorded at multiple scales for a subset
of individuals (n5500) and seed production of all flowerheads was measured for a further subset (n570). The probability of surviving to flowering
increased as YST cover increased. Average flowerhead production was
highest at intermediate YST cover and high grass cover. And, although
seeds per flowerhead did not show a directional trend with YST cover, total
viable seeds produced per individual decreased with increasing YST cover.
Overall, the average fecundity for YST was highest in intermediate YST
cover classes. Thus, although having many YST neighbors increases an
individual9s probability of surviving to flowering, that individual will on
average produce fewer seeds. These results may be suggestive of the interaction between competition and facilitation. Ongoing experiments are
attempting to tease apart these forces shaping YST population dynamics.
RIGGS, MICHAEL R. Department of Statistical Research, Research Triangle Institute, RTP, NC, USA. Nonparametric smoothed estimators of
population hazard functions.
Data from prospective studies of the fates of members of a cohort can be
used to estimate three interrelated functions: the cohort cumulative hazard
function H(t), the cohort hazard rate, h(t), and the cohort cumulative survival probability, S(t). The three functions can be estimated for either the
overall risk of death or for multiple competing risks of death from different
causes. In this paper, I focus on methods for the estimation of the hazard
function from data arising from experimental, quasi-experimental, and observational studies (e.g., monitoring studies). The hazard function is the
instantaneous rate of change of the cumulative hazard function, h(t) 5d/
dt[H(t)] When the survival time variable is age, the plot of h(t) gives the
probability of dying as a function of age. Thus the shape of h(t) reveals
how the risk of death changes with age in a population or cohort. Moreover,
it can be shown that the observed shape of a cohort survival curve (e.g.,
Kaplan Meier curves) is determined by the underlying hazard function.
The practical importance of the hazard function is further demonstrated by
the widespread use of parametric (e.g., exponential survival models) and
semi-parametric (e.g., Cox regression) survival models to estimate the ef-

Abstracts

281

fects of treatments and covariates on the underlying cohort hazard functions. Unfortunately, a nonparametric maximum likelihood estimator for
the hazard function does not exist; instead, a semiparametric smoothed
function is generally used to approximate h(t). Three nonparametric methods for obtaining optimally smoothed point and interval estimates of the
underlying cohort hazard functions will be discussed. These include splines,
kernel density, and penalized likelihood estimators. Examples of the application of these methods to data from experimental and observational
ecological studies will be used to illustrate how hazard plots can aid in the
interpretation of population survival and risk models.
RILLIG, MATTHIAS C.,* EMILY R. LUTGEN and CARL L. ROSIER.
University of Montana, Missoula, MT. Soil aggregation: The role of arbuscular mycorrhizal fungal species.
Soil structure (the arrangement of soil into aggregates and pore spaces) is
an important characteristic of natural or managed ecosystems, influencing
the abundance of soil-inhabiting organisms and rates of soil-borne processes. Soil biota exercise well-documented controls over soil aggregation,
with a key role attributed to arbuscular mycorrhizal fungi (AMF). AMF
are ubiquitous and abundant obligate biotrophs in the phylum Glomeromycota. While the general importance of AMF for soil aggregation is widely acknowledged, information concerning the role of diversity of AMF
species for soil aggregation is scarce. Given the large differences in physiology and life history strategy among AMF species, we hypothesized that
co-occurring AMF species will differ in their effects on soil aggregation.
Here, we specifically tested for potential differences among AMF species
in the production of the soil protein glomalin. There is a strong and positive
curvilinear relationship between soil glomalin pools and soil aggregate water stability. In greenhouse experiments (with compartmentalized pots separating a hyphal compartment from a root-hyphae compartment) using the
annual grass Bromus hordeaceus as plant host and several AMF species
(from the same fungal community and the genera Glomus, Acaulospora,
Gigaspora, and Scutellospora), we found significant differences in glomalin yield (expressed on a hyphal length basis). Due to the strong relationship between glomalin and aggregate stability, this result strongly points
to differences in soil aggregation as a function of AMF species. Also, since
glomalin yield was related to hyphal palatability across different species,
an indirect effect of glomalin on soil aggregation may exist via the consequences of glomalin for differential AMF hyphal persistence in soil.
RILOV, GIL.* University of Canterbury, Christchurch, New Zealand, New
Zealand. Scaling top-down effects: Can variable recruitment and fish
predation explain meso-scale mussel distribution?
Rocky intertidal community structure can vary across wave-exposure and
biogeographical scales. Differences may also occur at regional meso-scales
(1-10s kms), and may be driven by nearshore oceanographic conditions
that affect nutrient and larvae supply (bottom-up effects). In New Zealand,
it has been suggested that upwelling conditions on the west coast enhance
top-down predation effects, while on the east coast both effects are weak
and macro predators (seastars and whelks) are rare. However, there is considerable localized variation. Within 2 km of shoreline on the east coast
are sites with different mussel cover, species abundance and size structure.
Sites surrounded by sand and with no adjacent subtidal reefs (+S-ST) had
high mussel cover and plenty of small mussels. On sites with no sand and
with subtidal reefs (-S+ST), small mussels were rare in the low shore.
Using a comparative-experimental approach, I demonstrate that large mobile predators (probably fish) can have strong and rapid predation effects
on small mussels in the low shore; there, most mussels transplanted were
removed within a day in plots unprotected from such predators, while slower mortality was evident in the mid shore. There was a strong and rapid
predation effect on two sites (-S+ST), and no such effect on +S-ST sites.
Orders-of-magnitude differences in recruitment rates among sites were also
found, with the high values comparable to those measured in the west coast.
Fish can be important predators on small intertidal mussels, and can thus
play a major role in shaping mussel distribution and populations structure,
especially where recruitment is relatively poor or sporadic. I suggest that
highly localized oceanographic, biotic and demographic differences may
account for site-to-site variation. This work is part of the PISCO/Mellon
projects.

282

Abstracts

RINGOLD, PAUL L.,1,* TERESA MAGEE2 and JOHN VAN SICKLE.1

US EPA, ORD, Western Ecology Division, ringold.paul@epa.gov, Corvallis, OR; 2 Dynamac Corporation, Corvallis, OR. Distribution of selected invasive plants in riparian ecosystems of the western United
States.

Riparian ecosystems typically exhibit high levels of plant species richness,

physical disturbance, and interconnectedness; characteristics that may favor
establishment and spread of invasive plant species. To assess the magnitude
of this invasion, we organized an extensive survey over 12 western states
for 1 cryptogenic and 11 alien, invasive plant taxa (Arctium minus (Common burdock), Arundo donax (Giant reed), Bromus tectorum (Cheatgrass),
Carduus nutans (Musk thistle), Cirsium arvense (Canada thistle), Dipsacus
fullonum (Teasel), Elaeagnus angustifolia (Russian olive), Euphorbia esula
(Leafy spurge), Hedera helix (English ivy), Phalaris arundinacea (Reed
canarygrass), Rubus discolor (Himalayan blackberry), and Taramix spp.
(Saltcedar)). This work is part of the four year EMAP (Environmental
Monitoring and Assessment Program) Western Stream Pilot, which focuses
on the assessment of diverse stream properties, using a probability sample
of 1200 streams. Target species were selected based on invasive character,
widespread distribution, frequent or potentially increasing occurrence in
streamside settings of the study area, and for ease of identification. Field
crews were trained to recognize 6 to 11 of the target species, and each
crew searched for the list of species designated for the state(s) they sampled. Remeasurements by expert crews in some states and by repeat sample
visits in all states document a high level of repeatability. At 300 sites
sampled during 2001, the target weeds most frequently observed in the
states where they were surveyed included C. arvense at 31% of sampled
streams; E. esula at 21% and P. arundinacea at 13%. In general, and
consistent with numerous plausible mechanisms, all target species were
more frequently found on higher vs. lower order streams. For example, C.
arvense was found at 51% of surveyed streams with Strahler order 4 or
greater and only 19% of streams with Strahler order 1 or 2.
RINKES, ZACHARY L.* and BRIAN C. MCCARTHY. Ohio University,
Athens, Ohio. Eastern wild turkey (Meleagris gallopavo silvestris) litter
disturbance and its effect on hardwood regeneration.
Seed germination amongst hardwood tree species is strongly influenced by
the abundance of leaf litter, which affects water availability and light penetration. Wild turkeys create a heterogeneous forest floor mat by scratching
for food. The objective of this study was to evaluate this disturbance regime
and its effects on Quercus alba, Q. velutina, Fagus grandifolia, and Acer
rubrum seed germination. Two mixed oak forests in southeastern Ohio were
selected for study. Both contained various experimental management units
(Burned, Thinned, Thinned & Burned, and Undisturbed). Seeds were planted inside compartmentalized exclosures (N 5 8), each containing a control
(ambient litter) and treatment (litter experimentally scratched and removed). Treatments were monitored every two weeks for a six-week period. A 4-way ANOVA detected a significant difference among species (P
, 0.01) and species 3 treatment interaction (P , 0.01), but no significant
(P . 0.30) difference in germination between sites or among silvicultural
units. The nut species had better germination success (P , 0.05) in the
unscratched plots, while A. rubrum did not respond to the scratching treatment. Quercus spp. also exhibited a delayed germination response, as most
did not germinate until the latter part of the experiment. Our results suggest
that re-bounding turkey populations and their indirect effects (litter redistribution) may differentially affect the regeneration of species in oak-dominated forests.
RITCHIE, JERRY C.,1,* GREGORY W. MCCARTY2 and ERIK R. VENTERIS.2 1 USDA ARS Hydrology and Remote Sensing Laboratory, Beltsville, MD, US; 2 USDA ARS Environmental Quality Laboratory, Beltsville,
MD, US. Soil carbon redistribution in an Iowa agroecosystem.
Soil carbon is the largest component of the terrestrial carbon cycle. Soil
carbon varies with climate, topography, biological activity, soil movement,
and land management leading to large uncertainties in the soil carbon budget, especially in agricultural areas. This study was designed to determine
soil carbon redistribution in an Iowa corn field as related to soil movement.
Soil movement (erosion/deposition) was measured using radioactive fallout

cesium-137. Soil organic carbon had an order of magnitude difference in

concentration (0.5 to 5%) and was significantly correlated to soil cesium137 concentration and soil erosion/deposition rates. Sites of soil erosion
have lower concentrations of organic carbon (2.4 %) while sites of soil
deposition within the field had higher concentrations of soil carbon (3.4
%). This study shows the importance of being able to measure soil redistribution patterns within a field to understand soil carbon patterns and the
potential of developing or implementing better management systems to
increase carbon sequestration in agricultural areas.
RITLAND, DAVID B.* Erskine College, Due West, SC. Life is complicated: Interacting dimensions of complexity in a "simple" mimicry
relationship.
Defensive mimicry involves one prey species (the mimic) that gains some
protection from predation through its superficial resemblance to a second,
defended prey species (the model). The model/mimic relationship of monarch (Danaus plexippus) and viceroy (Limenitis archippus) butterflies has
for a century been widely cited as a clear-cut case of defensive mimicry.
However, studies over the past decade on viceroys, monarchs, and another
putative model, the queen (D. gilippus), have revealed that the viceroys
mimicry relationships are spatially and temporally dynamicanything but
"clear-cut." Here, I summarize our current understanding of how interactions among larval hostplant geography and phenology, butterfly palatability and abundance, and predator behavioral responses affect the continuing evolution of mimicry relationships.
RITTER, AMY F.* University of California, Santa Cruz, ritter@biology.
ucsc.edu, Santa Cruz, CA. Cannibalism between the tides? The indirect
effects of habitat variation on the abundances of different life history
stages of intertidal fishes.
Variation in habitat characteristics can have important influences on the
spatial distribution and densities of organisms, and these influences can
vary throughout the life history of an organism. I examined the effects of
habitat variation in intertidal pools on the adult and juvenile abundances
of intertidal fishes. Previously, I found that some species of intertidal fishes
demonstrate a positive relationship with the surfgrass or algal cover of
tidepools. In this study, I directly examined the role that surfgrass (Phyllospadix spp.) plays in determining both adult and juvenile abundances by
manipulating levels of surfgrass cover. I determined that adult fish of one
species, Oligocottus snyderi, demonstrate a positive relationship with surfgrass cover, but that surprisingly O. snyderi juveniles exhibit a negative
relationship. To determine if this pattern was due to an ontogenetic change
in habitat preference or due to interactions between adults and juveniles, I
also conducted a number of laboratory experiments. I found that despite a
preference for surfgrass cover in both life history stages, the presence of
adults caused juveniles to avoid adults, and thus surfgrass habitat. The
adult-avoidance behavior demonstrated by juveniles appears to be driven
by cannibalism. These field and laboratory studies suggest that the contrasting distribution of O. snyderi juveniles and adults with respect to surfgrass cover is a result of interactions between juveniles and adults, leading
to an ontogenetic shift in habitat use.
RIVERS, JAMES W.,1 MELINDA D. SMITH,2,* JEFFREY S. PONTIUS,3
PHILIP S. GIPSON4 and DONALD P. ALTHOFF.4 1 Department of Ecology, Evolution, and Marine Biology, University of California, Santa Barbara, CA, USA; 2 National Center for Ecological Analysis and Synthesis,
Santa Barbara, CA, USA; 3 Department of Statistics, Kansas State University, Manhattan, KS, USA; 4 Division of Biology, Kansas State University,
Manhattan, KS, USA. Long-term dynamics of grassland bird communities with and without anthropogenic disturbance.
Anthropogenic activities are predicted to change the composition of communities and promote the loss of species unable to withstand human-caused
perturbations. Yet, field studies investigating human impacts often lack appropriate reference sites and are limited in duration. We compared two
native tallgrass prairie sites in Kansas, one heavily impacted by humans
(Ft. Riley Military Installation) and one with minimal human impacts (Konza Prairie Biological Station), to assess the effects of human activities on
long-term dynamics of bird communities. We also used Breeding Bird Sur-

vey data to characterize the regional pool of potential colonizers and examine its influence on local dynamics. From 1991-2001, most of the small,
terrestrial landbird species (73%) recorded during breeding surveys were
found on both sites with similar relative abundance. For the study period,
mean annual richness on Ft. Riley (39.0 60.86SE) was similar to that of
Konza Prairie (39.4 60.61SE). Richness was maintained at markedly constant levels despite compositional changes because colonizations compensated local extinctions at both sites. These dynamics were driven primarily
by stochastic loss and gain of species that were restricted to woodland
habitats and present at low local and regional relative abundance. The similar community dynamics observed between the two sites suggests that
anthropogenic activities may mimic the natural disturbances (fire, grazing)
found at both sites and that species richness is an emergent property of
ecosystems often maintained within narrow limits.
RIXEN, CHRISTIAN1,2 and CHRISTA P.H. MULDER.1 1 University of
Alaska, Fairbanks, AK; 2 Swiss Federal Institute for Snow and Avalanche
Research SLF, Davos, Switzerland. Biodiversity and ecosystem functioning: Testing the insurance hypothesis in moss communities.
The relation of biodiversity and ecosystem functioning has been addressed
in an increasing number of studies in the last years due to growing concerns
about human-induced species loss in many ecosystems. Most studies have
concentrated on diversity-productivity relations. Only few studies have addressed the role of the biodiversity as an insurance for ecosystem functions
in a changing environment (insurance hypothesis) and mechanisms underlying interactions of diversity and ecosystem functioning. We carried out
greenhouse experiments on diversity and ecosystem functioning with moss
communities. We selected species from a larger species pool to create communities with different diversity levels. To test the relationship between
diversity and productivity, these communities were either exposed to experimental drought stress or kept at constant conditions. Humidity was
measured to describe the microclimatic conditions. Under constant conditions no relationship between biodiversity and biomass could be found.
When stressed with experimental drought, biomass increased with increasing diversity. Our results suggest that biodiversity can be an insurance to
maintain ecosystem functions in changing environmental conditions. Facilitative interactions between plants like maintaining a high humidity may
be an important mechanism for ecosystem functioning in divers communities.
ROBERTS, BRIAN J.,1,* JON J. COLE,2 MICHAEL L. PACE,2 DARREN
L. BADE3 and MATTHEW C. VAN DE BOGERT.2 1 Cornell University,
bjr11@cornell.edu, Ithaca, NY, USA; 2 Institute of Ecosystem Studies,
Millbrook, NY, USA; 3 University of Wisconsin-Madison, Madison, WI,
USA. Diel respiration measured using oxygen stable isotopes in fertilized, clear, and colored lakes.
Diel patterns in respiration rates were examined in three lakes as part of a
seasonal study on carbon cycling. Peter Lake received N and P fertilizations
resulting in a net autotrophic state while net heterotrophy was maintained
in Paul (clear) and Tuesday (colored) lakes. Continuous measurements of
O2 and CO2 were made throughout the experiment. Daytime rates of respiration were quantified by making multiple daytime measurements (2-6
sampling times) of both 16O and 18O from each lake on five different dates.
Respiration rates were greater during day than night throughout the experiment. Significant differences in day: night R ratios existed between lakes.
The highest daytime respiration rates were in the fertilized lake (Peter)
where day R was much greater than night R. The clear, unmanipulated lake
(Paul) exhibited intermediate day: night ratios while the day: night R ratios
in the colored lake (Tuesday) were consistently lowest. These results have
implications for accurately estimating gross oxygen evolution and consumption rates in lakes.
ROBERTS, KELLY A. and LLEWELLYN M. EHRHART.* Department
of Biology, University of Central Florida, P.O. Box 162368, Orlando, Florida. Impacts of beach nourishment on loggerhead and green turtles in
Brevard County, Florida.
Marine turtle nesting habitat in Florida is compromised by beach erosion
and coastal development. Beach erosion may be mitigated by beach nour-

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283

ishment projects which restore the beach profile and in most cases create
additional marine turtle nesting habitat. The impact of beach nourishment
on loggerhead (Caretta caretta) and green turtle (Chelonia mydas) nesting
on a Florida beach was investigated as the result of the Brevard County
Shore Protection Project, South Reach (Phase II). The approach to the
investigation involved examination of three factors that are integral to the
matter of marine turtle reproduction: 1) overall nest production, 2) nesting
success (ratio of nest to non-nesting emergences), and 3) reproductive success (hatching and emerging success). Results of this study indicate a significant decrease in nest production and a significant increase in non-nesting emergences, thus a decrease in nesting success for both species in the
nourished beach as compared to the control. Nests incubated in the nourished substrate produced equal (loggerhead) or significantly higher (green
turtle) reproductive success rates than in the adjacent non-nourished area.
This paper brings together the compilation, analysis and interpretation of
the effects of the nourished substrate on marine turtles.
ROBERTSON, KEVIN M.1,2,* and CAROL K. AUGSPURGER.1 1 Department of Plant Biology, University of Illinois U-C, Urbana, Illinois,
USA; 2 Tall Timbers Research Station, Tallahassee, Florida, USA. Spatial
segregation of floodplain tree species through differential mortality of
seedlings.
Cohorts of tree seedlings were followed and patterns of seedling recruitment were studied on an elevation gradient in the first stage of riparian
forest primary succession to determine 1) the degree to which spatial segregation among species arises as a result of differential mortality following
emergence and 2) the consistency in spatial patterns of survival among
years. In a previous study, pioneer trees on the Bogue Chitto River, Louisiana, U.S.A. were observed to be spatially segregated by species along
river bend point bars in association with an elevation gradient on each bar.
In this study, one natural cohort of seedlings consisting of ten species was
studied on one river bend for three years to measure annual changes in
distributions of seedlings with regard to elevation, soil texture, light, and
distance from the upstream end of the point bar. Also, four cohorts of
seedlings were studied at the 1-yr old stage in each of four years for 11
species to determine the year-to-year consistency of species distributions.
In general, mortality of seedlings and resulting changes in their spatial
distributions was greatest during the first year following emergence. Distributions changed from being greatly overlapped among species to being
segregated by species along the point bar, and changes were most strongly
associated with elevation. Locations of recruitment into the 1-yr age category differed among species and were consistent among years, suggesting
a predictable influence of the environmental gradient on spatial patterns of
seedling survival. Locations of species along the gradient corresponded to
life history characteristics measured in this and other studies. In conclusion,
spatial segregation of species along environmental gradients arises during
plant development through differential mortality that reflects plant life-history characteristics.
ROBERTSON, TRAESHA R.,* ERIN M. WALKER, JOHN C. ZAK and
DAVID T. TISSUE. Texas Tech University, Department of Biological Sciences, Lubbock, TX. Plant responses to variation in timing and magnitude of precipitation in Big Bend National Park.
Water availability is the primary limiting factor for plant growth, soil nutrient dynamics, and ecosystem productivity in arid ecosystems. Plant species have different responses to water availability that are dependent on
photosynthetic pathway, plant morphology, and root architecture. We tested
effects of predicted future changes in precipitation patterns and water availability by altering the precipitation received by the dominant plant species
of the sotol-grassland community: sotol (Dasylirion leiophyllum; C3), sideoats grama (Bouteloua curtipendula; C4) and brownspine prickly pear
(Opuntia phaeacantha; CAM). The first experiment investigated how a
25% increase in summer and winter precipitation affected plant photosynthetic performance, growth and species competition. A second experiment
used rainout shelters to exclude natural rainfall and then we added rainfall
pulses of different amounts and frequency to the plots. Plants received
rainfall as small but frequent pulses, pulses of moderate size and frequency,
or large but infrequent pulses to determine how a shift in pulse timing and
magnitude might affect plant photosynthetic performance. In the first year

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Abstracts

of study, following the summer watering of 2002 and the winter watering
of 2003, we found that the 25% increase in water availability significantly
increased the photosynthetic performance of grama (C4) during the summer
and sotol (C3) during the winter. However, the growth response (i.e. changes in leaf area, leaf volume, and biomass) was not altered for any species
in either season. In the rainout shelter experiment, small, frequent summer
rain pulses increased photosynthesis in grama, while larger, less frequent
pulses in both the summer and winter increased photosynthesis in sotol.
Therefore, differences in pulse size and frequency may differentially affect
photosynthesis in these desert species. It remains to be determined whether
differences in carbon assimilation will change long-term plant growth.
ROBINSON, JULIE A.,1,2,* SERGE ANDREFOUET,3 JENNIFER GEBELEIN,4 ALAN SPRAGGINS,1,5 EDMUND P. GREEN,6 MARCO
NOORDELOOS7 and LAURETTA BURKE.8 1 Earth Science & Image
Analysis Laboratory, NASA Johnson Space Center, julie.a.robinson1@
jsc.nasa.gov, Houston, TX, USA; 2 Lockheed Martin Space Operations,
Houston, TX, USA; 3 University of South Florida, St. Petersburg, FL, USA;
4
Florida International University, Miami, FL, US; 5 Hernandez Engineering, Houston, TX, USA; 6 United Nations Environment Programme, World
Conservation Monitoring Centre, Cambridge, UK; 7 International Center
for Living Aquatic Resources Management, World Fish Center, Penang,
Malaysia; 8 World Resources Institute, Washington, DC, USA. Global coral reef mapping and data distribution: Partnerships between remote
sensing scientists and international agencies.
Natural resource managers around the world desperately need maps of coral
reefs and adjacent land areas. The current level of knowledge about such
simple measures as the total area and locations of coral reefs in the world
is not sufficient as a baseline for monitoring change. Regional studies attempting to identify the risk factors for decline of coral reefs are also faced
with inadequate maps of reefs and adjacent land uses. At the same time,
local managers clamor for detailed reef habitat maps for monitoring smaller
scale changes in reef communities. These managers are making daily decisions that impact the health of coral reefs and the economies of the communities that depend on them. In a NASA-sponsored partnership between
remote sensing scientists, international agencies and NGOs, new efforts are
being made to develop the baseline global reef map that can be a foundation
for future more detailed investigation. Global mapping protocols developed
region-by-region allow improvement of the existing reef presence absence
map maintained by the United Nations Environment Programme-World
Conservation Monitoring Centre (UNEP-WCMC). At the same time, these
protocols produce a set of global reef areas maps showing major geomorphologic features and bottom types. We discuss the tradeoffs between different mapping objectives and abilities in meeting various user needs. A
set of partnerships between researchers and UNEP-WCMC, ReefBase (International Center for Living Aquatic Resources Management), and the
World Resources Institute have been built to improve the relevance and
distribution of global coral reef mapping data. We will describe the transition from research to operational methods using Landsat-7 data for operational production of land cover / shallow reef maps. We discuss the
challenges faced in developing GIS-based distribution networks, and demonstrate how UNEP-WCMC and ReefBase will distribute such maps in
support of the activities of international development agencies and local
resource managers.
ROBINSON, TODD M.P.* and KATHRYN L. COTTINGHAM. Dartmouth College, Hanover, NH. Differential nitrogen exploitation by
knapweed and bluebunch wheatgrass.
The replacement of endemic species by invasive species is an important
and growing problem with non-indigenous weed management costing up
to $5B annually in the U.S. For example, knapweed (Centaurea maculosa
and C. diffusa) is an invasive weed common in the western and midwestern
United States that replaces native grasses, increasing erosion and decreasing
forage quality. We used a 3-month greenhouse experiment to determine the
relative impacts of knapweed versus the native bluebunch wheatgrass
(Pseudoroegneria spicatum) on plant-available soil nitrogen. At the end of
the growing period, we used KCl extractions to extract plant-available nitrogen and determined nitrate and ammonium concentrations (1) in fresh
soil extracts and (2) extracts from soil incubated for 5 weeks without plants

to evaluate microbial effects. Ammonium concentrations were very low,

so we focused our analyses on nitrate. Multiple regressions indicate that
plant-available nitrate in both fresh and incubated soil declined with knapweed and bluebunch aboveground biomass; there was also a significant
interaction between species for the freshly extracted soil and a significant
quadratic term for knapweed in the incubated soil (Fresh: R250.67,
p,0.0001; incubated: R250.86, p,0.0001). These models strongly suggest
that knapweed and the native grass have nonlinear effects on plant-available nitrate concentration, but that the exploitation is asymmetric: knapweed appears to be the superior competitor. We conclude that belowground
competition for nitrogen may be an important component of knapweeds
invasive success.
ROBISON, LAURA A.,1,* SCOTT K. GLEESON,1 DEVETTA HILL1 and
TOM BIEBIGHAUSER.2 1 University of Kentucky, Lexington, KY; 2 United States Forest Service, Morehead, KY. Colonization of constructed
wetlands in the Daniel Boone National Forest.
In the Daniel Boone National Forest Area, a series of restoration projects
seeks to counter a long-standing trend of wetland loss by restoring former
wetlands and creating new ones in the area. It is hoped that the new wetlands will provide a wide spectrum of benefits, from water holding and
filtration, to increasing water availability to wildlife, to creating habitat for
hydrophytic plants. The focus of this study is a group of small to mediumsized ponds created during 1999, 2000, and 2001 as part of these projects.
Though no wetland plants were planted or seeded during construction, a
number of these plants have colonized the ponds on their own, and new
ecological communities have developed. To investigate the successional
progress of these communities, vegetation data was collected from each
pond, and compared with data from four nearby natural ponds. Vegetation
was sampled by creating a list of all species present either in the water or
within the hydrological fluctuation zone. Pond size, depth and soil type
were also measured. Data ordination of plant species composition showed
significant separation between natural and constructed community types.
This difference is interpreted as a combination of successional/colonization
effects and environmental conditions. The constructed ponds were located
in mowed clearings and may not closely resemble their natural counterparts
until their surrounding upland succeeds to forest.
RODRIGUEZ, SUSANA,1,* MARIA A. ORJUELA2 and GLORIA GALEANO.2,3 1 The Ohio State University, Department of Evolution, Ecology
and Organismal Biology, Columbus, Ohio, Usa; 2 Univesidad Nacional de
Colombia, Biology Department, Bogota, Colombia; 3 Instituto de Ciencias
Naturales, Univesidad Nacional de Colombia, Bogota, Colombia. Demography and life history of Geonoma orbignyana: An understory palm
used as foliage in Colombia.
Geonoma orbignyana is a common understory palm, which leaves are used
as foliage in Colombia. Currently, no information about its ecology or
conservation status is available. Our objective is to characterize the life
history and population dynamics of this palm as a first step to assess the
effect of leaf harvesting. More than 2000 palms were monitored from June
1999 to March 2000; we measured growth, mortality, and reproduction
success. Population dynamics were evaluated using Lefkovitch matrix model, and elasticity analysis. Results indicate that G. orbignyana behaves according to its understory condition. The growth rate is low (1.98 leaves/
year), the palm has a long lifespan, and growth and reproduction are related
with the light conditions in the forest. The density is high (21.061 palms/
ha), which is explained in part by its capacity of recovery after damage,
and the frequent gap formation in the forest. The population is growing
(l5 1.0585). Elasticity values indicate survival is the most important event
(0.77), although growth is higher than in other understory palms (0.204).
Because of the high density, the importance of growth transitions, and the
recovery capacity of this palm, we hypothesize G. orbignyana behaves as
an opportunistic species taking advantage of the forest gaps. This behavior
would make this palm a promising Non Timber Forest Product (NTFP)
because management plans could use this behavior to ensure the leaf availability and the persistence of the populations.

RODRIGUEZ-SAONA, CESAR R.* and JENNIFER S. THALER. University of Toronto, Department of Botany, 25 Willcocks St., Toronto, ON,
Canada. Distribution of arthropods in homogeneously and heterogeneously induced plant patches.
Plant responses to herbivore damage can affect future colonization of herbivores and their natural enemies. In nature, plants commonly occur in
patches, surrounded by other plants of the same species. Field experiments
were conducted to address how varying the number of plants damaged by
herbivores within a patch affect colonization of herbivores and their natural
enemies. Because herbivore damage causes changes in plants that can negatively affect the performance of herbivores and also induces volatiles that
attract natural enemies to these plants, we hypothesized that patches with
higher amounts of herbivore damage might contain reduced number of
herbivores, while recruiting higher numbers of natural enemies of the herbivores. To test this hypothesis, tomato plants were randomly assigned into
one of three patch types: homogenoeus patch (damaged), where all plants
within a patch were damaged by placing a S. exigua larva on each plant;
heterogeneous patch, where one plant per patch was damaged; and homogeneous patch (undamaged), where plants within the patch received no
damage. All patches consisted of three tomato plants. The data supported
our predictions that herbivores are more abundant on undamaged homogeneous patches, and predators are more abundant in patches with high
herbivore damage. Omnivores responded more similarly to herbivores than
predators. In heterogeneous patches, damage to plants did not affect the
number of arthropods on undamaged neighboring plants.
ROGERS, KEVIN H.1 and STEWARD T.A. PICKETT.2,* 1 The University
of the Witwatersrand, Johannesburg, Wits; 2 Institute of Ecosystem Studies,
Millbrook, NY, USA. Historic over-allocation of water resources: The
Kruger National Park case.
Water abstraction for new and expanding settlements and for agriculture
and forestry continue to reduce flows in the major rivers that traverse the
Kruger National Park in South Africa. Several of these rivers have been
perennial or predictably seasonal throughout the history of the park. However, the hydrographs have been increasingly altered, and flows have been
drastically reduced so that perenniality or regularity is threatened in several
of the large rivers. The connections between the rivers and the upland
savannas are proving to be a crucial link in the Kruger ecosystem. Yet,
even as the links are documented they are undermined by the altered flow
regimes. Adaptive learning is a management strategy that links communities outside the park in the upstream portions of the catchments, with researchers, and with managers in the Kruger National Park. The South African constitution mandates legal status to the environment and shared governance, therefore promoting a creative management approach to the anthropogenic drying of the Kruger Rivers. Similarities with urban
hydrological drought are noted.
ROHANI, PEJMAN.* Institute of Ecolgy, University of Georgia, Athens,
Georgia, USA. Noise and nonlinearity in epidemics of childhood diseases.
I will explore the relative importance of deterministic forces versus processes noise by studying the temporal dynamics of two childhood infections, measles and whooping cough, in England and Wales. I demonstrate
that epidemics of whooping cough are strongly influenced by stochasticity;
fully deterministic approaches cannot achieve even a qualitative fit to the
observed data. In contrast, measles dynamics are extremely well explained
by a deterministic model. These differences are shown to be caused by
their contrasting responses to dynamical noise due to different life-history
traits. I will also discuss the role of stochasticity in determining the relative
phases of epidemics when different pathogens "compete" for susceptibles.
ROHR, JASON R.,1,* ADRIA A. ELSKUS,1 BRIAN S. SHEPHERD,1
PHILIP H. CROWLEY,1 THOMAS M. MCCARTHY,1 JOHN H. NIEDZWIECKI,1 ANDY SIH1,2 and BRENT D. PALMER.1 1 University of Kentucky, Lexington, KY; 2 University of California at Davis, Davis, CA. Effects of the herbicide atrazine and food limitation on the survival, lifehistory, and behavior of streamside salamanders.
Amphibian populations can be affected adversely by biotic and abiotic
stressors that together can contribute to their global decline. Consequently,

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285

we evaluated the effects of food abundance (limited and unlimited food)

and exposure to ecologically relevant concentrations of atrazine (actual
concentrations: 0, 4, 40, 400 ppb), possibly the most abundantly used herbicide in the world, on the survival, life-history, and behavior of streamside
salamanders, Ambystoma barbouri, from embryo through metamorphosis.
In general, food and atrazine levels did not interact statistically, and atrazine
concentration was correlated positively with effect size. Exposure to
400ppb of atrazine decreased embryo survival and increased time to hatching and variation in hatching day. This greater hatching variation amplified
larval size variation, facilitating cannibalism. Both cannibalism and limb
loss due to biting were more common in food-restricted groups. Despite
the significant cannibalism, neither atrazine nor food restriction had overall
significant effects on larval survival. However, food-limitation increased
larval activity and decreased shelter use, while increasing atrazine concentrations decreased activity and shelter use. Our approach triggered a fleethen-freeze response by larvae exposed to 400 ppb of atrazine. Both food
restriction and 400 ppb of atrazine reduced larval growth and size at metamorphosis without affecting body condition, despite atrazine having no
effect on feeding rates through ontogeny. Food limitation delayed metamorphosis, while 400 ppb of atrazine shortened the larval period, but the
length of the larval period and size at metamorphosis were correlated positively regardless of food or atrazine level. Metamorphic responses to atrazine, food limitation, and release from competition were consistent with
the predictions of the Wilbur and Collins amphibian life-history model.
Because reduced growth and smaller size at metamorphosis can lower terrestrial survival and lifetime reproduction, resource limitation and environmentally realistic concentrations of atrazine have the potential to contribute
to amphibian declines in impacted systems.
ROHRS, JENNIFER K.* and CHRISTA MULDER. University of Alaksa,
Fairbanks, AK, USA. The effect of permafrost distribution on soil water
chemistry and understory nitrogen dynamics in a boreal forest watershed.
Caribou Poker Creeks Research Watershed (CPCRW) is a boreal watershed
where permafrost is discontinuous and distributed mainly in the valley
bottoms and north facing slopes. In this high latitude watershed, permafrost
is tightly coupled with aspect, tree shading, and moss insulation to determine vegetation composition. The goal of this study is to determine changes in soil water chemistry and understory seasonal foliar nitrogen dynamics
from the valley bottoms to the upland regions of high and low permafrost
hillslopes in the CPCRW. This investigation of foliar nitrogen dynamics is
based on the difference in permafrost coverage between 1) low and high
permafrost hillslopes and 2)valley bottoms and upslope regions of the watershed. Determining foliar nitrogen flux in the understory may be especially significant given the predicted trends in climate warming, permafrost
thaw, and resulting shifts to a more productive community composition.
The common understory species Rumex, Vaccinium uliginosum, Vaccinium
vitis2idaea,Ledum groenlandicum, and mosses Polytrichum and Pleurozium were sampled throughout the summer of 2002 to assess differences
in tissue nutrient concentrations from upslope to valley bottom regions.
Results from the 2002 field season show that active layer depth consistently
increases from the valley bottoms to the upland regions of southern facing
slopes. Northern facing, high permafrost slopes did not have this gradient
in thaw depth. Soil temperature consistently increases from the valley bottoms to the upland regions at all sites. In upland areas that have greater
thaw depth and nutrient availability, I expect that the understory vegetation
will maintain similar maximum foliar nutrient concentrations as vegetation
in areas with more extensive permafrost. However, I predict that the upland
understory species will take advantage of the increased nutrient availability
by decreasing the amount of nitrogen resorbed at the end of the growing
season.
ROMANUK, TAMARA N.1,* and JUREK KOLASA.2 1 University of Calgary, Calgary, Alberta, Canada; 2 McMaster University, Hamilton, Ontario,
Canada. Lessons from rock pools in Jamaica - microcosms, models
systems, or simply convenient ecological systems?
Coastal dissolution rock pools span a broad range of physical conditions
and are occupied by many aquatic invertebrates forming natural multitrophic communities. In the main, attributes of these communities (rank-

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abundance curves, diversity patterns, and others) either mimic or are within
range of other common ecological systems such as lakes, coral reef patches,
streams, or fragmented forests. Thus, using the rock pool communities, one
can gain insights into many general ecological phenomena that are less
tractable at larger spatial or temporal scales. We review recent insights
gained from the rock pool system. Specifically, using data spanning 12
years of observations and monitoring on 50 natural rock pools inhabited
by over 70 species of aquatic invertebrates, we show that (i) a global
gradient of diversity toward the tropics may be caused by differential extinction of habitat specialists in more variable habitats; (ii) relationship
between population stability and community diversity is modulated by habitat variability; (iii) metapopulations may respond to richness more consistently than local populations; and (iv) environmental forcing leads to a
considerable convergence of community structure following perturbation.
We compare these findings to those obtained from studies conducted in
larger systems and argue that any differences that can be identified between
the small and large systems do not affect the generality of results. We thus
conclude that rock pools offer a suitable system for research on general
and invariant properties of ecological communities and that their usefulness
is particularly high for testing ecological theory. Where system specific
questions are of great importance as, for example, in large and unique lakes,
natural microcosms offer only limited help.
ROOSENBURG, WILLEM M.1 and WILLIAM L. KENDALL.2 1 Department of Biological Sciences, Ohio University, Athens, OH, USA; 2 USGS,
Patuxent Wildlife Research Center, Laurel, Maryland, USA. Factors contributing to variation in population level process in the diamondback
terrapin.
We will present the analysis of a 16-year mark-recapture database of a
diamondback terrapin (Malaclemys terrapin) population in Chesapeake
Bay. Our population was studied from 1987 through 2002 and accumulated
more than 26,000 capture records of more than 9,000 individuals. While
sampling we employed a variety of capture techniques to minimize sex and
size related bias of a single trapping technique. During 2000 the study was
exposed to an oil spill. We collected data of age, size and reproductive
status of all individuals. We analyze these data with open-robust design
estimation techniques to derive standard demographic parameters and to
identify time-dependent variation in capture probabilities as well as estimates of fecundity, survivorship and immigration. In our analyses, we test
for the effects of environmental factors and the effects of the oil spill on
population-level processes.
ROOT, KAREN V.* Bowling Green State University, Bowling Green, OH.
A multispecies approach to ecological assessment and conservation in
Florida.
The conservation of ecosystems focuses on evaluating individual sites or
landscapes based on their component species. I developed methods to compute the community-level risk of extinction for a site and assign a multispecies conservation value using a spatially-explicit weighting of the species-specific habitat suitability maps by their respective species-specific extinction risks. These methods were applied to a set of species of South
Florida that were federally listed or of special concern to create a series of
multispecies conservation value maps. The multispecies conservation value
provided an independent measure of the value of a particular site based on
its ecological components using both the habitat suitability and risk of
extinction for each these species. The resulting maps highlighted areas that
were of greatest importance to the most vulnerable species, such as the
remaining forested habitat and critical wetland areas. Also, the results demonstrated that these methods were flexible and could accommodate the
quantity and quality of data available for each individual species both in
the development of the habitat suitability maps and the estimation of the
extinction risks. This method provides an important tool to prioritize conservation and recovery actions, such as the ongoing Everglades restoration,
and evaluate land preservation alternatives.
ROSE, JOSHUA S.* Duke University, Durham, NC, USA. Ecology of
exotic dragonfly species in different native assemblages.
To determine whether exotic species express the same ecology under different ecological contexts or whether these contexts alter their ecology, the

ecological niches of dragonfly species were quantified in three bioregions:

the Everglades of south Florida, the Lower Rio Grande Valley of south
Texas, and the main islands of the Hawaiian archipelago. Florida and Texas
each possess many more species of dragonflies than Hawaii, and have
many more types of predators and prey as well. Some dragonfly species
are indigenous to all three regions, but most to one or two; a few species
occur as nonnatives in two regions, or are native in one but have invaded
another. Several species in Texas appear to use shaded and dry habitats
more extensively than conspecifics in Florida. In Hawaii neither indigenous nor exotic species appear to express wider niches than their conspecifics in Texas and Florida, nor is niche overlap among species in Hawaii
perceptibly lower. However, populations of some species do appear to express different ecological niches between two or three of these regions.
Nonnative species occur predominantly in altered or artificial habitats in
Florida and Hawaii; Texas has only one invading species, which occurs
in apparently undisturbed habitat.
ROSENFELD, KRISTEN M.,* ELLEN DAMSCHEN, DEENA MURPHY-MEDLEY, THOMAS R. WENTWORTH and MARY B. WYER.
North Carolina State University, Raleigh, NC. Toward a more inclusive
ecology: Increasing career commitment through curricular intervention.
Current trends in science education encourage the integration of inclusive
material regarding women in the science curriculum. There are few studies
that document the impact of this material and those that exist are primarily
qualitative. This project is an NSF sponsored study by The Women in
Science and Engineering Project in the Womens and Gender Studies Program and the departments of Botany and Zoology at North Carolina State
University. It tests the theory that the inclusion of material regarding women in science increases retention and future career commitment. The study
was undertaken in a large introductory undergraduate ecology course taught
for two consecutive semesters. In 17 laboratory sections (approximately
400 students), 8 sections received supplemental material integrated into the
regular laboratory subject matter and 9 sections served as controls. Material
covered contributions of women overlooked in their course text along with
methodological, cultural, and social biases in science. Results after one
semester indicate a significant increase in commitment to a career in science for both men and women in treatment sections.
ROSENTHAL, DAVID M.,1,* FULCO LUDWIG,1 VOLKER STILLER2
and LISA A. DONOVAN.1 1 University of Georgia, Athens, GA; 2 University of Utah, Salt Lake City, UT. Differences in plant and soil water
relations and nutrient status on contrasting sides of a cold desert ecotone.
In arid ecosystems, soil texture is an important determinant of plant available soil moisture. There is a rapid shift from active sandy soils to stabilized
better developed soils along the ecotone between desert sand dunes and
adjacent off dune habitats. In general, soil clay and nutrient content increase
across this boundary. Plants on fine textured soils should exhibit more
negative water potentials and increased resistance to cavitation than plants
on coarser soils. At Little Sahara Sand Dunes, Utah, we studied plant available water and plant and soil nutrient content for three species of differing
life forms that occur along a narrow sand dune/desert ecotone. For all
species measured, seasonal plant predawn (Cpd) and midday (Cmd) water
potential were significantly higher on dunes than off dunes. Plants growing
on active sand dunes maintained relatively high Cpd (. -1.5 MPa)
throughout the summer season whereas plants growing off dunes were very
water stressed from July on (Cpd , -1.5 MPa). Hardly any water was
available for plants in shallow soils, but soil water potential increased with
depth. Water was available for plants at 50 cm depth and below on dunes,
but only below 75 cm depth off dunes. As predicted soil clay content was
higher at off dune sites. Total soil N concentration was more than four
times lower on dunes than off dunes. Overall soil P was similar across the
ecotone. However, soil P increased with depth on the dunes while the trend
was reversed off dunes. Plant N:P ratios were lower on the dunes. These
findings suggest plants may not root at greater depths on the dunes. We
tested the prediction that on dune plants have lower cavitation resistance
by measuring percent loss of xylem conductivity as a function of xylem
tension. Differences in cavitation resistance between congeners restricted

to either side of the ecotone support the observed differences in plant water
status for species that span the ecotone.
ROSENTHAL, SADIE K.,1,* DAVID M. LODGE1 and MARK W. KERSHNER.2 1 University of Notre Dame, Notre Dame, IN; 2 Kent State University, Kent, OH. Community effects of invasive crayfish: Evidence
from long-term, whole-lake surveys.
Previous laboratory studies and in-lake caging experiments suggest that
invasive rusty crayfish (Orconectes rusticus) reduce macrophyte abundance
and change species composition in north temperate lakes. Using long-term,
whole-lake studies, we test whether these effects are manifest in nature. In
1987 and 2002, we censussed crayfish and macrophytes in eleven lakes in
northern Wisconsin. Between 1987 and 2002, three of six lakes shifted
from low to high crayfish abundances and two of five lakes shifted from
high to low. The remaining six lakes changed little. Both lakewide macrophyte abundance and species richness per transect (of 43 total species
found) decreased as male rusties per trap increased (linear regression:
r250.418, p50.032 and r250.700, p50.001). These relationships were
driven by habitat patches with sand substrate where, as opposed to soft
organic sediments, we expected crayfish to move easily and have a greater
impact on plants. To examine whether macrophyte species composition
differed between high and low rusty lakes and whether composition
changed over time, we compared Detrended Correspondence Analysis results from 1987 with those from 2002. Analyses were based on the frequency of occurrence for the 18 most common macrophyte species over
all eleven lakes. We found that no axis delineated high and low crayfish
lakes in 2002, and only one newly invaded lake moved substantially in
ordination space between 1987 and 2002. We are currently examining two
possible non-exclusive explanations for this: (1) because crayfish are relatively nonselective consumers, they do not alter the occurrence of common
species, and/or (2) crayfish are selective towards particular species, but in
lakes where these species do not occur, composition remains unchanged.
In summary, these surveys suggest that rusty crayfish negatively affect
macrophyte abundance and diversity but that changes in species composition may be lake specific.
ROSS, MICHAEL S.* and PABLO L. RUIZ. Florida International University, Miami, Florida. Reduced freeze incidence associated with global
warming may promote mangrove encroachment in southeastern coastal marshes.
Coastal vegetation patterns in the southeastern US have been altered
through a combination of local and global forces. In South Florida, recent
expansion of mangrove vegetation into former brackish marsh has been
attributed to both sea level rise and local drainage activities. In this presentation, we document a second global mechanism by which this trend
may be exacerbated, i.e., through an anticipated reduction in the frequency
and severity of winter freeze events. We base this projection on vegetation
dynamics in a mangrove-dominated landscape that experienced two patchy
freezes within the seven-year period of study (1995-2002). Tall coastal
forests were buffered from a January 1966 freeze, but it proved to be a
potent force in the dwarf forests immediately inland. Damage differed by
species and was concentrated among individuals that formed or emerged
above the low forest canopy. Mortality or dieback of these dominant stems
had negligible effect on mangrove recruitment, but provided a window of
opportunity for herbaceous plants and mangrove advance regeneration. Five
years later, with species composition altered but biomass and canopy structure approaching the pre-freeze condition, a less intense cold temperature
episode re-opened the mangrove canopy. The landscape signature and ecological effects of the 2001 freeze, i.e., canopy thinning and expansion of
woody and herbaceous understory elements, were remarkably similar to
those of its predecessor. Although our observations pertain to a coastal
basin in which mangrove encroachment had already eliminated most of the
former graminoid component, they do suggest how repeated freezes in
mixed swamp communities may tip the competitive balance against sensitive woody plants. As climatic warming proceeds and freezes become
less pervasive along southeastern coastlines, we can expect mangroves to
become a more important element in undeveloped wetlands.

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287

ROSSI, ANTHONY M.,1,* PETER STILING,2 DANIEL C. MOON,2 MELISSA MURRAY1 and KELLY HUGHES.1 1 University of North Florida,
Jacksonville, Florida, U.S.A.; 2 University of South Florida, Tampa, Florida, U.S.A. Effects of gall size on host choice and parasitoid community
structure in Asphondylia galls.
The gall midge, Asphondylia borrichiae, attacks the stem tips of its host
plants, especially sea oxeye daisy (Borrichia frutescens). Female midges
oviposit near the apical meristem of the shoots, which produces a spherical
gall. Each larva develops individually in a chamber for up to three months.
Adult lifespan is short (1-4 days) and Asphondylia spend most (. 95%)
of their lives embedded within the tissue of the host plant. Immature Asphondylia are attacked by four species of parasitic wasps (parasitoids),
which vary greatly in size. Previous studies have suggested that parasitoid
community composition is determined primarily by ovipositor length. For
instance, the largest parasitoid, Torymus umbilicatus, dominates the parasitoid community in large Asphondylia galls because its long ovipositor
enables it to reach larval chambers when gall size is greatest and, in effect,
parasitize midges that are unreachable by the smaller species. In the current
study, we used yellow sticky traps, artificial galls and a bagging experiment
to determine the relative abundance of the four species of parasitoids within
patches of sea oxeye daisy and the susceptibility of Asphondylia galls to
parasitism. Several significant patterns were detected. For instance, galls
on short stems were 19% larger and 24% more abundant than those on tall
stems. Moreover, galls on short stems had a significantly lower rate of
parasitism (30%) compared to galls on tall stems (60%). Captures of parasitoids using artificial galls coated with Tangletrapy were significantly
higher on large galls attached to medium height stems, compared to either
small or medium galls from short or tall stems. The largest parasitoid, T.
umbilicatus, was the most common parasitoid collected from large galls,
while the smallest species, Galeopsomyia haemon, was the most abundant
parasitoid on small galls. Our results suggest that, while parasitoid community composition within host galls may be the result of differences in
ovipositor length among the parasitoids, interspecific preferences of parasitoids for different size classes of galls may also be responsible for parasitoid community structure.
ROSSON, JAMES F.* USDA Forest Service, Southern Research Station,
Knoxville, TN, USA. Temporal trends in tree species richness on the
lower Mississippi River Alluvial Plain, USA.
The conversion of bottomland hardwood stands to an agriculture land use
through much of the twentieth century has created interest in the sustainability and restoration of this forest resource in this important ecosystem.
One key indicator of forest health and sustainability is species richness.
Using U.S.D.A. Forest Inventory and Analysis data, tree species richness
was tracked on 1,200 sample plots measured in the 1970s and remeasured
in the 1990s on the Alluvial Plain of Arkansas, Louisiana, Mississippi, and
Tennessee. During this interval, mean tree species richness per plot significantly increased from 4.4 (S.E.M. 5 0.09) to 5.4 (S.E.M. 5 0.09), (n 5
1,200). Stands dominated by Quercus spp. showed a significant increase in
richness (6.03 to 6.37, n 5 310) while stands dominated by Taxodium
distichum showed no significant change in richness (4.53 to 4.50, n 5 118).
Although loss of forest land in this ecosystem has a negative impact on
species richness, preliminary findings indicate that complex interactions of
succession and degrees of disturbance in the remaining stands on the Alluvial Plain influence overall species richness.
ROSSON, JESSICA L.* and LINDA BUI.* Louisiana State University,
Baton Rouge, LA. Abiotic factors affecting Solenopsis invicta and Brachymyrmex sp. populations.
Ant populations may vary and often depend on factors such as temperature,
humidity, soil characteristics, and water availability. Many of these abiotic
factors influence the abundance and diversity of ants in habitats and landscapes. Our study focused on three weather factors, three soil characteristics, two water related factors, and three habitat characteristics. One hundred meter transects with food vials containing approximately 4g of vienna
sausage and 4 ml of honey were used to capture Solenopsis invicta and
Brachymyrmex sp. and monitor for ant abundance. In addition to the food
traps, hand collection and observation were used to document Brachymyr-

288

Abstracts

mex sp. abundance, and mound counts were used to document Solenopsis
invicta abundance. The abiotic factors will be analyzed with a multivariate
factor analysis, cluster analysis, logistic regression, and principle component analysis. These analyses will elucidate which of the factors are most
significant in determining Solenopsis invicta and Brachymyrmex sp. abundance.
ROTH, AGATHA-MARIE,1 PAUL KEDDY,1,* DANIEL CAMPBELL,1
HALLIE DOZIER2 and GLEN MONTZ.1 1 Southeastern Louisiana University, Hammond, LA; 2 Louisiana State University, Baton Rouge, LA.
Are pine savannas strongly or weakly structured by competition? A
two year removal experiment and its implications.
Plant communities can be strongly or weakly structured, that is with large
numbers of non-zero entries or with many zero elements in the community
matrix. Many removal studies show large numbers of non-zero elements,
and species tend to be organized into competitive hierarchies. Here we ask
to what degree competition organizes plant communities in wet longleaf
pine savannas. These savannas have exceptional numbers of plant species
including groups with Madrean, Caribbean and even South American biogeographic affinities. The Poaceae, Cyperaceae and Asteraceae show particularly large diversity, while families such as the Eriocaulaceae, Haemodoraceae, Melastomataceae, and Xyridaceae suggest biogeographic linkages to the Guyana Highlands of South America. The greatest diversity of
carnivorous plants in North America also occurs in pine savannas. We
describe diversity patterns in a Louisiana pine savanna and explore the
degree to which this vegetation is structured by competitive interactions.
Species-area curves were calculated from nested plots up to 200 m2 using
the Arrhenius equation [log Species 5 c + zlog(Area)]. The slopes were
high (overall z 5 0.30) and did not change with time since burn (6, 18 or
30 months) although shrubs increased in prevalence. A removal study was
performed for two years in two different habitat types (wet, dry) to evaluate
the competitive effects of the dominant grass (Andropogon virginicus L.)
upon the rest of the herbaceous flora. Mantel tests found that the removal
of A. virginicus had minimal effects upon the composition of other species
when compared to control plots. Pair-wise tests for removal effects showed
no effects for any single species or functional groups. The lack of a conspicuous competitive effect from an abundant species is unexpected, and
suggests that these communities are weakly rather than strongly interactive.
This could contribute to the high levels of biodiversity by reducing rates
of competitive exclusion.
ROTH, BRIAN M.,1,* STACY LISCHKA,1,2 THOMAS R. HRABIK,2 DAMON KRUEGER,2 NORMAN MERCADO-SILVA1 and JOHN J. MAGNSUON.1 1 Center for Limnology, University of Wisconsin-Madison, Madison, Wisconsin, USA; 2 Department of Biology, University of MinnesotaDuluth, Duluth, Minnesota, USA. Experimental removal of exotic rainbow smelt (Osmerus mordax) from Sparkling Lake, Wisconsin.
As part of a larger National Science Foundation-funded research project,
we are attempting to eradicate the exotic rainbow smelt (Osmerus mordax)
from Sparkling Lake, Vilas County, Wisconsin. Rainbow smelt have caused
the near extirpation of native yellow perch (Perca flavescens) and cisco
(Coregonus artedi) as well as a collapse in natural walleye (Stizostedion
vitreum)reproduction in Sparkling Lake and other lakes in northern Wisconsin. Remediation is an attractive option, but biocides and pathogen introduction are undesirable. As a consequence, we designed an eradication
program based on selective overexploitation of spawning smelt and increased predation on both juvenile and adult populations. We used gillnets
to capture smelt during the last three summers and fyke nets to capture
reproductive adults during the last two springs. In addition, the Wisconsin
Department of Natural Resources has stocked more than 400 adult walleye,
5,500 extended growth fingerlings, and 1.3 million fry in Sparkling Lake
the last three years as an effort to increase the predator population. Fishery
regulations on Sparkling Lake were altered to protect adult walleye. We
used a mass-balance model and fish Bioenergetic simulations to assess the
relative effect of gillnets, fyke nets, and predators on the smelt population,
estimated prior to the eradication effort at over 750,000 fish. Our results
indicate that the current walleye population is capable of consuming only
a small fraction of the smelt population. Gillnets captured more smelt than
predators, but are ineffective compared to fyke nets, which captured nearly

500,000 smelt the last two springs. The long-term effects of increased predation have yet to be determined. However, Bioenergetics simulations reveal that the walleye population must be substantially increased to successfully maintain a low smelt population.
ROUGHGARDEN, JOAN E.* Dept. of Biological Sciences, Stanford University, Stanford, CA, USA. Darwins theory of sexual selection false
and inadequate.
Darwins theory of evolution contains three propositions: species are descended from other species, natural selection causes evolution, and males
and females match distinct templates. The third piece of Darwins theory
is challenged by recent discoveries. Darwin wrote that "males of almost
all animals have stronger passions than females" and that "the female . . .
with the rarest of exceptions is less eager than the male . . . she is coy."
Darwin imagined that male characteristics evolve because females select
mates who are "more attractive . . . vigorous and well-armed," just as "man
can give beauty . . . to his male poultry" through selective breeding. This
theory, called "sexual selection," explains the evolution of exceptional male
ornaments like a peacocks tail, as the result of female choice, but now
appears falsified and inadequate overall. Ten findings across vertebrate species collectively refute sexual selection theory: many body types are not
solely male or female; multiple forms of males and females occur within
the same species; sex roles reverse; males forego opportunities to mate;
female choice is not for attractive genes, but to manage male power; family
size is negotiated between males and females; sexual mimicry has been
claimed but never demonstrated; same-sex sexuality is common; mating
promotes relationships independent of sperm transfer; and ornaments are
attractive in homosexual as well as heterosexual relations. A theory of
social selection to replace sexual selection should include two new principles: transactions of reproductive opportunity, and social-inclusionary
traits. A territory-controlling male blue-gill sunfish purchases marriagebroker services from a feminine male, using access to reproductive opportunity as currency. A penis in female spotted hyenas and same-sex sexuality
in female bonobos are required for inclusion in the groups controlling access to food and safety from predators. A female dies or fails to reproduce
without social-inclusionary traits. New approachs to gender and sexuality
must replace sexual selection theory.
ROULSTON, TAI H.,1,* AMANDA L. BREWSTER2 and STEPHEN A.
SMITH.3 1 Blandy Experimental Farm, University of Virginia, Boyce, VA;
2
Yale University, New Haven, Ct, USA; 3 Sarah Lawrence College, Bronxville, NY, USA. Body size constrains the generality of generalist bee
species.
In animal groups with extreme body size variation among taxa, dietary
differences commonly reduce competition for food. Bee species within a
habitat, however, commonly differ more than 100x in body weight but
consume the same basic food items: pollen and nectar. We envision three
possible outcomes of this potentially competitive interaction: 1) Segregation: bees of different size utilize flowers of different morphology, each
maximizing foraging profitability on a different set of hosts; 2) Convergence: all bee species converge on the most productive hosts, or 3) Bodysize dependent host breadth: host breadth is inversely proportional to
body size because foraging requirements, and thus the potential profitability
of hosts, scales with body size (i.e., all bees use the best hosts but small
bees utilize more hosts because more hosts meet their minimum foraging
requirements). To examine this question we established 3 1-hectare plots
and collected floral visitors to each of the main flowering plants for 1 whole
day. We found that, as predicted by size-dependent host breadth theory,
small bee species utilized significantly more host plants within a plot than
large bees. Larger bees were significantly associated with the most productive hosts (as measured by total nectar sugar per flower) but small bees
often used less productive hosts, which were, presumably still profitable
for their relatively more modest foraging requirements. We hypothesize that
two body-size dependent factors combine to yield this ecological pattern:
large bees, for which flight is relatively more cost efficient over longer
distances, find it more profitable to forage further while visiting only the
most productive hosts, while small bees find it more profitable to minimize
travel by visiting more species of hosts near their nest.

ROY, BITTY A.1,* and CHRISTA P.H. MULDER.2 1 University of Oregon,

Eugene, Oregon, USA; 2 University of Alaska, Fairbanks, Alaska, USA.
Global warming and plant pathogens in natural communities.
Temperature and humidity have direct effects on pathogen generation times,
over-wintering mortality, germination and transmission rates. Different
pathogens have different temperature and moisture optima, but because
more species are limited in their distributions by temperature minima, it
has been predicted that warming should increase both the abundance and
diversity of pathogens. The few studies that have been done to date suggest
that these predictions are correct. Even more interesting, and less well
understood, are the numerous indirect effects of climate on pathogens. We
will discuss current theory and data for three of these: (1) Climate change
will lead to some plant species being physiologically stressed, whereas
others will be performing better. How does the physiological status of host
plants influence resistance to pathogens? (2) Climate change will lead to
changes in host diversity. Does pathogen abundance and diversity respond
in predictable ways to increasing or decreasing host diversity? (3) Climate
change will lead to new combinations of hosts and pathogens interacting
with each other. How will hosts respond to attack by novel species of
pathogens?
ROY, MANOJIT* and MERCEDES PASCUAL. University of Michigan,
Ann Arbor, MI. Email: roym@umich.edu. Representing network heterogeneities in simple models of disease.
The role of population heterogeneities in governing disease dynamics can
be better understood by modeling the population as a network of connections, in which individuals interact through specific contact patterns that
vary from one individual to another. These contact patterns are known to
strongly influence the transmission characteristics of disease. They can be
explicitly represented in an individual-based model of disease that incorporates the full contact structure. Alternatively, the effect of such network
heterogeneity can be parameterized in simpler models of disease at the
population level. We consider here one empirical approach previously proposed for spatial heterogeneity and examine its applicability to general
networks. The population model consists of modified mean-field equations
(MMF), in which the functional forms are same as those expected for
homogeneous mixing but parameters are modified to account for heterogeneities. We compare the dynamics, both transient and long-term, generated by MMF and the individual-based simulations for small-word and
scale-free networks. We show that most network heterogeneities can be
parameterized by this simple approximation for different type of disease
models (SIS, SIR and SIRS).
ROYCE, GITANE L.,1,2,* JOHN A. ARNONE1,2 and HEATHER WEATHERLY.1,2 1 Desert Research Institute, University of Nevada, Reno, NV; 2 Desert Research Institute, University of Nevada, Reno, NV. NSF-IRCEB
interannual climate variability and ecosystem processes in tallgrass
prairie: Controls on root length density.
In the first year of a four-year study, we begin to explain controls on below
ground net primary productivity (BNPP) in an intact tall grass prairie ecosystem at the EcoCELL lysimeter facility. We look at how seasonal root
length density (RLD) relates to above ground biomass production, plant
species diversity, and soil moisture content across a growing season. A
simple line intercept method was used to quantify root length density along
the surface of a minirhizatron tube to a depth of 120 cm for each of 12
plots. Bi-annual above and below ground biomass harvests were conducted.
Above ground biomass was clipped to 10 cm, separated by species, and
dry mass measured for each species. Below ground biomass harvest were
conducted by extracting two 4 cm soil cores to a depth of 140 cm per plot.
Soil moisture content is continuously logged using vertically placed time
domain reflectometry segmented rods. Results indicate that RLD and species diversity relate linearly (R25 0.9997) which may indicate a species
niche effect on RLD. There is a bell shaped curve relationship between
RLD and biomass production. There is no simple relationship between
RLD and moisture content indicating that above ground controls on RLD
may be more important. Whether these relationships change in year two,
when the EcoCELL temperature will be increased by 4oC, will be important
information regarding terrestrial feedbacks to climate variability.

Abstracts

289

ROYO, ALEJANDRO A.1,2,* and WALTER P. CARSON.1 1 University of

Pittsburgh, Pittsburgh, Pennsylvania, USA; 2 USDA Forest Service, Northeastern Research Station, Warren, Pennsylvania, USA. Establishing the
tree regeneration template: The role of the understory layer and herbivory on seedling dynamics.
Competition from hay-scented fern (Dennstaedtia punctilobula) and excessive white-tailed deer (Odocoileus virginianus) browsing severely limit tree
recruitment throughout large portions of northeastern forests. Their combined effect threatens to reduce understory tree species composition to
species that are both shade-tolerant and browse-resistant, thus altering the
regeneration patterns that occur following overstory gap formation. If competition by the fern layer and deer herbivory regulate tree seedling establishment and diversity, we predict that gap creation in the fern canopy will
increase the recruitment, survival, and growth of less shade-tolerant species
in a manner comparable to overstory gap dynamics. Furthermore, we predict deer exclusion will increase recruitment, survival, and growth of
browse-intolerant or preferred tree species. We are testing these predictions
using a factorial field manipulation of deer herbivory using exclosures and
hay-scented fern cover via the creation of six different gap sizes. Currently,
we are monitoring the growth and survival of over 30,000 marked seedlings
at three sites within the Allegheny National Forest, Pennsylvania. Our initial census confirmed that the seedling layer at these sites is dominated
(.94%) by black cherry. However, by the second year fern gaps stimulated
germination of several species including birch, pin cherry, red maple, and
white ash, but not black cherry. Woody species richness increased under
fern gaps relative to controls (P,0.001). Fern gaps significantly increased
survival (P,0.001) and marginally increased growth (P50.059) of black
cherry seedlings. In contrast, deer exclusion did not affect richness, survival, or growth for this small (, 20 cm) seedling size class. The data
suggest that fern and deer impacts do not operate concurrently. Initial control of seedling layer is strongly limited by a dense fern understory and
deer may only exert control on the ensuing larger seedling size classes.
RUBINO, LUCY M.,* SARA L. WEBB and STEVE PASSAFARO. Drew
University, Biology Department, Madison, NJ, 07940. Comparison of
sympatric native and invasive maples: Five years of seedling bank dynamics.
Norway maple (Acer platanoides) has invaded intact deciduous forests of
eastern North America where it suppresses plant species richness and where
it has been targeted by restoration projects. We asked if the abundant seedlings of Norway maple behave like the seedling bank of the native sugar
maple (Acer saccharum), which it resembles in leaf morphology and shade
tolerance. From 1997 to 2002, we followed both maple seedling populations in 40 permanent 4x4 m plots within a New Jersey mixed Norway
maple/sugar maple forest. We recorded annual changes in seedling abundance and, in all but the initial sampling efforts, we distinguished new (first
year) maple seedlings. We found that both maples exhibited a major pulse
of seedling recruitment only during the first 2 year interval (1997-99).
However, neither species produced many new seedlings over the subsequent 3 years. This recruitment failure interfered with a restoration experiment; the removal of mature Norway maple trees from half of the study
area after the first census has not influenced the dynamics of the seedling
bank except during the mast seedling recruitment event just after tree removal. Both maples subsequently thinned out in both tree-removal and
control areas, losing 31% of seedlings between 1999-2002 due to seedling
mortality juxtaposed on poor recruitment. The two species differed, however, in their patterns of mortalitiy across the years (time x species interaction, P,0.0005). Closer scrutiny of survivorship from fall (2001) to
spring (2002) and from spring to fall (2002) revealed heaviest losses for
Norway maple during the winter but heaviest losses for sugar maple during
the summer. These results suggest that the two maples mast in synchrony
but differ in their temporal patterns, and thus in causes, of seedling mortality in this location.
RUBIO, MARIA G.,* JERRY W. DRAGOO and TERRY L. YATES. University of New Mexico, Albuquerque, New Mexico, USA. Microsatellite
DNA variation of Peromyscus maniculatus populations in New Mexico.
The deer mouse (Peromyscus maniculatus) is a common and widely distributed rodent known to transmit the hantavirus, Sin Nombre virus (SNV).

290

Abstracts

Genetic studies have revealed strong subdivisions among rodent populations, even at relatively fine scales. We hypothesize that genetic variation
among neighboring populations may be an effect of extinction and then
recolonization from a surrounding area. Analysis of the resident population
through time may shed some light on the origin of dispersers. Microsatellite
data when analyzed in the context of longitudinal studies will allow us to
investigate the basic genetic profile for the host population with an estimation of how it changes over time. Ultimately, we plan to use these data
to test various models associated with the epidemiology of SNV. We have
established parameters required to amplify 12 Peromyscus microsatellite
loci using DNA extracted from ear punches from P. maniculatus located
on 2 study sites in northwestern New Mexico. The loci we have examined
were obtained from Chirhart et al. (2000 - Mol. Ecol., 9:1669). The 12 loci
are as follows: Pml01 (125-195 bp), Pml02 (215-251 bp), Pml03 (224-273
bp), Pml04 (196-233 bp), Pml05 (191-225), Pml06 (129-197 bp), Pml07
(193-264 bp), Pml08 (204-256 bp), Pml09 (193-264 bp), Pml10 (162-198
bp), Pml11 (213-283 bp), Pml12 (139-187 bp). As of 1 March 2003 we
have analyzed approximately 50 of 98 animals for which we have DNA
extracted. All l2 loci were polymorphic and average 20 alleles per locus.
Expected heterozygosity was 0.92 whereas observed heterozygosity was
0.76. We will present population genetic data on approximately 100 animals
that have been marked and recaptured monthly from 2000 to 2002.
RUDGERS, JENNIFER A.* Indiana University, Bloomington, IN. Mechanisms of plant resistance: Plant architecture alters herbivory on a
coastal shrub.
Plants possess many traits that influence their resistance to herbivores. Traditionally, traits such as plant secondary metabolites and physical deterrents
(thorns, trichomes) have received the most attention in both natural and
agricultural systems, with less consideration given to characteristics of
plant architecture. Plant architecture may have particularly important effects on gall-forming insects, which benefit by causing plants to reallocate
resources. Architectural traits that reduce the impact of herbivores on plant
fitness are expected to experience positive selection; these traits may work
directly against herbivores or indirectly by modifying the effectiveness of
herbivores enemies, such as predators or parasitoids. Ultimately, differential herbivory may contribute to the maintenance of architectural polymorphisms. The perennial shrub, Baccharis pilularis (Asteraceae, coyote
bush), grows as two distinct architectural forms in coastal California, USA:
an upright form (1-4 m tall) and a prostrate form (0.1-0.2 m). Differences
in height, branching architecture, and leaf size persisted in a common garden comprised of multiple clones of the two forms, suggesting these architectural traits are genetically based. In a four-year field experiment, I
showed that plant architecture affected resistance to two gall-forming herbivore species. The upright form supported higher densities of lepidopteran
stem-galls (Gnorimoschema baccharisella), whereas the prostrate form received more bud galls formed by midges (Rhopalomyia californica, Diptera). Furthermore, for the midge, architecture did not alter the effectiveness
of parasitoids, which cause the majority of mortality in the field. Experimental reductions and additions of galls over four years demonstrated that
the prostrate form always had greater reproductive success than the upright
form, regardless of the level of herbivore damage. Thus, although plant
architecture strongly influenced the distribution and abundance of galling
insects, mechanisms other than differential herbivory must explain the
maintenance of the architectural polymorphism in Baccharis.
RUEDAS, MARCELA and TERESA VALVERDE. Lab. Especializado de
Ecologia, Facultad de Ciencias, Universidad Nacional Autonoma de Mexico (UNAM)., Mexico, D.F., Mexico. Distribution and abundance of
three sympatric columnar cacti (Neobuxbaumia sp.) with different degrees of rarity.
The genus Neobuxbaumia includes nine species of columnar cacti endemic
to the semi-arid regions of Mexico. In the Tehuacan-Cuicatlan region, wellknown for its high floristic diversity regarding xerophytic vegetation, this
genus is represented by three species which differ in their degree of rarity:
Neobuxbaumia macrocephala (the rarest), N. tetetzo (intermediate), and N.
mezcalaensis (the most common). We carried out a survey of 80 localities
within the Tehuacan-Cuicatlan region in order to investigate the ecological
factors that may be limiting their distribution and abundance, with the aim

of understanding their differing degree of rarity. At each locality we recorded several biotic and abiotic variables, as well as the density of the
Neobuxbaumia sp. population present. We subjected the data to a Principal
Component Analysis (PCA) and a Canonical Correspondence Analysis
(CCA) to identify the factors that account for the variation in the distribution and abundance of each of the three species. The results show that
the factors that explain the distribution of the most common species (N.
mezcalaensis and N. tetetzo) are mean annual temperature, altitude, rainfall,
and soil properties such as texture and organic matter content. N. mezcalaensis reaches population densities of 5000-16000 plants/ha, coexists with
Tecoma stans, Caesalpinia sp. and Aeschynomene sp., and is associated to
localities with relatively abundant rainfall. N. tetetzo shows population densities of 8000-14000 plants/ha, coexists with Cercidium praecox, Leucaena
sp. and Acacia cocliacantha, and is associated to high lalitudes and high
phosphorous content in the soil. The rarest species N. macrocephala shows
low densities (500-1200 plants/ha) and is associated to localities with high
soil calcium content; it coexists with Calliandra sp. and Aeschynomene sp.
Its distribution is limited to localities with specific values of the abiotic
variables recorded, conferring it a high habitat specificity. This accounts
for its higher degree of rarity.
RUIZ-BERNARD, IVELISSE1 and ARIEL LUGO.2 1 University Of Puerto
Rico, iveruiz@yahoo.com, San Juan, Puerto Rico, USA; 2 Int. Inst. of Tropical Forestry / USDA Forest Service, San Juan, Puerto Rico, USA. Restoration of biodiversity in degraded agricultural lands of the Luquillo
Mountains, Puerto Rico.
Forest regeneration on degraded abandoned agricultural lands is an ecological challenge because it requires establishing species-rich communities
that are self-maintaining over a long time. As most efforts of reforestation
in the tropics are recent, there is little evidence of tropical forest restoration
on degraded sites. We document the experience of a site planted about 60
years ago with 13 tree species on degraded abandoned agricultural land at
the Luquillo Experimental Forest. Plantings included native, alien, and native endemic tree species. We sampled 116 plots covering 9.4 ha in 1959
and in 1992 to assess the development of the planted forest. In 1959, we
found 38 tree species and by 1992 the species count increased to 81 tree
species including 86.6% native, 11.2 alien, and 2.2 native endemics to
Puerto Rico. Although six of the top twenty species were in the original
planting and two continue to dominate the site (Tabebuia heterophylla, a
native species, and Callophylum antillanum, an alien species), their presence has not hindered the natural enrichment of the site. The new restored
forest contains an array of tree species similar in number, but different in
composition, from native stands on nearby sites. The stand appears to be
regenerating normally and exhibits functional characteristics that are similar to those of mature native forests. This case study provides encouraging
promise for tropical forest restoration in degraded agricultural sites elsewhere in the tropics.
RUIZ-GUTIERREZ, VIVIANA,2 SCOTT R. LOARIE,1,* CAGAN H.
SEKERCIOGLU,1 GRETCHEN DAILY1 and PAUL R. EHRLICH.1 1 Stanford University, Stanford, CA, USA; 2 University of Costa Rica, Heredia,
CR. Effects of habitat modification on the understory birds of southern
Costa Rican countryside.
We sampled 10,548 birds from 192 species with mist nets from the Las
Cruces forest, small forest fragments, and coffee plantations around Las
Cruces Biological Station in southern Costa Rica. Our objective was to
understand the effects of forest fragmentation and land conversion on the
diversity and composition of the local understory avifauna. We sampled
both during dry (February through April) and wet (June through September) seasons in three different years between 1999 and 2003. We had 6929
captures in coffee plantations, 1935 captures in the Las Cruces forest, and
1584 captures in 3-5 ha forest fragments. The composition of the avifauna
was significantly different between Las Cruces forest, fragments and coffee
plantations. Understory insectivores were significantly more abundant and
speciose in the Las Cruces forest, while granivores, omnivores, and northern migrants were significantly more common and speciose in small fragments and coffee plantations. Forest-dependent species were mostly absent
from coffee plantations and were significantly under-represented in small
fragments. The hotter, drier, and more volatile microclimatic conditions in

small fragments, and especially in coffee plantations, were likely important

factors in the absence of numerous forest bird species from these sites.
Presence of a bird species in the agricultural matrix surrounding forest
patches was the best determinant of its occurrence in the small forest fragments. Poor dispersal ability between forest patches was the most likely
cause of the decline of many forest species, particularly in the superfamily
Furnarioidea, from small fragments. Better integration of human-dominated
ecosystems and small forest fragments into tropical conservation strategies
can significantly increase the capacity of sub-optimal habitats to support
native bird species.
RUMBAITIS-DEL RIO, CRISTINA M.1,2,* and CAROL A. WESSMAN.1,2
Cooperative Institute for Research in Environmental Sciences, Campus
Box 216, Boulder, CO, USA; 2 Department of Environmental, Population,
and Organismic Biology, University of Colorado, Boulder, CO, USA. Patterns of recovery of a wind disturbed and salvage-logged subalpine
forest.
1

This three-year study compares the initial rates of recovery of logged and
unlogged blowdown areas in a subalpine forest. The study site sustained a
catastrophic blowdown in 1997; portions of the blowndown forest were
salvage-logged in 1999. Rates of nitrogen cycling are significantly elevated
in blowdown areas relative to control areas of intact forest. Soil erosion
was significant following salvage-logging activities; consequently soils in
salvage-logged areas contain a shallower organic horizon as well as less
organic and inorganic nitrogen than unlogged blowdown areas. Summertime soil temperatures are on average 10 8C higher in salvage-logged areas
than in unlogged areas. Vegetation diversity and cover are greatest in blowdown areas and least in salvage-logged areas. Tree seedling in both salvagelogged areas and blowdown areas show evidence of a growth release, although seedling density is significantly reduced in salvage-logged areas.
The results of this study highlight the importance of organic matter in
maintaining biotic control over ecosystem recovery of nutrient cycling in
a wind-disturbed forest. Furthermore, this study shows that subalpine forests are relatively resilient to wind disturbance, despite the lack of resprouting in wind-damaged trees, which is common in wind-disturbed temperate and tropical forests. Finally, salvage- logging appears to have delayed ecosystem recovery of this wind-disturbed forest by removing tree
seedlings and soil organic matter, and by altering microclimate conditions.
RUOFF, RACHEL H.,1,* J D. LEWIS,1 ANDY MARTIN,1 GREG TURNER,1 KEVIN L. GRIFFIN2 and DAVID T. TISSUE.3 1 Calder Center, Fordham University, Armonk, NY; 2 Lamont-Doherty Earth Observatory, Columbia University, Palisades, NY; 3 Texas Tech University, Lubbock, TX.
Interactive effects of plant age and nitrogen supply on reproductive
allocation in Xanthium strumarium.
Reproductive output in annual plants generally increases with increasing
plant size. However, changes in resource availability and the length of the
vegetative growth phase may significantly alter reproductive output through
effects on both the absolute and relative allocation of biomass to reproductive structures. In this study, we examined the interactive effects of plant
age and nitrogen supply on allocation to reproduction in the determinate,
short-day, annual species Xanthium strumarium. Four cohorts of plants
were planted at 14-day intervals in a greenhouse. Five plants in each cohort
were given one of three nitrogen treatments (0.5, 1.5 or 5.0 mM NH4NO3).
Supplemental lighting was used to maintain an 18 h photoperiod during
vegetative growth. All plants were simultaneously induced to flower by
switching the photoperiod to 12 h. All plants were harvested at senescence.
Reproductive and total plant mass increased 270% and 334%, respectively,
with increasing nitrogen supply, and increased 200% and 166%, respectively from the youngest to the oldest cohort. However, total number of
fruit produced was highest in the second-oldest cohort, while mean mass
per fruit was highest in the oldest cohort. Both total number of fruit and
mean mass per fruit significantly increased with increasing nitrogen supply.
Plant age and nitrogen supply did not significantly affect the allometric
relationship between reproductive and total plant biomass. In contrast, the
allometric relationship between fruit number and plant size varied with both
plant age and nitrogen supply. These results suggest that the timing of
flowering affects reproductive output in Xanthium strumarium through effects on the mean mass per fruit and on the absolute allocation of biomass

Abstracts

291

to reproduction, but not on the relative allocation of biomass to reproduction. Additionally, these results suggest that nitrogen supply has a greater
effect on total number of fruit compared to mass per fruit in Xanthium
strumarium.
RUSS, ANDREW L.,1,3,* CHARLES M. BACHMANN,2 ROBERT A. FUSINA2 and JOSEPH BURKE.3 1 USDA Agricultural Research Service, Hydrology and Remote Sensing Laboratory, Beltsville, MD; 2 Naval Research
Laboratory, Remote Sensing Division, Code 7212, Washington, D.C.;
3
University of Maryland, Department of Geography, College Park, MD.
Assessing the spatial distribution of Phragmites australis on a MidAtlantic barrier island.
Phragmites australis (Common Reed) has become a concern to ecologists
due to its increasing expansion into wetland areas displacing grass species
such as Spartina patens and Spartina alterniflora. Phragmites was introduced to Hog Island, Virginia, a Mid-Atlantic coastal barrier island, in the
early 1990s. Hog Island is a part of the Nature Conservancys Virginia
Coast Reserve, a National Science Foundation Long Term Ecological Research Site (LTER). It is largely free from anthropogenic disturbance and
as such it offers a unique opportunity to study environmental factors influencing the spread of an invasive species. The spatial distribution of Phragmites on Hog Island was determined through the use of differential global
positioning system (DGPS) surveys, classificaton of PROBE2 hyperspectral
imagery, and examination of color infrared (CIR) aerial photography and
LIDAR data. A total of 102 Phragmites patches were mapped with DGPS,
encompassing an area of 61,740 m2. A Mixture Tuned Matched Filter
(MTMF) classification of the PROBE2 imagery identified pixels within 79
of the 102 surveyed patches. The 23 patches not identified by MTMF were
typically small or sparse, encompassing a total area of 3,080 m2. The LIDAR data suggest a greater than 500% increase in Phragmites coverage
on a portion of the island over the last 5 years. The influence of overwash
disturbance, edaphic factors and interspecific competition on the distribution of Phragmites was examined. Areas of overwash disturbance were
determined to be likely points of establishment. Phragmites appears to have
initially spread along the edges of inland lakes and into interior areas with
harsher edaphic conditions through means of clonal integration. Subsequent
dispersal of propagules lead to the emergence of isolated patches further
inland along the Myrica cerifera thicket/marsh interface.
RUSSELL, ANN E.,1,2,* DAVID A. LAIRD1 and ANTONIO P. MALLARINO.3 1 USDA-ARS National Soil Tilth Laboratory, Ames, IA, USA; 2 Iowa
State University, Department of Natural Resource Ecology and Management, Ames, IA, 50011; 3 Iowa State University, Department of Agronomy,
Ames, IA, USA. Long-term effects of nitrogen fertilization and crop
rotation on soil carbon in corn-belt agroecosystems.
We investigated effects of rate of N fertilization and crop-rotation type on
soil properties in Iowa agroecosystems, in comparison with an undisturbed
prairie. Our two rain-fed experimental sites at Nashua and Kanawha, IA
had been tile-drained and were under conventional tillage. The split-plot
experimental design for the past 23 y (Nashua) and 18 y (Kanawha) included three rotations: 1) continuous corn for grain (CC); 2) corn-soy (CS);
and 3) corn-corn-oats-alfalfa (CCOA). Each rotation had been subjected to
four N levels: 0, 90, 180 and 270 kg ha-1 y-1. We measured the following
soil properties in 2002 at six depth intervals over 0-100 cm: total soil
organic carbon (TSOC); soil inorganic carbon (SIC); total soil N; bulk
density; and pH. We measured other soil properties more intensively in the
0-15-cm depth at Nashua and the native prairie soil in 2001 and 2002. N
fertilization significantly lowered soil pH in all rotations (0-15 cm), but
fertilization had no significant effects on other soil properties. Rotation type
did have significant effects on soil carbon in the 0-15 cm layer. TSOC was
significantly higher (P50.05) under CCOA rotation (4194 g m-2) than CC
and CS rotations (3922 and 3690 g m-2, respectively at Nashua). All crop
rotations had significantly lower TSOC, ranging from 20.4 to 23.4 g kg-1
over all treatments, compared to 64.7 g kg-1 in native prairie. Under CCOA,
the following soil properties were higher compared to the other rotations:
microbial activity (P50.10); microbial biomass C (P50.01); particulate
organic matter C (P50.20); and resistant soil C (P50.05). Our results indicated that higher N fertilization did not result in increased soil carbon
storage under any crop rotation in these systems. Type of crop rotation,

292

Abstracts

however, did influence soil carbon sequestration and biological activity of

the soil, hence its capacity to supply nutrients for plant growth.
RUSSELL, COLIN A.,1,* DAVID L. SMITH,2 LANCE A. WALLER,3
JAMES E. CHILDS4 and LESLIE A. REAL.5 1 University of Cambridge,
Cambridge, England, United Kingdom; 2 University of Maryland, Baltimore, Maryland; 3 Rollins School of Public Health, Emory University, Atlanta, Georgia; 4 Centers for Disease Control and Prevention, Atlanta, Georgia; 5 Emory University, Atlanta, Georgia. A priori prediction of disease
invasion dynamics in a novel environment.
Predictive theories of disease emergence require the development of a
priori models of spatio-temporal host-pathogen dynamics. A major challenge to the development of such models has been the incorporation of
small scale heterogeneity. Initially we blended small-scale date-rich assessments of spatial heterogeneity into global models of disease spread, in
a mechanistic model for the spread of raccoon rabies across the state of
Connecticut. Our model incorporated spatial heterogeneity in local transmission rates depending on presence or absence of rivers and global longdistance transmission independent of local habitat conditions. We applied
the Connecticut model in an a priori attempt to model the spread of raccoon
rabies across New York State. Confronting the model predictions with the
observed data from New York townships provided us with an unusual and
powerful opportunity to test the predictive capabilities of our model as well
as examine the importance of initial conditions. Using the model with multiple points of disease introduction into the state we were able to correctly
predict 80% of the New York data (R2 5 0.80, y 5 0.95x, p , 0.0001).
We were also able to use deviations from model predictions to elucidate
the anisotropic effects of rivers and the interaction of wave direction and
other geographic features.
RUSSELL, ROBIN E.* and ROBERT K. SWIHART. Purdue University,
West Lafayette, IN. Modeling behavioral decisions of voles in complex
landscapes.
Present-day landscapes often consist of a patch work of land-cover types,
including areas heavily modified by human use. Wildlife species differ in
their ability to cross resource poor areas and willingness to venture into
areas that increase their exposure to predators. Transitions from one landcover type to another reflect a behavioral decision process which presumably relies on the animals assessment of risk versus gain in the new landcover type. We observed meadow voles, Microtus pennsylvanicus, in experimental landscapes to estimate their transition probabilities between habitat types.Voles were released within an enclosure containing habitat
patches connected by corridors and tracked by radio-telemetry.Vegetation
within the corridors was modified to represent four different levels of risk.
Transition probabilities from a patch to a particular corridor type were
determined using multi-strata modeling in program MARK. Transitions
from patches to corridors composed of tall vegetation (0.8m) and halfheight vegetation (0.4m) were significantly more common than transitions
to corridors devoid of vegetation or containing mown vegetation
(0.05m).However, bare- ground corridors were utilized more often than
expected and preferred over mown corridors.We previously conducted
computer simulations of animal movements in complex landscapes which
indicated that the movement rule utilized in the model influenced the abundance and distribution of the population. Our field results combined with
our simulations indicate that assumptions of optimal movements in landscape models may lead to inaccurate conclusions regarding the ability of
wildlife populations to persist in complex landscapes.
RUSSO, SABRINA E.,1,* PORTNOY STEPHEN2 and CAROL K. AUGSPURGER.2 1 University of Illinois at Urbana-Champaign, srusso@uiuc.edu, Urbana, IL, USA; 2 University of Illinois at Urbana-Champaign, Urbana, IL, USA. Evaluating recruitment limitation of a neotropical tree using a mechanistic model of vertebrate seed dispersal.
In plant communities recruitment limitation resulting from spatially restricted seed dispersal may promote species coexistence if the inability of
a better competitor to disperse to a site permits poorer competitors to establish there instead. However, descriptions of seed dispersion at landscape
scales are few, limiting quantification of restricted dispersal and its effect

on recruitment. We developed a spatially-explicit, mechanistic model that

predicts the landscape-scale seed-dispersion pattern of a neotropical nutmeg
tree species, Virola calophylla (Myristicaceae), by simulating the stochastic
process of seed dispersal. The model is based on field-collected data on
fruit production and total seed dispersal from source trees, densities of
dispersed and non-dispersed seeds, direct estimates of seed dispersal distances, and observations of the behavior and movement of the trees key
dispersal agent, the spider monkey (Ateles paniscus). No single statistical
distribution could be fit to the models predicted individual seed dispersal
curve, which was best described by a mixture distribution. We evaluated
the extent of dispersal limitation in V. calophylla and found that spatially
aggregated seed dispersal by spider monkeys at their sleeping sites produced a pattern consistent with dispersal limitation. Diurnal seed dispersal
events, which were spatially scattered, combined with the great range of
seed dispersal distances, produced a background of even, low-density seed
deposition in the landscape. Nonetheless, V. calophylla was strongly dispersal-limited at this site, in that few seeds reached potential 1-m2 establishment sites in one fruiting season. Furthermore, the large spatial variation in predicted post-dispersal seed densities has important implications
for species diversity-maintaining mechanisms. Describing the mechanistic
link between disperser activity and the pattern of seed deposition is fundamental to evaluating the role of plant-animal interactions in the maintenance of species diversity in the tropics.
RUTHIG, GREGORY R.* University of Virginia, grr4b@virginia.edu,
Charlottesville, VA. The influence of infectious disease on amphibian
egg-laying behavior.
Infectious diseases are poorly understood as selective forces on life history.
Since the threat of disease is often heterogeneous in the environment, one
would expect variation in traits that minimize the impact of diseases but
are costly in their absence. This expectation is tested in the host-pathogen
system of the southern leopard frog, Rana utricularia and the aquatic,
fungus-like chromista, Saprolegnia ferax. In the winter, R. utricularia mating pairs lay large communal egg masses, while in the fall they lay their
eggs in individual masses. Saprolegnia is a pathogen of amphibian eggs
whose transmission is facilitated by warmer water. Separating egg masses
may reduce Saprolegnia transmission in warmer months. This study recorded the threat of infection and the type of egg-laying behavior of the
frogs. This is the first empirical study to examine the interaction between
a pathogen and the environment and its effect on host life history. Understanding this interaction is particularly important in the face of global environmental change and worldwide amphibian extinctions.
RYAN, MICHAEL G.1,2,* 1 mgryan@fs.fed.us, USDA/FS Rocky Mountain
Research Station, Fort Collins, CO 80526, USA; 2 Graduate Degree Program in Ecology, Colorado State University, Fort Collins, CO 80523, USA.
Rate-based respiratory measurements without a system context are impeding a mechanistic understanding of the controls on respiratory fluxes.
Ecosystems scientists and ecophysiologists measure autotrophic and soil
surface CO2 efflux (soil respiration) to understand how these fluxes vary
among plant parts and with environmental variation. Such knowledge is
essential to develop scaling models and to predict how fluxes and ecosystem carbon storage will change with changes in species, biomass, nutrition,
climate, atmospheric decomposition and composition. Early measurements
of respiration rates assumed that these rates would remain constant (once
the sources of variability had been identified) and that inferences at the
system level could be drawn or inferred from measurements of rates. The
recent intense focus on the carbon cycle and its regulation has promoted a
renewed interest in respiration. However, the fundamental question motivating these measurements remains the same: will environmental changes
alter a plants carbon balance (its ratio of respiration to photosynthesis, R:
P) or the carbon balance of the ecosystem? My objective with this review
is to convince ecophysiologists to place respiration measurements into the
context of a larger plant or ecosystem carbon balance, because short-term
measurements of respiration rates cannot be used to infer ecosystem behavior. Autotrophic respiration rates are highly variable and plastic they
vary with temperature, metabolic activity, substrate availability, phenology,
and tissue type. Soil surface CO2 efflux is strongly linked to plant processes

through root and mycorrhizal respiration, and because much of the heterotrophic respiration consumes recently produced detritus. Both autotrophic
and soil respiration rates acclimate if conditions change. Finally, plants
can shift carbon allocation in response to shifts in resource availability.
This plasticity and structural changes make it difficult or impossible to infer
how R:P or ecosystem C storage might change from short-term measurements of autotrophic or soil respiration rates. I provide examples of rates
placed in the context of larger carbon balances, and some strategies for
assembling such budgets.
RYEL, RONALD J.,1,* A JOSHUA LEFFLER,1 MICHAEL S. PEEK,1
CAROLYN Y. IVANS2 and MARTYN M. CALDWELL.1 1 Department of
Forest, Range, & Wildlife Sciences and the Ecology Center, Utah State
University, 5230 Old Main Hill, Logan, UT, USA; 2 Department of Biological Sciences, Eastern Kentucky University, Richmond, KY, USA. Water conservation in Artemisia tridentata through redistribution of precipitation.
Water conservation is important for plants that maintain physiologically
active foliage during prolonged periods of drought. A variety of mechanisms for water conservation exist including stomatal regulation, foliage
loss, above- and below-ground allocation patterns, size of xylem vessels
and leaf pubescence. Using the results of a field and simulation study with
Artemisia tridentata in the Great Basin, USA, we propose an additional
mechanism of water conservation that can be used by plants in arid and
semi-arid environments following pulses of water availability. Precipitation
redistributed more uniformly in the soil column by roots (hydraulic redistribution) slows the rate at which this water can subsequently be taken up
by plants, thus prolonging water availability during periods of drought. By
spreading out water more uniformly in the soil column at lower water
potentials following precipitation events, water use is reduced due to lower
soil conductivity. The greater remaining soil water and more uniform distribution result in higher plant predawn water potentials and transpiration
rates later in the drought period. Simulation results indicate that plants can
benefit during drought periods from water storage following both summer
rain events (small summer pulses) and overwinter recharge (large spring
pulse). This mechanism of water conservation may aid in sustaining active
foliage, maintaining root-soil hydraulic connectivity, and increasing survival probability of plants which remain physiologically active during periods of drought.
RYKIEL, EDWARD J. Washington State University Tri-Cities, Richland,
Washington. Biocomplexity: the limits to ecological understanding.
Complexity lies at the heart of ecology. Yet, for the most part, ecologists
try to simplify whenever possible on the grounds that the complexity of
ecosystems exceeds our ability to understand interactions and system behaviors. My hypothesis is that ecosystems are in fact too complex for
humans to understand or control. Biocomplexity research is laudable and
will undoubtedly make incremental increases in our scientific understanding. But in the end, we may not have any greater degree of control over
the behavior of ecosystems and especially the biosphere. Virtually every
academic discipline contains some notion of complexity, which is usually
undefined in any operational sense. Therefore, it is generally impossible to
determine how system properties and behaviors differ with respect to complexity. Currently there are no definitions of complexity that are useful for
understanding the behavior of ecological systems. Ecological complexity
is most often associated with inability to predict system behavior. Complexity needs to be associated with some measurable property of the ecosystem if this concept is to express anything more than frustration. In addition, given some quantifiable level of complexity, some aspect of ecosystem behavior should be predictable. The fundamental reasons for studying biocomplexity are predictability, controllability, and recoverability. Can
humans predict and control the behavior of an ecosystem and in particular
the global earth system so as to restore, produce and sustain any desired
ecosystem behavior? Unless there are observable and measurable ecosystem properties associated with ecological complexity, the concept will remain largely intuitive and largely useless. I introduce the maximum complexity hypothesis which suggests that ecosystems approach a level of complexity that is sustainable within a particular environmental context. Complexity increases to this limit from below and decreases to this limit from
above.

Abstracts

293

SAAH, DAVID S.* and TRACY L. BENNING. University of California,

Berkeley, Berkeley, CA. Partitioning of heterogeneity in an object-oriented riparian boundary classification system for a South African savanna.
Riparian boundaries have structural characteristics that are distinguishable
from upland vegetation. These riparian elements within a savanna matrix
located in Kruger National Park, South Africa are the result of several
agents of spatial heterogeneity including fire, flood events and geomorphic
gradients. We developed a riparian classification system for this study region by integrating information on fire frequency, flood plain hydrology
and terrain unit data with ecologically relevant indices derived from a multispectral tassel cap analysis of Landsat ETM data. Results from this work
indicate that riparian boundaries account for 8% of the total classified area
and were found to be primarily located within the 500 year floodplain as
well as in areas with low fire frequency. The tassel cap greenness and
wetness indices were found on average to be 19% and 24% greater in
riparian boundaries compared to upland vegetation types respectively. Distinct structural differences along lateral gradients were also found on main
stem rivers. Main stem riparian boundaries were an average of 500 meters
wider on basalt geology compared to those found on granite substrate. In
addition, basalt vegetation tended to be sparser with an average canopy
cover of 13% while the typical canopy cover from the granites averaged
20%. An initial accuracy assessment of this classification was conducted
using a confusion matrix. Results from this assessment indicate an overall
accuracy of 93%.
SABO, JOHN L.,1,* WARREN P. PORTER2 and JIM REICHMAN.3 1 Arizona State University, Tempe, AZ; 2 University of Wisconsin-Madison,
Madison, WI; 3 National Center for Ecological Analysis and Synthesis,
Santa Barbara, CA. Thermal context dependency: The influence of climate and energetics on spatially explicit patterns of food web structure
and dynamics.
One of the greatest challenges to a general theory of food web dynamics
is understanding how food web structure varies with ecological contexts.
Contexts are diverse and include biotic factors such as the abundance or
recruitment of a particular species, as well as abiotic factors like flooding,
wave action or climate. Many of the classic experiments on intertidal communities as well as more recent studies of food webs in both aquatic and
terrestrial habitats suggest that context dependency is prevalent and a potentially useful framework for understanding and characterizing variability
in food web dynamics at meaningful spatial and temporal scales. One important context in terrestrial landscapes is the thermal environment. Local
climate is patchy at a variety of spatial scales across the landscape. The
goal of this talk is to explore how variability in the thermal environment
may determine spatial variation in food web structure (presence/absence of
herbivores and carnivores) and food web dynamics (i.e. interaction strength
between predators and prey) in terrestrial ecosystems. To do this, we first
apply newly developed biophysical models to predict spatially explicit patterns of individual activity and energetics for a suite of hypothetical predator-prey pairs varying in body size and mode of temperature regulation
(ecto- and endotherms). We then use predicted values of metabolic rates,
"discretionary energy" (the energy available for growth and reproduction),
and activity time from biophysical models as parameters in models of population growth for these predator and prey species. In this way we provide
a framework for linking landscape variables (slope and aspect) directly to
per capita rates of energy consumption and encounter. These per capita
rates in turn determine the relative growth rates and standing stocks of
predator and prey populations. We then illustrate how our model predictions could be used as testable hypotheses in field experiments designed to
evaluate how the strength of predator prey interactions and the prevalence
of trophic cascades vary across real landscapes.
SACCHI, CHRISTOPHER F. Department of Biology, Kutztown University
of PA, Kutztown, PA. Small mammal predation of flowering dogwood
seeds: the influence of seed density and leaf litter.
A putative benefit of seed dispersal is decreased seed predation resulting
from a reduced likelihood of detection of seeds by seed predators at reduced densities away from the parent plant. I studied, experimentally, the

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influence of seed density on predation of flowering dogwood, Cornus florida, seeds by small mammals, squirrels, or birds over a one month period
during the fall. Additionally, I studied the influence of leaf litter on seed
predation. Use of bird and squirrel exclosures indicated that seed predation
by small mammals, likely the white-footed mouse, Peromyscus leucopus,
was high at all seed densities. Added leaf cover provided protection to
seeds suggested by lower predation rates in litter addition treatments. Reduction of seed density did not lead to large decreases in predation rates
but increased leaf litter did permit an increased likelihood of seeds escaping
predation. Increased seed survival through escape from predation was not
an obvious benefit of seed dispersal for flowering dogwood seeds.
SADLER, KIM C.1 and CINDI SMITH-WALTERS.2 1 Middle Tennessee
State University, ksadler@mtsu.edu, Murfreesboro, TN; 2 Middle Tennesse
State University, csmithwa@mtsu.edu, Murfreesboro, TN. Raising ecological awareness in preservice educators and elementary students
through a cedar glade curriculum study.
The role of college biology faculty in preparing future teachers requires
they develop not only knowledge but also a basic understanding of the
nature of science. The cedar glade curriculum model was designed to raise
ecological awareness in preservice educators and elementary students by
teaching the processes of science through field study partnerships. Historically used as junkyards by local people, cedar glades serve as habitat to
over 350 plant species, with more than 25 species endemic to the limestone
glades. Viewing the cedar glades as a unique ecosystem composed of different plant populations, students select an endemic plant to research. Developmental plant processes are charted during the study; predictions and
inferences are made about the plant and other plants in that zone. Within
this contextual framework, the cedar glade study was designed to teach the
processes of plant ecology in two tiers, by biology faculty teaching preservice educators and preservice educators in turn teaching elementary students through partnerships. Interdisciplinary in scope, the curriculum design integrates math, language, and visual arts. The instructional model
involves concept formation (list, group, categorize), interpretation of data
(identify relationships, make inferences), and application of principles (predict, explain, hypothesize, support, verify). The integration of these skills
develops both the preservice and elementary students ability to use and
develop scientific ways of thinking, and in turn enables them to apply what
they know to their world beyond the classroom.
SAH, JAY P., MICHAEL S. ROSS, DAVID L. REED, PABLO L. RUIZ
and DAVID T. JONES. Florida International University, Miami, FL. Vegetation structure and composition along hydrologic gradients in Everglades Tree Islands.
Tree islands, an integral component of the landscape in the Shark River
Slough of the Everglades, have continuously been affected by managementoriented changes in water flow pattern. Our study determined the variation
in plant community structure within Shark Slough tree islands, and examined its associations with recent hydrologic conditions. Three tree islands, Black Hammock, Gumbo Limbo, and Satinleaf, were selected for
this study. Forests on each island were classified into hardwood hammock,
bayhead, and bayhead swamp. Hydrologic gradients in the islands were
calculated from elevation data collected at 5-10 m intervals along four
transects, one along the long axis of the islands, and three others perpendicular to it. Vegetation was sampled along the transects as well as within
three permanent plots of 25 x 25 m, 20 x 20 m, and 15 x 15 m established
in the hardwood hammock, bayhead, and bayhead swamp, respectively.
Tree, sapling, shrub and herb strata were sampled separately in a nested
design. The surface elevation decreased from hardwood hammock through
bayhead to bayhead swamp, suggesting deeper water and longer hydroperiod in reverse order of the above. On each tree island, the three forest types
differed significantly in species composition. Plant species diversity was
highest in the wetter types. Species hydrologic niches calculated for each
island using abundance-weighted averaging techniques indicated strong
correlations in species9 behavior among the three islands. Canopy height
exhibited a strong positive association with surface elevation, but understory cover showed an opposite trend. Litterfall data suggested that the
pattern of decreasing canopy height was also associated with a gradient of
decreasing site productivity, presumably as a result of hydrologically re-

lated factors. Differences in vegetation structure and productivity on tree

islands in relation to hydrology may help in establishing monitoring criteria
for the assessment of Everglades restoration efforts.
SALKELD, DAN.* James Cook University, Townsville, Australia. Parasite regulation of a lizard population.
Parasites can regulate host populations by reducing host survival and fecundity. I illustrate the impact of naturally occurring haemogregarines (parasitic protists) upon life history of the eastern water skink (Eulamprus
quoyii), an Australian viviparous lizard. Using mark-recapture data, and the
Anderson-Gordon model (1982), I show that haemogregarines do not affect
survival of their skink hosts. However, fecundity is reduced by parasitic
infection as heavily infected skinks produce smaller litters than skinks with
small parasite loads. Poulin & Vickery (1996) suggest that parasites can
also reduce fecundity by affecting female mate choice. The impact of protist infection upon lizard mating strategy (the extent of polygyny) is examined using evidence from microsatellite markers.
SALO, LUCINDA F.* USGS Snake River Field Station, Boise, ID. Introduced (Prosopis juliflora and Azadirachta indica) and native (Acacia
nilotica var. adansonia) trees spreading in Senegal, West Africa.
Plant invasions from the Old World to the New World are more frequent
than the reverse. Many of the New World plants that have become invasive
in the Old World are woody species and many of these were intentionally
introduced. Determining the age structure of a population is a first step in
understanding whether or not it is regenerating and in predicting whether
or not it may spread and become invasive. This study determined the size
structure, and mapped all individuals, of three woody species at a site at
Foundiougne, Fatick Region, Senegal. These included mesquite (Prosopis
juliflora), introduced from Central America, and neem (Azadirachta
indica), introduced from India. I also examined the only native tree regenerating at this site, Acacia nilotica var. adansonia. These three species were
all represented by large numbers of seedlings and progressively fewer individuals of successively larger size classes. This indicates that these species are regenerating successfully and that introduced species are spreading
from where they were planted. All other native tree species at this site were
represented only by a few mature, heavily used individuals. The Prosopis
and the Acacia species both appear able to regenerate, and to spread, due
to 1) spines that reduce browsing and rooting by livestock, 2) the ability
to resprout after browsing, 2) pods that are eaten and spread by livestock,
and 3) low frequency of fire, due to heavy grazing by livestock.
SALTONSTALL, KRISTIN.* University of Maryland Center for Environmental Science, Horn Point Laboratory, Cambridge, MD, USA. A historical perspective on a cryptic invasion: Phragmites australis.
Over the past century, one of the most conspicuous changes to marshes
along the Atlantic coast is the rapid spread of Phragmites australis. While
native to these communities over much of this range, evidence suggests
that the widespread invasions of this species are due to the introduction of
a EurAsian lineage of Phragmites which has both outcompeted native populations of the species as well as moved into sites where it was not found
historically. Native populations persist in some sites but little is known
about their ecology and whether or not they will be able to compete with
introduced populations. Clear determinations of human influences on this
invasion are elusive, however it is apparent that substrate disturbance, alterations in hydrology, and nutrient inputs facilitate establishment and
spread of the introduced lineage. Today introduced Phragmites is the dominant macrophyte in many intertidal environments and is considered poor
habitat by many due to its high standing biomass and ability to rapidly
accrete sediments thus altering marsh hydrology. Such characteristics may
prove valuable however in the face of sea level rise and recurrent nutrient
inputs due to coastal development.
SAMUELS, IVAN A.* University of Florida, Gainesville, Florida, USA.
Invasion of Chinese Tallow (Sapium sebiferum): Dispersal and recruitment limitation in multiple habitats.
Studies of exotic plant invasions have often focused on spread and impact
of these species, as well as the processes that facilitate invasion. However,

little attempt has been made to understand why particularly invasive plants
do not establish in potentially suitable habitats where dispersal of their
seeds may be high. I compared both dispersal and post-dispersal processes
of Sapium sebiferum (Chinese Tallow) in four habitats, two wet prairie
habitats where the plant is currently invading, and two adjacent forest habitats where it is absent, but where it occurs elsewhere in its non-native
range. I observed foraging birds to quantify the number of seeds removed
and post-foraging movements, and used point counts to estimate the density
of seed dispersing bird species in the non-invaded habitats. In addition, I
experimentally placed seeds in each of the four habitats to compare seed
predation, germination, and seedling survivorship rates. Of 30 species of
birds that visited trees, 53% ingested seeds and 20% carried seeds away
from parent trees. After foraging, these species frequently moved into forest
habitats, where they were commonly detected on point counts. Thus, dispersal limitation is unlikely to explain the absence of S. sebiferum in the
two forest habitats. Differences in seed predation are likewise unable to
explain recruitment limitation in forest sites, as it was significantly lower
than in wet prairie. On-going germination tests and monitoring of seedlings
will identify the stage/s that act as a barrier to establishment in forest
habitats, and the relative importance of these stages on the overall probability of recruitment.
SANDERS, NATHAN J.* Department of Biological Sciences, Humboldt
State University, Arcata, CA. Do we need to worry even more about
invasive ants?
It is abundantly clear that invasive species and global climate change threaten the integrity of many native communities and ecosystems. Recent reviews and research suggest climate change and the spread of invasive species may act in concert and that global change may exacerbate the spread
of invasive species. In this talk, I examine (i) the relationship among climate and ant species richness, (ii) how predicted climate change may affect
both species richness and the success of many invasive species, and (iii)
how predicted climate change will aid the spread of two of the most conspicuous invasive ant species in the US, Solenopsis invicta in the southern
US and Linepithema humile in the western US. Climatic variation is obviously related to variation in ant species richness, but the relationship
changes with scale. Likewise, the effects of predicted climate change on
ant species richness also vary with scale. The distributions of most invasive
species will be affected by changing climate, but probably not in systematic
ways. Finally, preliminary analyses suggest predicted climate change will
enhance the spread, and likely the impact, of both S. invicta and L. humile
on native communities. So, at least for these two species, we should worry
even more.
SANDERS, SUZANNE M.* and JAMES B. MCGRAW. West Virginia
University, Morgantown, WV, USA. Does breeding system contribute to
the rarity of Hydrastis canadensis?
Abundance of Hydrastis canadensis (goldenseal) in the eastern deciduous
forest is declining. This decline may be directly or indirectly linked to the
breeding system of this species. For example, this species reproduces primarily clonally, forming dense patches. These patches are often separated
by great distances, such that obligate outcrossing could limit seed production and/or viability. To test for breeding system type, we implemented
treatments on ramets from three natural source populations. To avoid difficulties associated with field pollination, this study was conducted in a
greenhouse. The location of flowering ramets were marked at all three
source populations in 2001. These were removed in March 2002 prior to
emergence and planted in a standard potting mix. Six treatments were implemented. Our results showed that 1) this species is not apomictic, 2) seed
set occurred via self pollination, both with and without assistance, 3) seed
set occurred via outcrossing with ramets from the same source population
and with ramets from other source populations and, 4) seed set of field
controls at the source populations was greater than that of the greenhouse
treatments. Tetrazolium testing showed that all seeds were viable. These
results suggest that breeding system type, and seed production, are not
factors limiting the abundance and spread of H. canadensis. However, seedlings are only uncommonly observed in natural populations. Seedlings can
be readily distinguished from small vegetative ramets because only cotyledons are produced during the first year of growth. This suggests seed

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295

germination may be a factor limiting abundance and spread of this species.

Changes in disturbance regimes and/or faunal composition of the eastern
deciduous forest may limit seed germination.
SANFORD, ROBERT L.,1,* JAYNE BELNAP,2 SARAH GOPALANI,1
NATHAN J. WOJCIK1 and MAXWELL J. RICHARDSON.1 1 University
of Denver, Denver, CO; 2 USGS - Biological Resources, Moab, UT. Cheatgrass (Bromus tectorum L.) and seasonal patterns of soil phosphorus
availability in arid ecosystems.
Seasonal changes in soil phosphorus (P) availability are not yet known for
many ecosystems. We report seasonal changes in several pools of soil phosphorus, including plant available P from the Mojave Desert and Colorado
Plateau. In addition we show that cheatgrass changes soil P fractions in
unexpected and ecologically significant ways. Monthly soil samples (0-10
cm) from four sites in Canyonlands National Park in southeast Utah, are
analyzed for P with a modified Hedley P fractionation method. Labile P
(plant available) peaks in spring and autumn with significant monthly variation. Surprisingly, HCl extractable P changes as well, with a pattern inverse to that of labile P. Each of these sites has considerable Bromus tectorum L. (cheatgrass) invasion. In a separate greenhouse study with four
contrasting soils (0-10 cm) from the Mojave desert CA and NV, we grew
cheatgrass for 100 days to test for soil P changes. Regardless of the soil
type, cheatgrass causes large (2x) increases in labile P and large (90%)
decreases in both NaOH and HCl extractable P. In addition, root biomass,
root nitrogen (N) and P as well as shoot biomass and shoot N and P vary
widely according to soil type. We propose that cheatgrass roots solubilize
large amounts of soil phosphorus seasonally, from P pools that are considered unavailable to most plant species, and that this solubilized P becomes
available for Broumus tectorum L. uptake. These results are discussed in
the contexts of root exudates, mycorrhizae and root rhizosphere effects on
desert soil P.
SANKARAN, MAHESH1,2 and DAVID AUGUSTINE.3 1 NERC Centre for
Population Biology, Imperial College at Silwood Park, Ascot, Berkshire,
UK; 2 NREL, Colorado State University, Fort Collins, CO, USA; 3 Syracuse
University, Syracuse, NY, USA. Consumer-decomposer interactions in
a semi-arid grazing ecosystem.
Ecosystem-level studies of producer-decomposer interactions have focused
primarily on plant production and soil texture as regulators of decomposer
abundance, but have rarely considered the role of grazers in mediating such
interactions. Here, we conducted replicated exclosure experiments at both
high and low levels of soil fertility to investigate the effects of large, mammalian grazers on decomposer biomass and activity patterns in a semi-arid
grazing ecosystem in Kenya. Within only 2 years of grazer exclusion, microbial biomass increased in fenced grassland across all levels of soil fertility, indicating that grazers regulate decomposer populations in this system
by depressing the quantity and/or quality of plant inputs to soils. However,
across all study sites, microbial biomass was highly correlated with soil
carbon content, suggesting that landscape-scale constraints on soil organic
matter content are even more important than grazing in regulating microbial
abundance. As soil organic carbon levels increased, both microbial biomass
and the fraction of soil carbon accounted for by microbes increased, and
this was associated with an increased efficiency of resource utilization by
microbes and lowered carbon turnover rates in soils. Our results support
previous ecosystem-level studies showing that microbial biomass and
growth are constrained by plant production and soil C availability. In addition, our findings also demonstrate that decomposer abundance can be
influenced by an ecosystems trophic structure, with significant reductions
in biomass occurring as a result of herbivores diverting plant carbon away
from soils.
SANTIAGO, LOUIS S. University of Florida, Department of Botany, PO
Box 118526, Gainesville, FL, USA. Linking plant physiological ecology
to ecosystem science: Effects of life history traits on leaf decomposition.
Plant life history traits such as wood density, photosynthesis and anti-herbivore defenses, were compared with leaf litter decomposition rates (2
years) on a broad selection of plant growth forms including palms, lianas,
canopy trees and pioneer trees in Panamanian lowland wet forest. Leaf

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nitrogen had a strong positive effect on leaf decomposition rate, whereas

anti-herbivore defenses had a strong negative effect on decomposition. Leaf
nitrogen also had a strong positive effect on leaf photosynthetic rate and
anti-herbivore defenses were negatively correlated with photosynthesis.
Wood density appeared to constrain suites of traits. Species with low wood
density exhibited high photosynthetic and decomposition rates. In contrast,
species with low wood density exhibited leaves with lower photosynthetic
rates that were relatively slow to decompose. In general, wood density is
negatively correlated with plant growth rate and therefore reflects variation
along an axis of life history traits. The observation that leaf photosynthesis
and decomposition are driven by the same interdependent suite of characteristics suggests that the axis of plant life history traits can be extended
to decomposition. Therefore, the effects of plant species on decomposition
at the ecosystem scale can be understood in terms of specific allocation
patterns that reflect plant strategies.
SARNELLE, ORLANDO.* Department of Fisheries and Wildlife, Michigan State University, East Lansing, MI, USA. Keystone-predation revisited: Herbivore effects on phytoplankton diversity and grazing-resistance.
Keystone predation models predict that a selective predator can promote
coexistence among competing prey, and so enhance prey diversity, when
the predator selectively kills the competitively-superior prey. This implies
that there is a tradeoff between resistance to predation and competitive
ability at low resource levels among prey species. In this case, keystone
predators promote both prey diversity and dominance by more resistant
prey. It is also possible for a selective predator to promote dominance by
prey species that are less resistant to predation, if there is a tradeoff between
resistance to predation and competitive ability at high resource levels
among prey species. However in this case, keystone predation theory predicts that the predator is not likely to promote prey diversity. I present
experimental evidence demonstrating that Daphnia, a selective grazer, can
act as a keystone predator (i. e., promote phytoplankton diversity) without
shifting the phytoplankton community to dominance by more resistant species. There appear to be multiple strategies for dealing with high predation
pressure among the phytoplankton, such that a selective predator can increase prey diversity by simultaneously promoting dominance by species
that are resistant to predation and species with high maximum growth rates
that are vulnerable to predation.
SAUER, SUZANNE L. and ROGER C. ANDERSON.* Illinois State University, Normal, Illinois, United States. Modifying distance methods to
estimate historical tree density from General Land Survey Records.
Various researchers have used the Government Land Office (GLO) survey
records to determine historic vegetation through the application of the distance methods to witness tree data. We test the suitability of applying modifications of the random pairs method or the quarter method to GLO records
from the Shawnee National Forest Purchase Area in southern Illinois. The
witness tree data was evaluated with criteria based on the distribution of
the witness trees around the section or quarter section points. As a whole,
the data set did not meet all criteria for either method. Data were subdivided
into groups meeting specific criteria. For interior points with two witness
trees, the random pair method with an exclusion angle of 202.93 degrees
provided the best modification of the distance methods to convert witness
tree distances to tree density. A derived correction factor of 0.74 to convert
the mean witness tree distance to the square of the mean area was determined by regressing exclusion angle against correction factor. Wilcoxon
sign-rank tests indicate that there are significant differences in the calculated square root of the mean based on distance method used (quarter method v. random pairs S53665417, p,0.001; random pairs vs. the derived
0.74 correction factor S53893716, p,0.001).
SAUNDERS, COLIN J.,1,* JAMES F. REYNOLDS,1 J P. MEGONIGAL2
and BERT G. DRAKE.2 1 Department of Biology, Duke University, Durham, NC, USA; 2 Smithsonian Environmental Research Center, Edgewater,
MD, USA. Modeling the past 500 years of organic matter accumulation
in a Chesapeake Bay salt marsh.
Evidence from salt marshes along the Atlantic Coast indicates the composition of plant species has changed in the last century, attributed to

changes in land use, sea-level, and salinity. Ecosystem models are needed
to explore the consequences of changes in species composition on ecosystem structure and function. The goal of this study is to determine whether
changes in species composition improve predictions of soil accumulation
in a Chesapeake Bay salt marsh. We developed five alternate ecosystem
models, ranging from simple to highly mechanistic, that incorporate changes in C3/C4 plant composition but differ in their assumptions about plant
growth, plant chemistry and belowground structure that accompany changes in species composition. We use two data sets, including soil cores and
elevation change measured in a 13-year CO2 enrichment study, to calibrate
and validate the models. We found the simplest model, in which species
changes are ignored, generated the highest overall likelihood to the observed soil profiles. However, we also found that excluding species-specific
root profiles, associated with changes in species composition, generated the
highest likelihood to observed soil profiles of the specific organic chemistry
types. Finally, models that included species-specific root profiles matched
observed changes in elevation better than models that did not. In summary,
while a simple model that excludes changes in species composition (and
species-specific root profiles in particular) may be useful as a baseline
predictor of soil accumulation, the importance of root profiles in successfully predicting elevation change suggests important feedbacks between
elevation change and species composition. Given that the relationship between sea-level rise and elevation change may control processes such as
erosion, saltwater intrusion, and potential habitat loss, understanding the
links between species changes, belowground structure, and elevation
change will be crucial for successfully predicting long-term dynamics of
marshes in the Chesapeake Bay.
SAUNDERS, THOMAS J.* and MICHAEL E. MCCLAIN. Department of
Environmental Studies, Florida International University, Miami, FL, USA.
Nitrogen dynamics in surface and subsurface waters of a montane forest, headwaters of the Amazon, Peru.
We investigated the dynamics of runoff and nitrogen cycling in a pristine
first-order catchment in a montane rainforest of the Peruvian Amazon. The
study-site is located at 2520 masl in a region characterized by steep hillslopes (;45 degrees) and over 2,600mm/yr of precipitation. Suction-lysimeters and piezometers were installed along an upland-riparian-stream transect and samples were collected weekly over a 9-month period and analyzed for NO3-, and NH4+. Storm events were sampled for precipitation,
throughfall, overland flow, soil-water, groundwater, and stream flow as circumstances permitted. Hydrologic data from a v-notch weir 30m below the
site, a weather station within 300 meters distance, and from riparian piezometers demonstrate abrupt storm-induced fluctuations in both hydraulic
head and discharge, resulting in a marked pulse in system nitrate concentrations and export through a storm cycle. During baseflow conditions,
significant differences (P,0.01) in average NO3- concentrations are found
between upland soil-water (27.2628.3mM) and both riparian ground water
(4.364.3mM) and stream water (1.662.0mM). Ammonium concentrations
remained consistent between upland soil-water and riparian groundwater,
at 3.663.0mM and 3.764.4mM respectively, and drop to 2.762.3mM in
stream water. Preliminary dissolved organic nitrogen data suggest mean
values much greater than those reported for total inorganic nitrogen in all
compartments. Initial measurements of soil N2O fluxes have been low. Although riparian NH4+ concentrations are generally very low when compared
against sites studied in the lowland Amazon and in other tropical catchments, resulting streamwater ammonium concentrations are found to be
slightly higher. In contrast, riparian nitrate concentrations in the Amazon
headwaters site are generally higher than those found in other tropical riparian zones, with both of these trends partially owing to marked differences in the hydro-geomorphic makeup of the regions studied. Overall
patterns of N concentrations within upland/riparian/stream compartments
demonstrate that these montane forest catchments are exceptionally efficient in retaining nitrogen.
SAVAGE, VAN M., * JAMES F. GILLOOLY, JAMES H. BROWN,
GEOFFREY B. WEST1,2 and ERIC L. CHARNOV.3 1 Santa Fe Institute,
Santa Fe, NM, USA; 2 Los Alamos National Laboratory, Los Alamos, NM,
USA; 3 University of New Mexico, Albuquerque, NM, USA. Effects of
size and temperature on population growth.
1,2,

1,3

Since the time of Malthus, population growth has been recognized as providing a critical link between the performance of individual organisms and

the ecology and evolution of species. We present a theory that shows how
the intrinsic rate of exponential population growth, rmax, and carrying capacity, K, depend on individual metabolic rate and resource supply rate.
To do this we construct equations for the metabolic rates of entire populations by summing over individuals, and then combine these populationlevel equations with Malthusian growth. These individual-level and population-level processes are inextricably linked because metabolism sets both
the demand for environmental resources and the resource allocation by
individuals to survival, growth, and reproduction. We use this theory to
explain how and why rmax scales with body size and temperature. Data for
algae, protists, insects, zooplankton, fishes, and mammals confirm these
predicted scalings. The metabolic flux of energy and materials also dictates
that the carrying capacity for populations increases with increasing body
size and temperature. We then extend the theory to argue that most of the
variation in mortality and fecundity rates is also determined by body mass
and temperature. These predictions are supported by data for instantaneous
mortality rates of marine fishes in the field. This theory links rates of
metabolism and resource use of individuals to life history attributes and
population dynamics for a broad assortment of organisms, from unicells to
mammals.
SAX, DOV F.* and STEVEN D. GAINES. University of California,
sax@lifesci.ucsb.edu, Santa Barbara, CA. Species diversity: from global
decreases to local increases.
Current patterns of global change may strongly affect patterns of biodiversity at global, regional and local scales. At global scales habitat destruction and the introduction of exotic species are contributing to declines in
species diversity. At regional and local scales evidence for declines in diversity are mixed, and recent work suggests that diversity may commonly
be increasing. Here we examine evidence that indicates how species diversity is changing across spatial scales for plants, birds and freshwater
fishes. This evidence suggests that global decreases in diversity are commonly contrasted by increases in diversity at regional and local scales. This
evidence also shows that the magnitude of increases in diversity may be
highly predictable for particular taxonomic groups on islands and within
continental regions.
SCHAAFSMA, HOSKI* and JOHN M. BRIGGS. Arizona State University, Tempe, Arizona. A legacy on the modern landscape of prehistoric
farming in the Sonoran Desert of Arizona.
In order to measure the legacy effect of prehistoric farming in the extant
plant community within the Sonoran Desert, research was conducted on
prehistoric Hohokam agricultural fields (abandoned ca 750 years B.P.).
Quarter-point sampling transects were used to sample the woody and cacti,
plant communities, while the herbaceous plant community was sampled
with 100m x 0.5m transects. Results from the point-quarter sampling provided data on spatial distribution and density of the woody and cacti communities on and off fields. Replicate off-field transects were located in
similar geographic settings. Prehistoric farmers altered the soils on some
fields, with little soil alteration on others. The total plant density for woody
species on the soil-altered fields is estimated to be 822 plants/ha while a
total plant density in off field locations is estimated at 4795 plants/ha, a
large decrease in density on abandoned fields. Similar trends are documented in species richness. Fields with altered soils show a significant
change (t 5 4.53, p 5 .002) in the species density of modern vegetation
over non-farmed areas. Fields with minimally altered soils exhibit little
change in the modern plant community when compared to non-farmed
areas. Thus the extant vegetation in the Sonoran Desert can be impacted
by human activities occurring over 750 years ago.
SCHADE, JOHN D.,1 SARAH E. HOBBIE,2 RYAN SPONSELLER1 and
SCOTT L. COLLINS.3 1 Arizona State University, Tempe, AZ; 2 University
of Minnesota, St. Paul, MN; 3 University of New Mexico, Albuquerque,
NM. Spatial and temporal variation in islands of fertility in a Sonoran
Desert landscape.
The influence of canopy trees and shrubs on soil nutrient cycling and understory plants is complex and context-dependent. Canopy plants can exert
positive, negative or neutral effects on nutrient cycling rates and produc-

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297

tion, composition, diversity and nutrient content of understory plant communities, depending on local environmental conditions and position in the
landscape. We studied spatial and temporal variation in the influence of
mesquite (Prosopis velutina) on soil moisture and nitrogen cycling rates,
and understory vegetation along a topographic gradient in the Sonoran
Desert. We found significant increases in both soil moisture and N along
the gradient from desert to riparian zone. In addition, mesquite canopies
had positive effects, relative to open areas, on soil moisture in the desert,
and soil N availability, potential N mineralization rates and microbial biomass N in both desert and intermediate terrace, particularly during wet
months. Biomass of understory vegetation was highest and species richness
was lowest in the riparian zone. Canopies had a positive effect on biomass
in both desert and terrace, and a negative effect on species richness in the
terrace. The effect of the canopy depended on landscape position, with
desert canopies more strongly influencing soil moisture and biomass and
terrace canopies more strongly influencing soil N and species richness.
Individual species distributions suggested interspecific variation in response
to water- vs. N-availability strongly influences species composition at both
patch and landscape position levels.

have been found that enable the reproductive synchronization requisite for
mast seeding to occur. These cues tend to be linked with large-scale climatic phenomena like El Nino-Southern Oscillation, the frequency and
intensity of which are expected to increase as global temperatures rise. We
examine through empirically based models how climate change could alter
the frequency, intensity, and synchrony of masting, and examine impacts
on plant populations, seed consumers and their ecological associates. The
results depend strongly on the phenotypic plasticity of the masting cue
response. The linkage between oak masting, mice, and gypsy moths provides an informative case study. In temperate zones, warming is expected
to have disproportionate impact on the presence and depth of snow, which
can have inverse effects on many herbivores (whose foraging is hindered
by snow) and plants or seeds (for which snow can provide protection from
extreme cold and herbivores). We review the case of invasive musk thistle,
in which biological control agents may interact synergistically with snow
cover to suppress the population. These mechanisms represent but a small
subset of possible effects of climate change on plant-animal interactions.
However, focusing on cues, constraints, and other nonlinearities is likely
to accelerate our understanding of how plant-animal interactions will
change as the world warms.

SCHAEFFER, SEAN M.,1,* SUSAN ZIEGLER,1 JAYNE BELNAP2 and

R.D. EVANS.1 1 University of Arkansas Stable Isotope Laboratory, Department of Biological Sciences, University of Arkansas, Fayetteville, AR,
USA; 2 U.S. Geological Survey, Forest and Range Ecosystem Science Center, Moab, UT, USA. Alterations in soil microbial carbon and nitrogen
cycling in an arid Colorado Plateau grassland with Bromus tectorum
invasion.

SCHEESSELE, ERIN A.,* TIFFANY S. GARCIA and ANDREW R.

BLAUSTEIN. Oregon State University, Corvallis, Oregon. Indirect effects of ultraviolet-B radiation on amphibians: using mesocosm experiments to examine UV-B effects on trophic level interactions.

Previous studies have shown that Bromus tectorum invasion in arid Colorado Plateau grasslands has significantly changed the quality and quality
of litter deposition. Our objectives were to determine what effects these
changes in substrate quantity and quality may have on microbial utilization
of carbon (C) and nitrogen (N). We measured net inorganic C and N mineralization, and the d13C of bulk organic matter, respired CO2, and microbial
phospholipid fatty acids (PLFA) from soils of C3 (Stipa hymenoides) and
C4 (Hilaria jamesii) native grasslands invaded by a C3 annual grass (Bromus). After 24 h incubation under controlled conditions, carbon mineralization was lower in non-invaded compared to invaded plots (1.25 6 0.09
and 1.52 6 0.18 mg CO2-C g-1 soil h-1 respectively). Total PLFA C increased with invasion (6.1 6 0.5 and 4.6 6 0.4 mg C kg-1 soil for invaded
and non-invaded respectively), along with a decrease in the relative abundance of fungal PLFA biomarkers. Net inorganic N mineralization (NH4+
and NO3-) was lower in invaded compared to non-invaded plots in C3 stands
(-0.32 6 0.03 and 1.04 6 0.11 mg N g-1 soil respectively). In non-invaded
C4 plots, the d13C of a bacterial fatty acid, a15:0 (-19.1 6 0.3 ), was
lower than that in invaded C4 plots (-21.3 6 0.4 ) and the bulk organic
matter (-21.5 6 0.5 ). The d13C of mineralized C was also lower in soils
from non-invaded C4 plots compared to invaded C4 plots (-20.3 6 0.8 and
-22.0 6 0.3 respectively). Greater C and N mineralization and decreased
fungal abundance in invaded plots suggests proportionally greater bacterial
activity in invaded compared to non-invaded plots. Isotopic data suggests
that microbes in C4 non-invaded plots are utilizing a significantly greater
proportion of C4 organic matter than in C4 invaded plots.
SCHAUBER, ERIC M.,1,* DAVE KELLY,2 RICHARD S. OSTFELD3 and
CLIVE G. JONES.3 1 Cooperative Wildlife Research Laboratory, and Department of Zoology, Southern Illinois University, Carbondale, IL; 2 Department of Plant and Microbial Sciences, University of Canterbury, Christchurch, New Zealand, New Zealand; 3 Institute of Ecosystem Studies, Millbrook, NY. How will climate change affect plant-animal interactions?
A look at cues and constraints.
Many of lifes processes change in a continuous manner as environmental
conditions change within particular bounds, but small additional environmental shifts can cause disproportionate effects. As starting points from
which to examine potential effects of climate change on plant-animal interactions, we focus on two such nonlinearities: cueing of mast-seeding and
seasonal constraints. Mast-seeding (or masting) is a powerful ecological
driver that both improves reproductive success of participating plants and
initiates chain reactions that ripple through animal communities. For several plant species, disproportionate responses to subtle temperature cues

298

Abstracts

While most of the existing information on the role of ultraviolet-B (UVB) radiation in aquatic ecosystems is at the organismal or suborganismal
level, it is essential to investigate the community and ecosystem level responses to UV-B. Previous studies have found counterintuitive increases in
productivity under elevated UV-B due to differential sensitivity between
algae and their consumers. We used a three-trophic level mesocosm experiment to explore community level responses to ambient UV-B. We hypothesized UV-B exposure would differentially affect adjacent trophic levels of a simple food web with potential effects on the life history characteristics of amphibian larvae. For the three-trophic level food web, we used
a natural algae assemblage as the primary producer, a natural zooplankton
community as the primary consumer and long-toed salamander larvae (Ambystoma macrodactylum) as the top predator. When exposed to ambient
UV-B, we predicted the algal community would experience a change in
species composition and would tend to increase in biomass. We predicted
the density of the primary consumer (zooplankton) would be correlated
with UV-B exposure. The top predator, long-toed salamander larvae, would
most likely be negatively affected by a combination of direct and indirect
UV-B effects. We manipulated levels of UV-B exposure and trophic complexity, having either two or three trophic levels. At regular intervals we
measured species composition and biomass of the primary producer trophic
level, abundance and size and age distribution of the primary consumer
trophic level, and mortality and growth rates of the top level consumers.
The results of this experiment will be discussed.
SCHEINER, SAMUEL M.* National Science Foundation, Arlington, VA.
Path analysis and measuring natural selection.
Crucial to understanding the process of natural selection is characterizing
phenotypic selection. The traditional method uses multiple regression to
dissect total selection into portions due to direct selection and indirect selection through other traits. However, such an analysis can mis-identify the
causal components of phenotypic selection because the wrong causal model
is being specified. Path analysis allows for more complex causal structures
to be analysed. Path analysis also address two long-standing problems with
selection analyses: the problem of missing traits and environmental bias. I
show how path analysis can result in a different understanding of patterns
of selection, and it strengths relative to a multiple regression approach. I
also discuss available software that makes such analyses as easy to do as
multiple regression.
SCHIEL, DAVID R.1,* and BERNARDO R. BROITMAN.2 1 University of
Canterbury, Christchurch, New Zealand, New Zealand; 2 University of California, Santa Barbara, California, USA. Large-scale sampling in coastal
ecosystems: ecological patterns and processes in New Zealand, Chile
and Western USA.
There is considerable debate about the relationships between the distribution and abundance of coastal species and nearshore oceanographic pro-

cesses. Large-scale surveys along the coastlines of Western USA, Chile

and New Zealand show that ecological patterns have no simple relationship
with general oceanographic conditions characterized through mean sea-surface temperature or upwelling indices. Abrupt ecological discontinuities are
not accompanied by evident changes in general oceanographic conditions.
The heterogeneous coastline of New Zealand further challenges the predictive power of general oceanographic conditions. Here, the east and west
coasts have distinct rocky shore communities, regardless of the broad distribution of most species, and considerable localized variation. Apparent
ecological discontinuities are associated with changes to coastal currents
and geomorphology, degree of wave exposure, and sedimentation regime,
all of which interact in determining ecological patterns. We highlight some
of the lessons from these sampling programmes and the challenges arising
from using long term / large-scale sampling to link localized and broader
phenomena.
SCHILE, LISA M.* and SUSAN MOPPER. University of Louisiana, Lafayette, Lafayette, LA. Leafminer performance on brackish and freshmarsh Iris hexagona under different salinity treatments.
The effect of plant stress on plant-insect interactions is a highly debated
topic. My research addresses the plant stress hypothesis which states that
plants under physiological stress are of higher nutrient content and thus
serve as better hosts for insect herbivores. I focused on the effect of salinity
stress on the performance of the leafminer, Cerodontha iridiphora, on a
native wetland Louisiana iris, Iris hexagona. A common garden experiment
was conducted to address this question. The experiment used I. hexagona
plants collected from populations on Marsh Island, a brackish marsh located off the gulf coast of Louisiana. Salinity treatments of 0 and 8
NaCl were employed to test the effects of salinity-stress on leafminer recruitment, performance, and mortality. I monitored the number of mines
per treatment during the period of active feeding and the weight of larvae
after pupation. In addition, I measured the amount of leaf senescence on
mined and un-mined leaves to evaluate the effect of mining activity on
leaves under stressed and non-stressed conditions. There was no significant
difference in leafminer pupal weight on stressed and non-stressed plants;
however, significantly more mines were located on non-stressed plants.
Leaves with mines had significantly more senescence than un-mined leaves.
This effect was the strongest on salinity-stressed I. hexagona, resulting in
complete leaf death. Salinity stress significantly increased total foliar nitrogen as predicted by the hypothesis, but it did not improve leafminer performance. These data suggest that the plant stress hypothesis is not applicable in this system. More comprehensive research is currently being conducted to further examine this question in another common garden experiment using iris populations from Marsh Island and an intermediate-fresh
marsh located along the gulf coast.
SCHILLING, ERIK B.* and GRAEME LOCKABY. School of Forestry
and Wildlife Sciences, Auburn University, Auburn, Alabama, USA. Productivity and biogeochemistry of two contrasting floodplain forests.
Nutrient circulation patterns were examined to provide insight into the nature and driving forces behind the variation in production observed within
floodplain forests. Located in Georgia, USA, the Satilla River floodplain
(SAT), an oligotrophic, blackwater river system, and the Altamaha River
floodplain (ALT), a eutrophic, redwater river system, represent geochemical
and biogeochemical contrasts typical of many Southeastern floodplain forests. Measurements of aboveground net primary production (ANPP), litterfall nutrient contents, and resorption proficiencies were made in 1999
and 2000 within plots covering a range of topographic positions across
both floodplains. A greenhouse fertilization experiment was also used to
examine seedling growth responses to N and P additions. For the ALT,
ANPP (2-yr mean 5 10.9 Mg ha21 yr21) exceeded the SAT by 2 Mg ha21
yr21. Litterfall production within the SAT and ALT averaged 5.3 and 5.7
Mg ha21 yr21, respectively. Slightly larger amounts of N circulated through
litterfall on the SAT (40 kg ha21 yr21) compared to the ALT (37 kg ha21
yr21); however, P circulation in litterfall on the SAT (4.0 kg ha21 yr21) was
more conservative than the ALT (8.0 kg ha21 yr21). The relationship between litterfall mass and litterfall P content, coupled with efficient P circulation, highly proficient P resorption, and growth responses to P fertilization suggest P limitation on the SAT. On the ALT, litterfall N/P ratios

(,6.0), efficient N circulation, and proficient N resorption suggest N limitation. SAT base cation circulation patterns suggest low circulation and
high efficiency of use for Ca and K compared to the ALT. Conversely, the
ALT displayed low Mg circulation and high use efficiency. Relationships
between productivity and nutrient circulation from these two systems appear to be strongly driven by differences in P availability and suggest that
a broader array of nutrients limit productivity in oligotrophic compared to
eutrophic floodplain forests.
SCHLESINGER, WILLIAM H.* Nicholas School of the Environment and
Earth Sciences, Duke University, Durham, NC, USA. Global role of
rangelands in biogeochemical cycles.
Arid and semiarid systems cover nearly 30% of the Earths land surface
and harbor nearly 20% of the human population. Many of these lands are
used for grazing and managed as "rangeland." Not contributing a large
fraction to global NPP, soil carbon sequestration, or nitrogen cycling, can
arid lands be ignored in studies of global biogeochemistry? Studies of past
changes in the transport of dust to the polar ice caps indicate that the
signature of arid lands on global function has varied widely-with large
transports of dust at the Last Glacial Maximum affecting planetary albedo,
ocean productivity, and the transport of loess and soil microorganisms to
distant lands. Model predictions of transient mid-continental drought with
global warming compel us to understand local biogeochemical processes
in deserts, to predict how they might change in the future. Studies in the
Jornada Basin (LTER) in southern New Mexico and other sites in the arid
Southwest show that the biogeochemistry in deserts is often determined by
the distribution of the sparse plant cover-with semiarid grasslands showing
a relatively uniform distribution of soil nutrients in contrast to a patchy
distribution in associated with vegetation in shrublands. Nutrient losses by
erosion are greater in shrubland habitats. However, when semiarid grasslands are lost in favor of desertified shrublands, the total nutrient pool in
the landscape shows relatively little change. Rather, desertification is associated with a redistribution of soil nutrients on the landscape. Future
studies of biogeochemistry in arid habitats should focus on the episodic
removal of soil fertility by wind erosion.
SCHMALZER, PAUL A.* and TAMMY E. FOSTER. Dynamac Corporation, Kennedy Space Center, Florida. Characteristics of long-unburned
scrub on the Merritt Island/Cape Canaveral barrier island complex
before restoration.
Restoration of long-unburned scrub on the Merritt Island/Cape Canaveral
complex has been underway since 1992. Florida scrub ecosystems are
shrublands maintained by relatively frequent fire. With fire suppression and
landscape fragmentation, some scrub has reached a size structure resistant
to fire under typical prescribed burning conditions. Without burning, the
habitat suitability of scrub for a variety of species declines. We obtained
pretreatment data from 186 permanent 15 m line-intercept transects in 29
stands across the barrier island complex. On acid soils (11 stands) oak-saw
palmetto scrub occurred with Quercus myrtifolia, Q. geminata, Q. chapmanii, Serenoa repens, and Lyonia ferrugineaas the dominant species. Carya floridana was important in only one stand, as was Ceratiola ericoides.
Acid scrub soil series include Paola, Pomello, Astatula, Orsino, and Immokalee. On less acid to circumneutral soils, (Cocoa, Welaka series) oaksaw palmetto scrub (6 stands) occurred although with ericads less abundant.
On younger, alkaline soils (Canaveral, Palm Beach series), community
composition changed to coastal scrub (12 stands). A shrub form of Quercus
virginiana was the dominant oak; Q. geminata, Q. chapmanii, and ericads
were absent. Quercus myrtifolia occurred in transitional areas, but Serenoa
remained abundant. Myrcianthes fragrans and Persea borbonia increased
in abundance in coastal areas. Mean height of unburned stands ranged from
1.5 - 7.5 m; height differences reflected different periods of fire suppression
and probably different growing conditions.
SCHMIDT, BENEDIKT,1,2,* REINER FELDMANN3 and MICHAEL
SCHAUB.4 1 University of Zurich, Zurich, Switzerland; 2 KARCH, Bern,
Switzerland; 3 Pfarrer-Wiggen-Strasse, Menden, Germany; 4 University of
Bern, Bern, Switzerland. Demographic contributions to population
growth and decline in a salamander.
Amphibian populations are declining worldwide. Efficient conservation action requires that we understand which stage(s) in the complex life cycle

Abstracts

299

of amphibians contribute most to population growth (or decline) and which

stage(s) are most sensitive to environmental change. Here, we study the
demography of a stationary and a declining population of the salamander
Salamandra salamandra, and assess whether an observed decline in one
population is due to a change in recruitment or adult survival. The two
populations were studied for c. 20 years and all salamanders were known
individually. We analyze this data set using a multistate capture-mark-recapture model with an unobservable state. Our analysis shows constant
adult survival and variable recruitment in the stationary population and
declining adult survival and constant recruitment in the declining population. This suggests that the decline is due to a change in adult survival,
which was probably caused by a change in forest management. The common paradigm in amphibian population ecology is that population growth
is determined at the larval stage. Our results show that this may not always
be the case.
SCHMITZ, DENINE M. and DUNCAN T. PATTEN. Montana State University, Bozeman, 59717. Riparian patch types as indicators of watershed hydrologic complexity.
Watershed characteristics and climate drive catchment hydrology, which
controls the magnitude of flood disturbance regimes. These, in turn, drive
riparian vegetation dynamics. This study in the northern Greater Yellowstone Ecosystem (GYE) quantifies the relationship between watershed characteristics and the spatial structure of riparian patch types using hydrology
as the common element. In the GYE, watershed alterations such as fire and
grazing have modified flood magnitude and frequency consequently changing the character of associated riparian areas. Applying the Intermediate
Disturbance Hypothesis, a gradient of frequency and magnitude of hydrological events should produce successional diversity represented by riparian
patch type structural variability. Riparian patch types were ranked relative
to average flood frequency and stream power. Flood frequency and magnitude were related to watershed characteristics using canonical correlation
analysis. Once hydrologically linked, riparian patch types were then related
to watershed characteristics. Recently burned watersheds in the northern
GYE tended to produce riparian sites with early seral stages, which were
characteristically simple in vertical and horizontal structure. Riparian sites
in unaltered watersheds were generally diverse in patch type spatial structure. DCA Axis 2 scores show that patch types dominated by deciduous
and herbaceous species appear to represent seral stages better than those
dominated by coniferous species with those dominated by deciduous species representing intermediate stages. Thus, regions of a watershed with
riparian canopies dominated by deciduous and herbaceous species are likely
to yield more information through remote sensing techniques than those
dominated by coniferous species. These data suggest that the character of
the riparian zone through linkages to surface hydrology may be indicative
of basin attributes, a useful connection for watershed management.
SCHNEIDER, DAVID C.* Ocean Sciences Centre, Memorial University,
St. Johns, NL, Canada. Spatial and temporal scaling in minuscule ecosystems.
Some habitats are extensive, while many habitats are scattered sparsely
across the landscape. Such habitats can be viewed as a collection of minuscule ecosystems, in which species interactions are shaped by highly
variable densities of component species. If units are naturally small then
scale-up from a sample to the landscape is accomplished relative to environmental gradients, taking into account the size of each unit. Where small
size is imposed the magnitude of the reduction becomes another spatial
scaling factor. Temporal variance is expected to increase with increasing
isolation and decreasing size. Minuscule ecosystems are promising for observational and experimental studies of non-equilibrium dynamics.
SCHNURR, JACLYN L.1,* and BEVERLY S. COLLINS.2 1 Sheldon Jackson College, jschnurr@sj-alaska.edu, Sitka, AK; 2 Savannah River Ecology
Lab, Aiken, SC. Fire suppression and Quercus falcata seedling recruitment in southeastern forests.
Without fire, southeastern pine forests undergo succession to mixed pinehardwood stands. We transplanted Quercus falcata (southern red oak) seedlings into 10 sites on the Savannah River Site (SRS), near Aiken, SC, to

300

Abstracts

determine if this common upland oak shows increased recruitment in the

absence of fire. The sites included unmanaged dry ridgetop sandhills (N52)
and more mesic pine-hardwoods on slopes (N52), and managed stands
with different burn histories [unburned since 1950 (N52), burned in the
mid-1990s (N52) and recently burned (N52)]. We asked: Does survival
of transplanted Q. falcata seedlings differ among community types and
land management treatments? We measured initial seedling height and stem
diameter, and soil moisture in each plot in April, 2002. We censused survival in May, July, and October and measured seedling growth in October.
Canopy openness above each plot was measured in midsummer. Seedling
survival differed among the sites, with greatest survival at the sites unburned since 1950. Canopy openness significantly decreased seedling survival, while soil moisture in May increased survival. However, there was
no effect of community type on seedling growth: seedlings that survived
grew equally well in all sites. Our data indicate that fire suppression favors
establishment of Q. falcata, ultimately decreasing pine dominance in upland southeastern forests.
SCHOENNAGEL, TANIA,1,* MONICA G. TURNER,2 ANDREW FALL3
and DANIEL M. KASHIAN.4 1 University of Colorado-Boulder, Boulder,
CO; 2 University of Wisconsin-Madison, Madison, WI; 3 Simon Fraser University, Vancouver, BC, Canada; 4 Colorado State University, Fort Collins,
CO. The influence of climatically altered fire regimes on successional
patterns across the Yellowstone landscape.
Fire regimes throughout the Rocky Mountains are expected to vary with
climate change over the next century, yet landscape consequences of possible shifts in fire regimes remain unknown. We created a spatially explicit
probabilistic model using SELES to examine how variation in climate will
affect the fire regime and consequently the age and density of lodgepole
pine stands across a c. 500,000-ha landscape of Yellowstone National Park.
Based on a 2 x CO2 climate change scenario for the region, fire regimes
are expected to reduce mean stand age from 250 to 200 yrs. This decline
in mean age was not associated with significant change in mean stand
density across the landscape. Stand densities remained relatively unchanged
as significant self-thinning by lodgepole pine occurs predominantly in
stands , 100 yrs old, after which densities remain relatively constant over
time. However, the introduction of spatial variation in ignition probabilities
influenced the response of stand densities to altered fire regimes. When
low-elevation, highly serotinous stands preferentially burned, stand densities across the landscape increased and were more variable relative to highelevation stands where serotiny is low. Although adaptive responses to fire
frequencies are not represented in our model, serotiny may increase at
higher elevations with more frequent fire, resulting in denser stands across
the landscape. In response to climate change scenarios considered for this
region, the age structure of the landscape is expected to become younger,
yet stand structure appears to be much more resilient to wide variation in
fire regimes.
SCHOEPPNER, NANCY M.* and RICK A. RELYEA. University of Pittsburgh, Department of Biological Sciences, Pittsburgh, PA. Damage, digestion, and defense: Behavioral and morphological plasticity in response to different chemical cues.
Organisms change a wide range of behavioral, morphological, and lifehistorical traits in response to predators, but for the organism to correctly
match their defensive phenotype to the environment there must be reliable
cues about the current level of predation risk. In aquatic systems, chemical
cues are given off by both predators (kairomones) and prey (alarm pheromones) and both cues can induce prey defenses. While alarm pheromones
and kairomones induce changes in prey phenotypes, these cues could play
different roles in inducing different traits. In this experiment, we examined
the effects of nine consumed and nine crushed diets on the behavior and
morphology of gray treefrog tadpoles (Hyla versicolor). The consumed
diets released a combination of kairomones and alarm pheromones, while
the crushed diets only released alarm pheromones. We found that behavior
and morphology were affected by predator diet as well as by differences
between crushed and consumed prey.

SCHOETTLE, ANNA W.,1,* CARRIE BERGER1 and VERONIQUE H.

BONNET.2 1 Rocky Mountain Research Station, Fort Collins, CO, USA;
2
Matcom at Rocky Mountain Research Station, Fort Collins, CO, USA.
Proactive conservation options for bristlecone pine forests in the presence of an exotic pathogen use of fire?
Bristlecone pine (Pinus aristata) is at risk for infection by the exotic pathogen white pine blister rust (Cronartium ribicola). The rust is already contributing to mortality of limber pine (Pinus flexilis) in the northern Rockies
and into Colorado. The rust infection front is less than 100 km north of
the edge of the distribution of bristlecone pine; bristlecone pine forests will
be impacted in the near future. Proactive management is critical for these
slow growing species because the pathogen reduces cone production and
young trees established from rust resistant seed will not be reproductive in
the field until they are 40 to 60 years old. Protection of seed source stands
from wildfire while utilizing nearby burns as colonization and natural selection opportunities may accelerate the development of rust resistant pine
populations. We demonstrate that bristlecone and limber pine regenerate
well after large fires. In the Packer Gulch Fire of 1978 in central Colorado,
limber pine, and to a lesser extent bristlecone, establish throughout the
burned area whereas Douglas fir establishment is limited to near the unburned edge. These differences are likely a result of the mode of seed
dispersal (bird vs. wind dispersal). The density of Ribes ssp., the alternate
hosts for the blister rust, also increased in the burned area. While fire may
promote pine regeneration and, in the presence of the rust, possible natural
selection for rust resistance it may also increase the risk of the rust on the
landscape.
SCHOOLMASTER, DONALD R.* University of Michigan, Ann Arbor,
MI, 48104. Impact of the angiosperm parasite Cuscuta gronovii on a
wetland plant community.
Ecologists have long recognized that non-specific parasites are capable of
impacting the nature of their hosts interactions with other species and that
this can have a significant role in shaping community structure. Here, I use
an angiosperm host-parasite system to study the impacts of a generalist
parasite on host community structure. The parasite, Cuscuta gronovii, an
annual in the family Convovulaceae, grows in wetlands where it is most
often observed parasitizing the annual plant Impatiens capensis (Balsaminaceae), and the rhizomatous perennials Solidago patula (Asteraceae), Aster puniceus (Asteraceae) and Eupatorium maculatum (Asteraceae). Using
field observations and experiments, I tested the hypotheses that C. gronovii
infectiveness and virulence varied with the species of host. Field surveys
show that although C. grononvii is found to infect a number of host species
at the end of the growing season, all of the original infections occurred on
one species, I. capensis. These results suggest that only one of the many
host species is available to the parasite early in its life cycle. A field experiment where C. gronovii seedlings where tied onto potential host plants
confirmed this result. Twenty-one percent of 60 seedlings tied to I. capensis
were able to establish infections, whereas no seedlings tied to Solidago
patula or Aster puniceus were able to establish infections. Observations
also demonstrate that the virulence of C. gronovii varies among host species. Field grown I. capensis plants show no significant effect of C. gronovii infection. However, infected Solidago patula and Aster puniceus generated less biomass and fewer rhizomes compared to uninfected plants.
These data suggest that C. gronovii may be important factor in shaping
wetland plant community structure by reducing the cover of clonal perennials and thus facilitating the annual species I. capensis.
SCHOONOVER, JON E.,1,* B. G. LOCKABY1 and PHILIP L. CHANEY.2
School of Forestry and Wildlife Sciences, Auburn University, 108 M. W.
Smith Hall, Auburn, AL, USA; 2 Department of Geology and Geography,
Auburn University, 104A Tichenor Hall, Auburn, AL, USA. Changes in
water quality across an urban-rural landscape gradient: The West
Georgia project.
1

An interdisciplinary approach is being used to examine relationships between economics, biotic and abiotic indicators, and anthropologic variables
across the developing landscape of the Middle Chattahoochee Watershed
in western Georgia. A two-phase, watershed-scale study was established to
develop relationships between land use and water quality across an urban-

rural gradient. During phase 1 (i.e. year one), biweekly grab samples were
collected from pine, mixed forest, pastoral, developing, and urban watersheds ranging in size from 600-2500 ha. Nutrient loads, total suspended
solid load, dissolved organic carbon, and fecal coliform counts have been
determined. These data are being used to generate water quality prediction
models that are based on relationships between percent land use within a
watershed and water quality parameters. During the second phase (i.e. year
two), the models will undergo calibration and testing by predicting water
quality within the original database as well as newly chosen watersheds.
Also, during the second phase, sediment origin will be estimated within
the original 20 watersheds. The determination of sediment origin will be
achieved through iron and textural analyses of sediment combined with
channel morphology measurements taken in the headwaters of the streams.
Preliminary data show that fecal coliform and chloride are tightly related
to landuse, with values being the highest within the urban watersheds followed by pastured watersheds. Total dissolved solids are highest within the
urban and developing watersheds, while nitrate is highest in both urban
and pasture dominated watersheds.
SCHULZ, KURT E.,1,* TRAVIS BURLEYSON,1 STEPHANIE ALBRECHT,1 JOHN ZASADA,2 DAVID BUCKLEY3 and THOMAS
CROW.2 1 Southern Illinois University Edwardsville, Edwardsville, IL,
USA; 2 North Central Forest Experiment Station, Grand Rapids, MN, USA;
3
University of Tennessee, Knoxville, TN, USA. Ontogeny of herb layer
suppression by Acer saccharum regeneration thickets.
Sugar maple (Acer saccharum) dominated forests of Upper Michigan, USA
experience a natural regime of canopy gap formation at a rate of ca. 2%
per year. Selective harvest systems in the region employ multiple tree selection distributed over ca. 20-25% of stand area on a 20 year rotation.
Observations of natural forest understories following gap formation suggest
that understory herb communities lose species and decline in density in
response to thickets of regenerating maple. Regeneration thickets are commonplace in managed stands, and may imperil most of the understory in
the course of several harvest cycles. This study examined natural herb
populations across a gradient of regeneration thicket age. In general herb
cover and richness decline with time, showing the greatest depression in
plots containing large numbers of large saplings 5-10 cm dbh. Data distributions are "triangular", with wide ranges in cover and richness at low
density, and narrower ranges (and means) at high density. Light environments beneath thickets were compared across four developmental stages:
seedling/small sapling phase, large sapling phase, small tree phase, mature
phase. Both direct beam and diffuse light penetration are markedly lower
in the large sapling phase; the other phases are comparable. Reduced light
in the large sapling phase corresponds with particularly dense canopies at
zenith angles that obstruct midmorning and midafternoon sun.
SCHUPP, EUGENE W.1,* and REBECCA R. SHARITZ.2 1 Forest, Range,
and Wildlife Sciences, and The Ecology Center, Utah State University,
Logan, UT; 2 Savannah River Ecology Lab, Aiken, SC. Effects of burial,
litter, and canopy openess on Aesculus pavia seedling emergence.
Successful seedling emergence is influenced by many potentially interacting factors. For large-seeded species in particular, emergence is expected
to be maximum with seed burial and low insolation, conditions which
should increase the ability of seeds to hydrate. We experimentally tested
the effects of seed burial (3 levels; buried in soil beneath litter, beneath
litter on surface of soil, exposed on top of litter) and insolation (2 levels;
closed canopy, 10.2% openness; open canopy, 14.2% openness) on emergence of the large-seeded understory tree Aesculus pavia L. in bottomland
forest in South Carolina, USA. Seeds were sown in Fall in poultry wire
cages to protect from squirrels. Seedling emergence was recorded from 13
February through 29 March, after which no additional seedlings emerged.
The most obvious result was that emergence from exposed seeds (5%)was
significantly less than emergence from buried and litter seeds (both 65%).
Seedlings from buried seeds emerged significantly more rapidly than seedlings from litter seeds, but by 12 March cumulative emergence from buried
and litter seeds did not differ. The effects of canopy type were not straightforward. For buried and litter seeds emergence was significantly greater
beneath closed (77.5%) than open (52.5%) canopy, while canopy did not
affect emergence of exposed seeds. These results reveal complex effects

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301

of the environment on A. pavia seedling emergence. Completely exposed

seeds are not successful even if not consumed by seed predators. The advantage of burial in the soil, however, depends on whether there is an
ultimate selective advantage to earlier emergence in this species; if there
is, burial will be advantageous, if not, buried seeds will gain no advantage
over those that work down into the litter on their own. Importantly, only
small differences in canopy openness resulted in large differences in emergence.

conclude that many invertebrate inhabitants of ephemeral wetlands disperse

poorly. Hence, each wetland acts as an island for these poorly dispersing
species and the preservation of each habitat becomes critical in a landscape
that is increasingly fragmented.

SCHURER, MAAYKE, BRANDON S. SCHAMP* and LONNIE W.

AARSSEN. Dept of Biology, Queens University, Kingston, Ontario, Canada. If a tree bends in the forest, do we have any idea why?

The link between ecosystem structure and function is fundamental to our

understanding of ecology. Structures in urban areas are dynamic and difficult to define. This makes studying the structure-function link in cities
challenging. Anderson et al. (1976), an industry standard in land cover
classification, confounds structure and function thereby limiting the ability
to test their relationship. In addition, the resolution of this classification
scheme is not appropriate to accurately define the spatial heterogeneity
characteristic of urban systems. We developed a new classification that
separates structure from function and can be applied at finer spatial resolutions. The classification uses three components to define structure: 1)
building type and density, 2) vegetation texture and density, and 3) the
presence/absence of parking lots. We applied this classification to three
study areas along an urban to rural gradient in the Gwynn Falls Watershed,
Baltimore, Maryland. Using false-color infrared air photos, study regions
approximately 18 km2 were classified using on-screen polygon delineation.
Class richness and frequency, as well as measures of patch size, shape, and
distribution, were calculated using Patch Analyst (Remel et al. 1999). Regions closer to the city center are more heterogeneous, with 19% more
patches than suburban areas. Patches in urban areas are, on average, 19%
smaller than patches found in suburban areas. With cities becoming an
increasing component of land use cover, scientists can use this classification
in combination with social and biogeochemical datasets as a critical tool
to better understand the structure of cities and explore the functional dynamics of urban systems.

Understory saplings of deciduous trees commonly display a 9bent9 posture,

but the reasons for this are unknown. We explored three hypotheses: (1)
The 9biomechanical constraints9 hypothesis suggests that bending is due to
a relatively high stem length / diameter ratio resulting from strong apical
control and rapid vertical growth in low light conditions; (2) The 9light
harvesting9 hypothesis suggests that tree bending in the understory represents a shade-avoidance strategy since a more lateral growth form could
display leaves with less overlap across a greater horizontal area, thus increasing the potential for light harvesting under low light conditions; and
(3) The 9shade-avoidance9 hypothesis suggests that understory saplings exhibit directional bending toward sources of greater incident light (e.g. forest
edges and canopy gaps). We collected data on tree height, stem length,
stem diameter, and canopy light transmission for understory saplings of
three species in woodlands of southeastern Ontario. Sugar maple (Acer
saccharum), white ash (Fraxinus americana), and basswood (Tilia americana) saplings exhibiting a range of 9bendedness9 were sampled from three
height classes ranging between approximately 0.5 3.0m. Canopy light
transmission was assessed by image analysis of hemispherical canopy photographs using computer software (Gap Light Analyser). We found evidence supporting the biomechanical constraints hypothesis for mediumsized basswood saplings, but for no other size class or species. The light
harvesting hypothesis was supported for large-sized sugar maple saplings
only and the shade avoidance hypothesis was supported for small-sized
basswood saplings. Both shade-avoidance and shade-tolerance strategies
may be age/stage specific for these three tree species. It is also possible
that some tree bending in the understory is due to damage from ice loading
during the ice storm of 1998, or from herbivory induced activation of one
or more lateral meristems in place of the terminal meristem. Further data
is being collected to address these questions.
SCHWARTZ, STEVE S.* and GABRIEL COSYLEON. Oklahoma State
University, schwass@okstate.edu, Stillwater, OK. Lack of similiarity in
ephemeral wetland zooplankton communities among Oklahoma ecoregions.
Although ephemeral wetlands are globally ubiquitous our knowledge of
function and relatedness across the landscape is weak. We have studied the
similarity of microcrustacean communities of ephemeral wetlands across
Oklahoma. The biota of these habitats has evolved in a highly fragmented
and dynamic landscape and so must persist regionally by dispersal in time
or space. The relative importance of these strategies is unknown but recent
studies suggest that spatial dispersal is common. Given the intimacy of the
terrestrial landscape with these shallow (,1 m) habitats we tested the hypothesis that terrestrial ecoregions can be used to predict community assemblages of aquatic microcrustaceans. In the spring of 2001 and fall of
2002 we sampled ephemeral wetlands across Oklahoma. Species richness
of the zooplankton was determined for each habitat and compared within
and between ecoregions. In two adjacent ecoregions (Central Irregular
Plains and Ozark Mountains) that were examined in detail, 26 species were
identified in 21 habitats but most species were rarely encountered, with
each species found on average in 2.1 habitats with only 3.3 species/pond.
Means for both measures were not significantly different between the two
ecoregions. A cluster analysis based on relative abundance of species
showed that the species assemblages in these ecoregions are largely distinct, with few habitats placed in the incorrect cluster. The results of a
distance-similarity analysis indicates that there is no relationship between
the distance between habitats and the similarity of their faunas both within
and between ecoregions. Consequently, based on all these analyses, we

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Abstracts

SCHWARZ, KIRSTEN,* MARY L. CADENASSO and STEWARD T.A.

PICKETT. Institute of Ecosystem Studies, Millbrook, NY, USA. Toward
understanding ecological structure and function in urban systems: a
new land cover classification.

SCHWENDENMANN, LUITGARD1 and RUBEN LARA.2 1 Institute of

Soil Science and Forest Nutrition, University of Goettingen, Goettingen,
Lower Saxony, Germany; 2 Center for Tropical Marine Ecology, Bremen,
Bremen, Germany. Factors influencing dissolved organic matter and
dissolved inorganic nutrients in a mangrove forest near Braganca
(Para, Brazil).
This study focused on depth distribution and seasonal variation of dissolved
organic carbon and nitrogen, dissolved inorganic nutrients (ammonium,
nitrate, nitrite, ortho-phosphate and silicate) and salinity of waters from a
mangrove forest and an adjacent tidal creek located at the northeast coast
of Para (Brazil). Water samples were taken at different depth from boreholes and piezometers located within the mangrove forest and from the
tidal creek at neap and spring tides during dry and rainy season, respectively. In addition, sediment from the mangrove forest has been analyzed
for carbon, nitrogen, salinity and hydraulic properties. As surface and
groundwater inflow from neighboring areas is impeded, depth distribution
and seasonal changes of organic matter, nutrients and salinity can only be
attributed to the in/output of these elements via tidal creek, rainwater and
to biological, biogeochemical and physical processes within the sediment.
Compared with creek water, mangrove water was enriched in dissolved
organic carbon and nitrogen, dissolved inorganic nitrogen and silicate because of the input of organic matter and subsequent biological transformation. Thus, these mangrove forests may act as a source for carbon and
nitrogen to the adjacent tidal creek and estuary. The subsurface distribution
of organic matter and inorganic nutrients was mainly governed by biological and geochemical processes, but vertical and lateral transport processes
also seemed to play a role. Both processes were strongly influenced by
physical sediment characteristics in the different profile layers. Whereas
dissolved organic matter, silicate and salinity showed a distinct seasonal
pattern, dissolved inorganic nitrogen and ortho-P were invariant to seasonal
changes.
SCHWILK, DYLAN W.* and DAVID D. ACKERLY. Stanford University,
Stanford, CA. Functional diversity along climatic gradients: A spatial
model.
In heterogeneous environments, the distribution of plant species is the outcome of a series of biotic and abiotic filters: the community assembly

process. Recent developments in neutral models emphasize the importance

of incorporating non-equilibrial processes (e.g., ecological drift) in models
of niche-structured community assembly. Here, we examine community
diversity of sessile organisms in response to spatial variation in habitat.
Our framework combines the potential for ecological drift with non-neutral
competition for space in a heterogeneous environment. We constructed a
spatially explicit model to simulate the distribution of species in a onedimensional spatial landscape with an underlying gradient in environmental
conditions. Our objective is to examine the influence of niche breadth,
dispersal distances, community size (total number of individuals) and the
breadth of the environmental gradient on levels of species and functional
trait diversity. In our model, organisms response to the environmental
condition is described by a Gaussian fitness function and species vary in
the location of their fitness mode along the gradient. As expected, equilibrium diversity decreased with increasing niche breadth. Increasing dispersal
distance, however, increased diversity. Diversity increased linearly with increasing community size, consistent with neutral theory. Increasing the
range of the environmental gradient, however, also increased diversity and
niche-packing. The effect of environmental range is not linear: diversity
levels off as environmental range approaches zero and stochastic processes
become more prominent. Apparent limiting similarity emerges from the
effect of niche breadth and the stochastic effect of finite populations.
SCOBELL, SUMMER A.* University of Miami, Coral Gables, FL. GIS
and path analysis: Examining associations between the birds, the bees
and plant sex in Echinocereus coccineus (Cactaceae).
In plant families that exhibit dioecy, an association between the predominant pollinator type and the plant9s mating system has often been reported.
Plants with more specialized pollinators (e.g. birds, bats) are predominantly
hermaphroditic while species with depauperate, generalist or small pollinator fauna tend to be dioecious. It is hypothesized that the latter pollinator
types may increase self-fertilization rates of hermaphroditic plants, and the
inbreeding depression in selfed seeds then selects for dioecy. An alternative
hypothesis is that selection in arid environments favors dioecy because this
system allows more efficient allocation of scarce water resources to male
and female functions. There is also the possibility that the association between water resources and dioecy is indirect: A habitat shift to drier areas
may be associated with loss of more effective pollinators, which is proposed to select for dioecy. Echinocereus coccineus (Cactaceae) is an excellent model system to test hypotheses about links between pollinator type,
water resource gradients and dioecy because it possesses contemporaneously both hermaphroditic and dioecious populations growing over a wide
rainfall gradient. E. coccineus flowers appear specialized to hummingbirds,
but are also pollinated by small Halictid bees. Distribution and abundance
of these pollinator types varies across the plant9s geographic range and
dioecy appears to be associated with areas of low hummingbird abundance.
Dioecious populations also appear to be associated with drier, low altitude
areas. In order to better quantify these associations, a GIS database was
compiled containing data from over 500 herbarium specimens of E. coccineus, annual rainfall, and hummingbird abundance. The data from each
population was then analyzed using multiple regression and path analysis
to assess the relative direct and indirect influence of the predominant pollinator type and mean annual rainfall on the presence of dioecious populations in this species.
SCOFIELD, DOUGLAS G.* d.scofield@umiami.edu, University of Miami, Coral Gables, FL. Obgenetica: A flexible, individual based, genetically explicit modelling platform for population simulation.
I describe the design and initial capabilities of Obgenetica, a flexible, extensible stochastic simulation platform for the exploration of individualbased evolutionary questions within potentially complex population and
environmental models. Individuals are modeled via their genomes, with
behaviors and fitnesses dependent upon genotype and encompassing environment. Populations in turn are modeled as regulated collections of individuals occupying environmental patches which transfer individuals via
corridors; the specification of environmental structure is extremely flexible
and no particular model is implied by Obgeneticas "patch" and "corridor."
A script language is used to specify characteristics of individuals, populations, environments and simulation experiments; many published models

may be directly specified in the script language. Obgenetica is free, opensource, largely written in Javay, and employs straightforward, biologically-motivated object hierarchies. User extensions of any size are easily accommodated and strongly encouraged.
SCOTT, FINLAY AM.* and ALASTAIR GRANT. University of East Anglia, Norwich, Norfolk, UK. Sensitivity of density dependent populations to environmental variability.
Environmental variations (weather, toxin exposure, etc) alter demographic
parameters of individuals. How do these perturbations feed through into
effects on populations? Density dependence can reduce the effects of environmental stochasticity, but it is also possible for the two to interact in
a way that amplifies the environmental noise, as in the case of Dungeness
Crab and Soay Sheep. However, we lack a general framework to predict
how populations will respond to environmental noise. Here we perturb two
and three stage class population models, including the LPA model of Tribolium, with white noise, focussing on those parts of parameter space
where the unperturbed population has a stable equilibrium. We assess responsiveness by measuring the variance of the input and output time series,
and examine the ability of different elasticities to predict population responsiveness. The effect of these perturbations on the population can be
characterised using elasticities. The elasticity of the density independent
growth rate is effective as a general guide but for some forms of density
dependence the one-step elasticity performs better. Density dependence
may reduce impacts, but this depends crucially on its form and which parts
of the life history are impacted by the environmental shocks. Short-term
population consequences are sometimes greater than would be inferred
from conventional elasticities. The extent to which this occurs can be predicted by the magnitude of the stability determining eigenvalue.
SCOTT-DENTON, LAURA E.,* SARAH A. SCHLIEMANN, BRANT
BACKLUND and RUSSELL K. MONSON. University of Colorado, Boulder, CO. Tree girdling and soil carbon cycling at the Niwot Ridge
Ameriflux Site, Colorado.
The importance of the linkage between recent photosynthesis and belowground carbon cycling is becoming more clear as ecosystem-level carbon
flux datasets become larger and temporally dense. This study attempted to
examine this linkage by measuring soil carbon pools and soil respiration
in a Colorado subalpine forest after cutting off fresh photosynthate to plots
by girdling trees and trenching soil. Ten plots of approximately 10m2 were
established. In five of these plots all the trees (2-4) were girdled during the
winter while the trees were dormant. After snowmelt, the circumference of
the girdled plots was trenched through the organic soil layer to a depth of
approximately 30cm. Soil samples were taken from the organic layer
throughout the season, including the time before snowmelt. Soil respiration
measurements were taken after snowmelt on established arrays of collars.
Soil respiration was shown to be similar between the girdled and control
plots at the beginning and end of the season but to be reduced in the girdled
plots by approximately 50% at the mid-summer maximum. Soil microbial
biomass was also depressed in the mid-summer maximum in the girdled
plots, but seemed to show an ephemeral peak during snowmelt in the girdled plots that may represent a stimulation of the soil heterotrophs. Soil
fungi as measured by soil ergosterol was not significantly effected by girdling, nor did it vary significantly throughout the summer, though it seemed
to follow a similar pattern to the total microbial biomass. Soil soluble
carbon content showed a large peak during snowmelt in the control plots
that was absent from the girdled plots. The magnitude and timing of these
results suggest that the influence canopy above-ground processes can have
critical short-term effects on soil carbon cycling and the below-ground
community structure.
SEABLOOM, ERIC W.,1,* ANDY P. DOBSON2 and DAVID M. STOMS.3
National Center for Ecological Analysis and Synthesis, Santa Barbara,
CA; 2 Department of Ecology and Evolutionary Biology, Princeton University, Princeton, NJ; 3 Institute for Computational Earth System Science,
University of California, Santa Barbara, CA. Abiotic and biotic constraints on invasions: Vascular plants in California.

Invasion by exotic species is one of the most serious threats to global

biodiversity. The distribution of exotic species in the landscape is the result

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303

of complex feedbacks between anthropogenic disturbance, abiotic conditions, distance to sources of introductions, and the richness of the native
species pool. We have combined a unique data set containing the distribution of the nearly 5000 native vascular plants in California with data on
human impacts, climate data, and plant productivity. We use these data to
investigate the relative importance of factors constraining the distribution
of exotic plants. At the state scale, both native and exotic richness are high
in areas with high plant productivity. In addition, exotic species richness
also is higher in low-elevation coastal areas with high levels of human
activity. These areas with high levels of human activity and exotic species
also have the highest numbers of species threatened with extinction. This
pattern suggests that similar processes are increasing rates of both invasion
and extinction.
SEARS, ANNA L.W.* and PETER CHESSON. University of California,
Davis, CA. Spatial storage effect moderates species interactions in diverse ecosystems.
While many researchers have studied variation in plant competition intensity along environmental gradients, it has previously been difficult to quantify how environmental variation impacts plant population and community
dynamics. Spatial storage effect theory provides a set of tools to explain
how patchiness or gradients in the environment, combined with speciesspecific differences in density dependence under different environmental
conditions, can affect population persistence and the maintenance of species diversity. Using data from a variety of previously published neighborhood plant competition studies, we show that positive covariance between plant response to the environment and competition [Cov(E,C)], a
key component of the spatial storage effect, is widespread in diverse ecosystems. This covariance, which is only found in the presence of environmental variation and at higher spatial scales, acts to increase populationlevel intraspecific competitive effects. Theory predicts that when Cov(E,C)
is found for high-density, but not low-density species, it promotes species
diversity. We found, for the response variables tested, Cov(E,C) was not
always closely correlated with species density, which implies that in some
of these systems, spatial variation may promote competitive exclusion.
Cov(E,C) was also found to increase net population-level intraspecific competition intensity in systems where experiments showed a net positive effect
of neighbors (facilitation) at local spatial scales. The methods developed
for this study use new statistical techniques to extract previously inaccessible information from conventional experimental designs. Spatial storage
effect theory appears to provide a promising context for studying a wide
variety of metacommunity types and processes, and gives a depth of mechanistic understanding that was not possible using previous techniques.
SEARS, ROBIN R.1,2 1 Columbia University, New York, NY; 2 The New
York Botanical Garden, Bronx, NY. Trees in Amazonian agroecosystems: Ecological and economic benefits of smallholder timber management.
In Amazonia, the over-extraction of traditional timber species has provided
an incentive for private landholders to maintain and enrich impoverished
forests and agricultural fallows with valuable trees on their landholdings.
By enrichment planting and promoting the natural regeneration of timber
and fruit trees in their agricultural production systems and forests, Amazonian farmers restore the ecological function and reestablish the economic
value of high-graded forests. Near Iquitos, Peru, smallholder farmers on
the floodplain actively manage 22 tree species for production of timber,
fruits, medicine, and household items, as well as for maintenance of ecological function of this seasonally flooded ecosystem. One native tree, Calycophyllum spruceanum (Rubiaceae), is prominent in these managed
stands providing timber, fuelwood and construction material for local and
commercial use. This species, a fast-growing hardwood, can account for
up to seventy percent of stand basal areas in agricultural fallows or forests
where farmers use an array of silvicultural techniques to promote it and
other species. This and other timber species respond well to the environmental conditions of the swidden agricultural systems and silvicultural
practices of floodplain farmers. While the traditional approach of establishing parks and other protected areas has been recognized to play an
important role in conservation and the provision of ecosystem services, I
present data that demonstrates the high value of the tree component of

304

Abstracts

diverse and productive agroecosystems outside of protected areas on the

floodplain for conservation and sustainable use. The data suggest that
smallholder forestry should be considered as one of several conservation
practices promoted by government and non-government development and
conservation agencies.
SEKERCIOGLU, CAGAN H.,1,* GRETCHEN C. DAILY,1 PAUL R. EHRLICH,1 SCOTT LOARIE1 and VIVIANA RUIZ-GUTIERREZ.2 1 Stanford
University, Stanford, CA, USA; 2 Universidad Nacional de Costa Rica, San
Jose, San Jose. Habitat use and movements of native forest birds in
southern Costa Rican agricultural countryside.
We used radio telemetry to study the habitat use, activity budgets, and
movements of three bird species that are found in native forest and coffee
plantations of southern Costa Rica. We obtained 7522 locations from Turdus assimilis (n527), Tangara icterocephala (n553), and Catharus aurantiirostris (n542), the species ranked from most to least forest-dependent. We caught birds in coffee and radio-tracked them to understand their
dependence on remnant native vegetation patches. Home range size and
frequency of long-distance movement was positively correlated with the
species forest dependence. T. assimilis and T. icterocephala preferred forest fragments, riparian strips and remnant forest trees, mostly avoided coffee plantations, and moved frequently between suitable habitat patches.
Even though remnant forest trees and riparian strips covered 3.7% and
4.8% of the land area, respectively, they made up 9%-49% of T. assimilis
and T. icterocephala home ranges, and were especially important in the
dry season (25%-49%). C. aurantiirostris, an edge/second growth species,
was highly sedentary, opportunistic in its habitat preference in the wet
season, and preferred second growth in the dry season. Home range size
for C. aurantiirostris and T. icterocephala decreased significantly in the
dry season, concentrating around food resources in secondary growth and
fruiting trees, respectively. Native habitat patches were cooler, more humid,
had reduced microclimatic fluctuations, and increased in importance during
the dry season. Remnant patches of native vegetation in Costa Rican agricultural countryside provide crucial resources for these bird species that
evolved in forest, but now have to live in an increasingly deforested landscape. Active management of tropical human-dominated ecosystems can
significantly increase their capacity to support native bird species.
SELMANTS, PAUL C.* and STEPHEN C. HART. Northern Arizona University, Flagstaff, Arizona, USA. Nitrogen cycling across an Arizona
semiarid woodland soil chronosequence.
We examined nitrogen (N) pools and fluxes under pinyon pine (Pinus edulis) and one-seed juniper (Juniperus monosperma) canopies as well as in
intercanopy spaces along a three-million-year soil chronosequence in northern Arizona. We expected that N would accumulate in the soil with ecosystem development, differences in soil N content among the three canopy
types would lessen with time, and the d15N signature of both soils and
foliage would become more depleted with soil age. Interspace total soil N
increased roughly seven-fold from 25 g m-2 at the youngest site (1,000 yr)
to 184 g m-2 at the oldest site. Total soil N was consistently higher under
tree canopies than in the interspaces across the chronosequence, with the
largest differences among canopy types at the youngest site. Microbial
biomass N under pinyon and juniper canopies increased uniformly from 4
to 25 g m-2 across the chronosequence, while interspace microbial biomass
N increased from 2.5 g m-2 to 19 g m-2 at the second oldest site (750,000
yr) and then declined to 16 g m-2 at the oldest site. Contrary to our predictions, both total soil d15N and foliar d15N of the two dominant tree
species became more enriched across the chronosequence, suggesting an
increased rate of N loss with soil age. The dramatic increase in gross rates
of nitrification across the chronosequence suggests that either leaching of
NO3- or denitrification are potential pathways leading to enrichment of both
soil and foliar d15N.
SENDEL, TANIA T.* and DAVID J. CURRIE. University of Ottawa, Ottawa, Ontario, Canada. Can changes in bird population density at particular sites be related to changes in climate?
Population densities of many bird species are known to have changed significantly over the last half-century, and these changes have varied across

space. It has been hypothesized that these changes may be due, at least in
part, to climate change, which has also varied regionally. Since climate
models predict further dramatic changes for this century it is important for
conservation biologists to understand the extent and manner in which species respond to climate change, both in their efforts to maintain species
adaptability through land allocation for conservation and in management
of individual species. We ask the question: to what extent can changes in
bird population density at particular sites be related to changes in climate
at those sites? We investigate this question using data from the North American Breeding Bird Survey, one of the largest and most comprehensive
biodiversity databases available, and climate data from the National Climatic Data Center. Two-dimensional climate envelopes (temperature and
precipitation) were created for each of over 200 passerine species. We
determined whether climate change at each survey route was moving away
or toward the centroid of the climate envelope. After correcting for observer bias in the survey data, we tested the relationship between population trends at the survey route level and regional climate trends from 1970
to 2001.
SEYMOUR, MICHAEL A.* and LINDA M. HOOPER-BUI.* Louisiana
State University, Baton Rouge, LA, USA. Population estimates of red
imported fire ant, Solenopsis invicta Buren: Mound numbers versus
food traps.
Red imported fire ant, Solenopsis invicta Buren, is an invasive, exotic pest
species that has been implicated in the decline of native vertebrates and
invertebrates. Mound construction by this species makes nest detection and
quantification an easy method for estimation of fire ant populations. Researchers usually compare mound numbers across plots, habitats, and landscapes and often rely on mound numbers to quantify the effects of biological, chemical, and cultural control methods. Mound density is also useful
in examination of fire ant impact on animals and ecosystems. Large
amounts of precipitation may result in increased mound construction, and
high temperatures and low precipitation may result in decreased mound
construction. In order to determine if the number of nest mounds could be
used as a reliable indicator of fire ant population, I evaluated 10 sites three
times in spring, summer, and fall 2002 and three times in spring 2003.
Habitat type ranged from city park to patches of closed canopy forest.
Twenty foil-wrapped scintillation vials, ten containing approximately 4 g
of vienna sausage and ten containing approximately 1 ml of honey, were
placed along a 66.7 m line transect. I collected the vials after 50 minutes
and trapped the recruited ants inside. In addition, active ant mounds within
2 m of each side of the transect were counted. Total number of ants at each
site were pooled and compared to mound counts.
SEZEN, ZEYNEP* and KATRIONA SHEA. The Pennsylvania State University, University Park, PA. Dynamics of the invasive thistle Carduus
nutans and its specialist herbivore Rhinocyllus conicus: Effective biocontrol in patchy environments.
Musk thistle, Carduus nutans, is an invasive weed of major economic
importance and is listed as the second most noxious weed in the US. Its
specialist herbivore, the seed feeding weevil Rhinocyllus conicus has commonly been used for biocontrol, unfortunately with variable success. We
hypothesize that colonization and establishment of the insect herbivore relative to the patch dynamics of its host strongly determine the success or
failure of biological control efforts. To test these ideas, we have developed
a metapopulation model with an explicit spatial structure, incorporating
both plant dispersal and insect movement, and human-mediated insect augmentation. The model explores how the invader and its herbivore spread
through a landscape, and suggests ways in which biocontrol can be improved by strategic insect releases at the plant invasion front.
SHAPIRO, JOSSLYN B.,1,* JAMES D. LEWIS,2 DAVID T. TISSUE3 and
KEVIN L. GRIFFIN.1 1 Lamont-Doherty Earth Observatory of Columbia
University, Palisades, New York; 2 Louis Calder Center and Biological Station, Fordham University, Armonk, New York; 3 Texas Tech University,
Lubbock, Texas. Response of leaf respiration in the light to elevated
CO2 concentration, nitrogen availability and temperature.
In this study we investigated the leaf level response of respiration in the
light to CO2 concentration, nitrogen availability and short-term temperature

variation using the herbaceous species Xanthium strumarium. Respiration

that takes place during hours of illumination is a vital component of a
plants total energy and carbon balance and has been largely unstudied with
respect to possible responses to environmental changes. In this experiment,
the Kok method was used to measure respiration in the light (RL) and
respiration that takes place in the dark (RD) and yielded significantly lower
values for RL versus RD across CO2, nitrogen and temperature treatments.
Percent inhibition of respiration in the light was 30% for plants grown in
ambient CO2 and 18% and 35% respectively for plants grown in elevated
CO2 with a high and low nitrogen supply. RL values significantly increased
with increased CO2, nitrogen and temperature, and there was a strong CO2
by temperature interaction in plants grown in elevated CO2 and a high
nitrogen concentration. Furthermore, a significant negative correlation between RL and C:N concentration was shown. Additionally, data indicate
that the percent of respired carbon with respect to assimilated carbon increases as temperature increases for all CO2 and nitrogen treatments from
a low of 2.6% in the elevated CO2, high nitrogen treatment to a high of
11.4% in the elevated CO2, low nitrogen treatment. Using RL values in
place of often used RD values decreases estimated total daily respiratory
carbon losses by 18-35% across treatments. Since RL responds independently from photosynthesis to environmental variables, quantification of
respiratory carbon loss during the day has the potential to impact our understanding of the role of terrestrial vegetation in the global carbon budget,
and our ability to predict under which circumstances ecosystems will act
as sources versus sinks for carbon.
SHARITZ, REBECCA R.,1 STEVEN J. HARPER,1,* DONALD W. IMM,2
BEVERLY S. COLLINS,1 KATHRYN MADDEN1 and TRACEY TUBERVILLE.1 1 Savannah River Ecology Laboratory, Aiken, SC; 2 U.S. Forest
Service, Savannah River Institute, New Ellenton, SC. Impacts of military
training and land management on rare species in southeastern fall-line
sandhills communities.
In the southeastern U.S., the federal government has extensive land holdings near the fall line that occurs between the Piedmont and Coastal Plain.
Sandhills found within this region, which are remnants of ancient coastal
dunes, are nutrient-poor habitats on sandy, xeric soils. Extending from the
Carolinas, through Georgia and into Alabama, fall-line sandhills support a
unique flora and fauna, including a suite of rare plant and animal species.
The Department of Defense must address simultaneously the habitat sensitivities of these species along with demands associated with military training and other land-use activities. This complex challenge requires the integration of diverse information with understanding of processes operating
at multiple spatial and temporal scales. The goal of this collaborative research is to develop methods to evaluate effects of military training and
land management activities on the sustainability of sandhills communities
and associated threatened and endangered species. Forests on these military
installations are typically managed to promote open pine woodlands as
habitat for the endangered red-cockaded woodpecker (Picoides borealis),
though it is not known whether these efforts are also beneficial, or possibly
detrimental, to sandhill species. We have conducted GIS analysis and field
sampling at Fort Benning, Fort Gordon, and the Savannah River Site (SRS)
to delineate sandhills communities from adjacent forests. Results indicate
that sandhills tree composition is dominated by four oak species, with canopy openness ranging from 20-39%. Soil characteristics include high sand
content, low soil moisture, and low nutrients. Methods have been developed
to determine the microhabitat requirements of rare plant species, and to
experimentally re-introduce the gopher tortoise (Gopherus polyphemus) at
SRS. Future work will include extensive field surveys and experiments to
quantify the impacts of forest management practices on sandhills communities across a broad array of military land-use conditions.
SHAW, REBECCA.* The Nature Conservancy, 201 Mission Street, 4th
Floor, San Francisco, CA. Invasives, climate change, and conservation
challenges.
In a priority setting assessment, invasive species were identified as one of
the most serious and pervasive threats across The Nature Conservancys
Conservation Areas. The threat of climate change is also recognized to be
as menacing to The Nature Conservancys long-term goal of conserving
biodiversity. The scientific study of invasions under a changing climate is

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in its infancy but it is certain that invasive species will have expanding
opportunities for establishment under changing climate. When environmental conditions change such that native communities are no longer adapted to the environmental conditions of the habitat, the invasion of exotic
species could drastically alter community composition and successional
patterns. These issues are of particular concern to conservation organizations that must plan now to protect biodiversity in the future. While The
Nature Conservancy of California has been developing strategic plans for
addressing these threats individually, a more comprehensive approach to
addressing simultaneous and interacting, multiple global changes is needed.
This presentation will describe the current approaches to the invasive species and climate change threats and present an outline for a more comprehensive approach to multiple threats.
SHEIK, CODY,* TAYLOR OLIVER and LINDA L. WALLACE. University of Oklahoma, Norman, OK. Grasshopper herbivory of Solidago speciosa varies with distance from Juniperus virginiana in tallgrass prairie.
Earlier work has shown that Solidago speciosa is a favored forage for
grasshoppers in a tallgrass prairie landscape in central Oklahoma. This
work also indicated that plants closer to Juniperus virginana trees may be
more heavily grazed than those farther away. We examined possible reasons
behind this including an examination of abiotic factors, plant community
structure and physiological responses of S. speciosa to herbivory. We found
that soil temperature and light increased significantly with distance from
the tree, but that air temperature and relative humidity were unaffected.
Significant community differences were found at the canopy edge, but not
where the majority of herbivory was occurring. We also found no photosynthetic differences between grazed and ungrazed S. speciosa. However,
a significant negative relationship existed between grazing intensity and
distance from trees. Grazing intensity was greater on shorter plants, but no
relationship between distance and height was found. We hypothesize that
trees may represent a safe haven from avian predation, making forage
plants closer to trees more at risk for grasshopper herbivory.
SHER, ANNA A.,* DAVE SPENCER, SARA SWEET and JOE DITOMASO. Compensatory growth response across two nutrient regimes
by invasive Arundo donax.
Arundo donax (giant reed) is a pervasive invader of riparian systems and
is currently a candidate for biological control. We investigated response to
simulated herbivory for one year (2/02-2/03) of twelve Arundo individuals
grown in 1.86 m3 containers with horizontal mini-rhizotrons at 30 cm intervals. Plants received one of four treatments: either uncut or subjected to
cuttings that removed half of the above ground biomass, with either high
or low nutrient levels (approx. 24 mg/wk inorganic N vs. 1.6 mg/wk). We
monitored growth and architecture above ground with a 3-D digitizer, and
below ground with a mini-rhizotron camera. Photosynthesis and transpiration rates were measured using a LI-6400. A less-detailed, companion
study in the field on 8 pairs of plants (cut and uncut) was simultaneously
conducted. All 12 potted plants had roots .1 m deep within 5 months.
Cut plants had the highest rates of branching and the highest relative
growth rate (RGR) as measured by shoot height and number, leaf number,
and increase in mean diameter of new shoots. Nutrient treatment had no
effect on branching rates or RGR in most cases. Cut plants also had higher
photosynthetic rates than uncut plants, as did high nutrients relative to low
nutrients. The same photosynthetic patterns were seen for field plants. Our
results suggest that A. donax is capable of vigorous compensatory growth
in response to above ground herbivory, even under low nutrient conditions.
SHERRY, REBECCA A.,* LINDA L. WALLACE, YIQI LUO and NANCY ZEHRBACH. University of Oklahoma, Norman, OK. Phenological
differences among winter annuals in warmed plots in tallgrass prairie.
The phenology of winter annuals in warmed and unwarmed plots in tallgrass prairie in central Oklahoma were followed for a 9-month period (October to July). Twenty 1.22m by 2.44m plots with a surrounding 0.3m
buffer were observed monthly from October 2002 to February 2003, and
weekly beginning in March 2003. Half of the plots had two 1500 watt
Kalgo infrared heaters hanging in the center of the plots 1.4m above the
ground. The remaining ten plots have dummy heaters in the same position.

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The first two centimeters of soil in heated plots was 4 oC warmer than soil
in unheated plots during February 2003. Heating began in mid-February
2003. Japanese brome, Bromus japonicus, germinated by the beginning of
October 2002 and is the dominate winter annual in all plots. Two suites of
annual forbs were followed. The first set of forbs included small bluets
(Hedyotis crassifloria), ridgeseed spurge (Euphorbia glyptosperma), and
henbit (Lamium sp.). Hedyotis germinated in early November, while Euphorbia and Lamium germinated in late November. Hedyotis was the first
plant to bloom at the site, in late February. A second suite of species,
dominated by Rumex sp., germinated in January and consisted primarily
of rosette forming annuals. These rosette-forming annuals were much less
numerous than the first suite of forbs. Clipping of the plots for biomass
measurements in the fall of 2002 may have been deleterious to perennial,
non-rosette forming forbs but may have benefited winter annuals by eliminating the canopy of standing dead tallgrass. The relative influence of daylength and temperature on the phenology of winter annuals will be discussed.
SHIPLEY, BILL.* Departement de biologie, Universite de Sherbrooke,
Sherbrooke, Quebec, Canada. Translating from the language of causality
to the language of probability (and back . . .).
Ecologists invoke the notion of causality to express an asymmetric relationship between a cause (C) and its effect (E): CE. Probability theory,
the official language of statistics, cannot express such asymmetrical relationships and instead describes only symmetrical relationships between random variables. This has led to a century of confusion between the relationship between causality and statistics, and has resulted in ecologists
consistently mistranslating their causal claim (CE) as E5f(C). Here, I
give a user-friendly introduction to an alternative mathematical language
that can express asymmetrical relationships (the theory of directed graphs)
and describe the link between directed graphs and probability distributions.
I then present a new, general and robust statistical test (d-sep test) of path
models that allows one to test causal claims without physical manipulation,
and also how the same logic can be used to explore observational data and
generate causal hypotheses. I conclude by describing the relationship of
these methods to structural equation models.
SHIRK, JENNIFER L.* and MARIANNE E. KRASNY.* Cornell University, Department of Natural Resources, Fernow Hall, Ithaca, NY, USA.
Can student inquiry lead to valid data? Seeking mutual benefit in a
student-scientist partnership.
Over the past twenty years, scientific organizations have been soliciting
volunteer assistance to broaden the geographic range for monitoring studies. Simultaneously, high schools challenged to meet standards for science
inquiry seek resources in the scientific community for exposure to science
content and processes. As a result, a variety of student-scientist partnership
programs have evolved to enhance high school classroom research. While
adult-focused citizen science projects are now widely accepted and scientists utilize data collected by trained volunteers, few organizations incorporate student-collected data into monitoring reports. For partnerships between schools and scientists to be sustainable, mutual benefit must be
sought, and validating student-collected data may provide incentives for
scientists involvement with schools. Recent reports indicate that student
data can reliably be used for biophysical and biochemical studies, but student skills in biodiversity research have not been rigorously examined. This
study compares data collected by students with those collected by professional herpetologists, following identical protocols for monitoring amphibian populations. In addition, addressing school inquiry standards which call
for deeper student involvement in research than rote data collection, we
examine the effect of student participation in the design of their driving
research question on the quality of data they collect. Schools in the project
received identical protocols training, but half of the classrooms additionally
defined their own research question through a facilitated process. By interviews and observations this project investigates the effect of the inquiry
process on students demonstrated research skills. Preliminary results indicate that students who helped develop a class research question demonstrated project ownership and commitment to accurate research.

SHORTLE, JAMES S.,* ROBERT E. OCONNOR, ANN FISHER and

JAMES FINLEY. Penn State University, University Park, PA, USA. Socioeconomic dimensions of environmental indicators and their applications to coastal resource management.
Useful indicators of the condition coastal ecosystems must have ecological
validity and reliability. But they must also be meaningful to and relevant
for intended audiences. These audiences include decision makers in environmental and resource management and planning agencies, as well as
stakeholders to whom decision makers must be responsive. In this paper
we examine the implications of (1) human perceptual and attitudinal dimensions of the types of information that different audiences find useful
for documenting trends, setting priorities, and selecting management activities and (2) risk communication methods for the choice and presentation
of indicator information to environmental managers and other audiences.
The paper includes results from interviews with state and federal environmental managers in the Atlantic Slope Region to identify their use of environmental indicators and their informational needs and preferences.
SHUMAN, BRYAN N.,1,* STEPHEN T. JACKSON2 and THOMPSON
WEBB.3 1 University of Minnesota, Department of Geography, Minneapolis, MN; 2 University of Wyoming, Department of Botany, Laramie, WY;
3
Brown University, Department of Geological Sciences, Providence, RI.
Responses to Past Global Change: Biogeographic Change in the Southeastern United States over the Past 21,000 years.
Climate change since the end of the last ice age has had a significant impact
on the distribution of plant species in eastern North America. During the
last glacial maximum, 21,000 years ago, northern conifers (including one
extinct species) dominated the vegetation of the southeastern United States.
Temperate and cool-temperate hardwoods also had widespread distributions, but like endemic species, were isolated to locally suitable habitats.
Comparisons of sparse fossil pollen and plant macrofossil data from the
Southeast with independent paleoclimate data have shown that these plant
communities changed dramatically in response to subsequent climate
change. For example, an expansion of the southern species of pine (Pinus
spp.) in the southeastern United States between 10,000 and 8000 years ago
coincided closely with an increase in available moisture that also raised
lake levels along the coastal plain. However, a long-term increase in the
abundance of many taxa, particularly temperate and northern hardwoods,
requires further explanation. Abundance of these taxa increased between
17,000 and 11,000 years ago and maybe related, in part, to the simultaneous
rise in atmospheric carbon dioxide content. An important consequence was
an increase in the number of locally-dominant tree taxa (and in the evenness of abundance among tree taxa).
SHURETTE, RYAN, GEORGE CLINE and ROBERT E. CARTER.* The
effects of hardwood mid-story removal in Montane Longleaf pine (Pinus palustris) stands on breeding bird communities in the Talladega
National Forest, Alabama.
Recent research in Longleaf pine forest management has focused on the
endangered Red-cockaded woodpecker (Picoides borealis; or RCW), however the Longleaf pine ecosystem has been shown to provide critical habitat
for a number of other avian species as well. While bird communities of
low elevation and Coastal Plain Longleaf forests have been well studied,
montane Longleaf ecosystems have received little attention. In this study,
I compare preliminary results of breeding bird inventories in open montane
Longleaf stands managed for RCWs (via mechanical hardwood mid-story
removal and burning) versus unburned stands with relatively dense hardwood mid-stories. Species richness (n540, managed; n542, unmanaged)
and species diversity indices (Shannon-Weiner, H51.36 managed;
H51.41 unmanaged) were similar for both management practices. While
data are too limited to support statistical inferences, notable numbers of
breeding species in managed stands seem to be displaced by a different
community type as hardwood mid-story is reduced. Species of special concern (e.g., Aimophila aestivalis and Sitta pusilla) were observed more frequently in managed versus unmanaged stands. This study should improve
understanding of the role montane Longleaf pine ecosystems play in regard
to breeding bird communities.

SIEDSCHLAG, SARAH,* CHERYL SWIFT,* KIM SCHRUM and BRYAN STIMMLER. Whittier College, Whittier, CA. Determinants of aquatic plant cover in a Southern California river.
Humans commonly alter the San Gabriel river channel by constructing rock
dams that create pools suitable for swimming. Adjacent pools and riffles
were compared in order to determine the effect of lower current velocity
on sediment accumulation and type as well as plant cover. Transects were
established across the steam channel. Percent cover for algae and five sediment size classes and fine sediment depth were measured at meter intervals. Water surface velocity was measured by estimating the time a float
took to travel a pre-determined distance. The velocity was twice as high
in the riffles as in the pools. Very fine sediment was twice as deep and
occupied twice as much of the coverage area in pools compared to riffles.
Percent cover of the algae Chara was three times higher in pools than in
riffles. The results of this study raise the question of whether higher algae
cover is the result of increased fine sediment accumulation or whether
higher algae cover traps increased amounts of sediment leading to accumulation. Alternatively, both fine sediment accumulation and algae cover
may be independently associated with decreased current velocity. A large
proportion of the watershed above the study site burned in late summer of
2002, and three large precipitation events resulted in both substantial sediment loads in the stream channel and the increased discharge washed away
the rock dams providing an opportunity for a follow up study to further
investigate the relationship between current velocity, algae cover and fine
sediment accumulation. This study provides evidence that human use of
the San Gabriel River results in changes in fluvial geomorphology that in
turn change the habitat quality of the river.
SIEVING, KATHRYN E.* and THOMAS A. CONTRERAS. Wildlife
Ecology and Conservation, 110 Newins-Ziegler Hall, University of Florida,
Gainesville, Florida, USA. Winter flocking for foraging or predation
benefits, or both?
The relative importance of foraging and predation-related benefits is variously weighted in discussions of principal selective forces generating and
maintaining multi-species bird flocks. We hypothesize that life history characteristics related to reducing predation risk and increasing foraging efficiency are both important; specifically, that one of the factors by itself is
probably not sufficient to generate flock participatory behavior. To test our
hypothesis we compared the anti-predation and foraging traits of two
groups of understory bird species in north-central Florida; those that participate as satellite species in organized winter flocks with Eastern Tufted
Titmice (Baeolophus bicolor) vs. those species that vigorously participate
in predator-mobbing aggregations, but do not follow titmice while foraging
(35 species total). Our central prediction is that flocking and mobbing species should not be distinguishable on the basis of anti-predation related
traits (assuming they share potential predators since flock satellites also
mob), but that they would diverge in foraging-related traits (assuming that
mobbing only species do not gain foraging efficiency by flocking). Using
logistic regression, we tested for the ability of 2 predation-related variables
(body mass and escape tactic (4 categories)) and 3 foraging-related variables: foraging location (3 categories), foraging tactic (4 categories, and
diet breadth (5 categories) to predict group membership (either flock satellites or mobbing aggregations). Supporting our central prediction, foraging tactic was the principal predictor of group membership (model chisquare 5 20.15, P , 0.001; -2LLR 5 24.1), providing 92.3% correct
identification of flocking species, and 75% for mobbing-only species. Most
flock species are foliage gleaners, and most mobbing only species are
ground gleaners. Body mass was secondarily important in predicting group
membership with mobbing-only species being heavier (p,0.05) but all
birds weigh significantly less than the reported mean weights for potential
avian predators in the study region.
SIH, ANDREW. University of California-Davis, asih@ucdavis.edu, Davis,
CA. An experimental study of the predator-prey behavioral response
race.
Space use by predators and prey, and in particular, patterns of spatial overlap between predators and prey are crucial elements of the predator-prey
interaction. Numerous studies have shown that if prey are fixed in space

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307

(e.g., prey are immobile), then predators tend to forage in areas with more
prey. The outcome is a positive spatial association between predators and
prey. Conversely, numerous other studies show that if predators are spatially fixed (e.g., caged), then prey avoid areas with more predators. The
result is then a negative spatial association between the two. Surprisingly
few studies, however, have looked at the dynamics of the behavioral response race that occurs in the common, natural situation where both predators and prey can respond to the other. When should prey win the race
(resulting in a negative spatial association between the two), and when
should predators win (as indicated by a positive spatial association)? Here,
I first review the basic idea of some extant game theory on this predatorprey behavioral race, and then present experiments examining this interaction involving treefrog tadpole prey (Hyla regilla) and predatory salamander larvae (Ambystoma tigrinum). The experiments address: 1) whether
tadpoles in the absence of predators follow the matching prediction of
simple ideal free distribution; 2) whether tadpoles avoid caged predators;
and 3) the outcome of the race (i.e., the pattern of spatial association between tadpoles and salamanders) when both are free to respond to the other.
Some results corroborated existing theory; however, some other results did
not fit the predictions of existing theory. These suggest new, broader approaches to modeling the predator-prey race.
SILIM, SALIM, HARBANS L. SEHTIYA and HORMOZ BASSIRIRAD.*
University of Illinois at Chicago, Chicago, Illinois. Control of nitrogen
uptake responses to elevated CO2 in tree seedlings.
An important factor that regulates the uptake of nitrogen (N) by plants is
the physiological uptake capacity of the root system. A number of studies
have shown that kinetics of root N uptake can change dramatically in response to CO2 enrichment, but the effects are not consistent among species
or between inorganic N forms. Theoretical considerations suggest that such
inconsistencies must involve changes in: 1) supply of sugars to the root
and/or 2) internal N pools i.e., elevated CO2 elicits species-specific pattern
of responses in root carbohydrate and internal N pools. We examined the
effects of elevated CO2 (720 versus 360 Pa) on N uptake and assimilation
in seedlings of Acer saccharinum, Fraxinus pennsylvanica and Ulmus
americana. Elevated CO2 significantly increased ammonium uptake in all
species, but had little effect on nitrate uptake. Ammonium uptake increased
the most in Ulmus followed by Acer and Fraxinus. Total root and leaf
soluble carbohydrates were higher under elevated CO2. The activity of glutamine synthetase, increased significantly in roots of all species under elevated CO2 while that of nitrate reductase did not. Consequently, in all
species root ammonium concentration was significantly lowered at high
CO2. Total root amino acids and nitrate concentration did not respond to
growth CO2. From these data we conclude that ammonium uptake responses to high CO2 may have been mediated by internal root ammonium concentration and a positive control via root carbohydrate status.
SILLIMAN, BRIAN R. and MARK D. BERTNESS. Brown University,
Providence, Ri. A trophic cascade regulates salt marsh primary production: Will over-harvesting of blue crabs trigger massive die-off of
southern salt marshes?
For nearly 5 decades, the prevailing theory of salt marsh ecology has been
that bottom-up forces, such as nutrient availability, are the primary factors
regulating marsh primary production. However, experimental manipulation
of the dominant marsh grazer (the periwinkle, Littoraria irrorata) and its
predators (e.g. blue crabs) showed that top-down forces are also strong
determinants of marsh plant production. Periwinkle grazing can convert
one of the most productive grasslands in the world into a barren mudflat
within 8 months. Marine predators regulate the abundance of this plantgrazing snail. Thus, top-down control of grazer density is a key regulatory
determinant of marsh grass growth. The discovery of this simple trophic
cascade implies that over-harvesting of snail predators (e.g. blue crabs) may
be an important factor contributing to the massive die-off (10s of km2) of
salt marshes across the southeastern United States. We have surveyed and
installed exclusion cages at 4 die off areas in GA and LA (in LA with L.
Stanton and I. Mendelssohn) and found 100 m long snail fronts with
.2,500 ind./ m2 grazing down marsh grass at up to 14 m/ year. Largescale, multi-site snail tethering experiments and monitoring of predator
abundances over three years in GA marshes showed that top-down control

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Abstracts

of Littoraria decreases significantly with local blue crab declines. These

results show that (1) successful management of blue crab fisheries may be
critical for long-term health and persistence of East Coast salt marshes and
(2) the bottom-up paradigm of salt marsh ecology and its application to
other systems needs to be re-evaluated.
SILVERA, KATIA,1,2,* LOUIS S. SANTIAGO,1 EDWARD A. SCHUUR,1
NORRIS H. WILLIAMS,1 W M. WHITTEN1 and KLAUS WINTER.2
1
University of Florida, Gainesville, FL, USA; 2 Smithsonian Tropical Research Institute, Balboa, Panama, Panama. Evolution of photosynthetic
pathways in the Orchidaceae: Evidence from stable isotopes and phylogenetic analysis.
We investigated the carbon stable isotopic composition of orchids from
Panama and Costa Rica to determine if photosynthetic carbon assimilation
occurs predominantly by C3 or Crassulacean Acid Metabolism (CAM).
Carbon isotopic composition of leaf material showed a bimodal distribution
with most species exhibiting values near 227, suggesting a C3 pathway,
or around 212, suggesting a CAM pathway. Leaf thickness was related to
photosynthetic pathway. Thicker leaves often signified a predominantly
CAM pathway, whereas thinner leaves were related to C3 photosynthesis.
When incorporated into the orchid phylogeny, the distribution of photosynthetic pathways suggests that C3 is the ancestral state and that CAM
evolved more than once in the Orchidaceae. This analysis is part of a larger
project in which approximately 1600 species will be analyzed from living
and preserved specimens for a greater understanding of the evolution of
photosynthetic pathways in orchids.
SIMMONS, JEFFREY A.,1,* ERIN M. LAWRENCE1 and THOMAS G.
JONES.2 1 West Virginia Wesleyan College, Buckhannon, WV; 2 Marshall
University, Huntington, WV. Acid mine drainage effects on stream ecosystem health in West Virginia.
Acid mine drainage (AMD) from coal mining affects approximately 6,500
miles of streams in the Appalachian region. Raw or untreated AMD is
highly acidic and contains elevated concentrations of Fe, Mn, Al and other
heavy metals. Currently much of the AMD discharged from active mining
operations is first treated to raise the pH and reduce concentrations of heavy
metals. The effects of raw AMD on biotic diversity and leaf decomposition
have been studied but impacts of treated AMD on streams has been overlooked. Our objective was to determine if discharges of treated AMD affected the health of stream ecosystems. We used an index of biological
diversity as well as two measures of ecosystem function to assess ecosystem health. Four streams from each of three categories were selected randomly from a GIS database of West Virginia stream reaches. AMD streams
had inputs of untreated acid mine drainage, Treated Streams had inputs of
treated AMD and Reference streams had no inputs of acid mine drainage.
From each stream water samples were collected on three dates for chemical
analysis. Periphyton growth on ceramic tiles and benthic macroinvertebrate
diversity was measured in each stream. Leaf decomposition bags were incubated for six weeks in Reference and Treated streams. Chemical analysis
of stream water verified that the streams were placed in the correct category. Periphyton growth was significantly reduced in AMD streams
(p,0.05), however Treated streams were no different from Reference
streams. In terms of macroinvertebrates AMD streams had significantly
fewer taxa, lower diversity index and fewer EPT taxa than the other stream
types (p,0.05). Treated streams had less than half the number of taxa
compared to Reference streams (p,0.05). Leaf decomposition was significantly slower in Treated streams than in Reference streams (p,0.05).
Thus, although treated AMD is much less toxic than raw AMD, it still has
substantial impacts on species diversity and leaf decomposition thereby
reducing ecosystem health.
SIMMONS, STEVEN P.,* DIANNE V. OWEN and JOHN C. VOLIN.
Florida Atlantic University, Davie, FL. Implications of water velocity for
phosphorus capture by Stormwater Treatment Areas in the Florida
Everglades.
Artificial wetlands are increasingly being used as filters of wastewater with
high nutrient loads. This is especially true in Florida, where Everglades
restoration includes construction of six Stormwater Treatment Areas

(STAs) totaling 16,000 hectares. The primary objective of this research

was to investigate the effects of water movement on phosphorus uptake by
periphyton. Our hypothesis was that phosphorus uptake by a periphytondominated system would correlate positively with water velocity. We tested
this hypothesis within constructed mesocosms that allowed us to test the
sole effect of flow rate, keeping other factors, such as hydraulic retention
time and hydraulic loading rate, equal across treatments. Treatment water
velocities were approximately 0.22 cm s-1 and 2.0 cm -1, which are achievable levels within an STA. The mesocosms were installed at the outlet of
an STA and thus received water from which much of the biologically
available phosphorus had already been assimilated. After twelve weeks of
treatment, biomass in the high flow rate was 20% higher than that in the
low flow rate. Plant tissue phosphorus did not differ significantly between
the two treatments. The greater biomass accrual in the fast treatment therefore represents a greater removal of phosphorus from the water column.
Information obtained from this study can be used to optimize water movement within STAs, leading to greater operational efficiencies and a lower
final concentration of phosphorus in water discharged to the Everglades.
SIMONSON, SARA E.,1,* DAVID T. BARNETT,1 THOMAS J. STOHLGREN1,2 and MICHAEL IELMINI.3 1 Natural Resource Ecology Laboratory, Colorado State University, Fort Collins, CO; 2 Fort Collins Science
Center, U.S. Geological Survey, Fort Collins, CO; 3 National Wildlife Refuge System, U.S. Fish and Wildlife Service, Arlington, VA. Invasion of
the National Wildlife Refuges.
We report great success in Phase I of the National Invasive Species Survey,
an electronic questionnaire of all federally designated U.S. Fish and Wildlife Service National Wildlife Refuges. We conducted this survey to assess
the current status and management of invasive plants, non-indigenous animals and emerging diseases of plants and animals throughout the refuge
system. We summarize the responses of managers, administrators and biologists from more than 440 national wildlife refuges (representing almost
90% of the organizational units with direct land management responsibilities). This initial snapshot of the survey results is based on the 670 + nonnative plants, non-indigenous animals and emerging diseases reported by
refuge managers, biologists and administrators. These refuges include an
impressive diversity of terrestrial and aquatic habitats and range in size
from 0.5 acres to over 26 million acres. The survey results will be available
to the refuge system through on-line databases with custom search capabilities for information at local, regional and national scales. An invasive
species web portal will contribute to improved support for management
action on the refuges and informed decision-making at local, regional and
national levels. National-scale datasets will be combined with the refuge
system information to encourage early detection of invading species and
improve prediction of vulnerable habitats. Dataset integration is an important first step in preventing harmful effects of future invasions in the
worlds largest network of public lands and waters set aside specifically
for the protection of wildlife.
SIMPSON, COLLEEN M.* and MARY L. REID. Department of Biological Sciences, University of Calgary, Calgary, Alberta, Canada. When a
tree falls in the forest: Resource concentration after forest management
benefits bark beetles.
The resource concentration hypothesis (sensu Root 1973) predicts that herbivores will be most abundant in areas of high host species concentration
(large patches of low vegetational diversity). Here, we apply the resource
concentration hypothesis to explain increased pine engraver bark beetle (Ips
pini (Say); Scolytidae) abundance in thinned stands, as compared to natural
stands. Forest thinning, the selective harvest of a proportion of trees, results
in changes to the biotic and abiotic characteristics of the remaining stand.
After thinning, stands become more open, evenly spaced, and dominated
by a single tree species. Increased wind speeds in thinned stands may
contribute to the observed increase in input of freshly downed trees. We
found pine engravers to be more abundant in thinned lodgepole pine stands,
even up to seven years after thinning harvest. The resource concentration
effects of thinning (greater input of host material with less non-host material) is a novel explanation for the increased abundance of secondary
bark beetles in thinned boreal forest stands.

SIMS, DANIEL A.,1,* WALTER C. OECHEL,2 HONGYANG LUO2 and

JOHN A. GAMON.1 1 California State University, Los Angeles, CA; 2 San
Diego State University, San Diego, CA. Relationships between vegetation greenness and ecosystem CO2 flux in southern California chaparral: A test of the functional convergence hypothesis.
Correlations between vegetation greenness and CO2 flux have been found
across a wide range of ecosystems. This observation led to the functional
convergence hypothesis stating that capacity for CO2 fixation should be
curtailed whenever a limitation in the availability of any resource prevents
the efficient exploitation of additional capacity. Remote sensing data used
to support the functional convergence hypothesis have all been gathered
over long time periods (one year or more) and across diverse ecosystems.
In this study, we tested the functional convergence hypothesis within one
ecosystem (southern California chaparral) and collected spectral reflectance
data at monthly intervals. Ecosystem CO2 flux was measured continuously
with the eddy covariance technique. In order to better match the temporal
and spatial scales of the spectral reflectance measurements to those of the
eddy flux, we installed an automated tram system within the eddy flux
tower footprint. This system measured hyperspectral reflectance over a 100
m transect throughout the diurnal cycle. Since we observed large effects
of diurnal and seasonal variation in solar elevation angle on NDVI, we
defined the inherent vegetation greenness as NDVI at a solar elevation
angle of 40 degrees. Large seasonal and inter-annual variation in ecosystem
carbon flux rates, resulting from annual summer droughts and the severe
drought of 2002, provided a strong test of the relationships between CO2
flux and vegetation greenness. CO2 flux was strongly correlated with NDVI
at a constant solar elevation angle (i.e. vegetation greenness) but was only
weakly correlated with NDVI measured at noon. In contrast to previous
studies, this correlation between CO2 flux and vegetation greenness did not
imply a constant light use efficiency (LUE) of the vegetation. In fact LUE
varied widely, approaching values associated with crop plants during favorable environmental conditions and dropping to zero during the severe
drought. Our results suggest that it may be possible to predict CO2 flux
directly from vegetation greenness without any assumptions about LUE.
SINGER, MICHAEL S.* and YVES CARRIERE. University of Arizona,
Tucson, AZ. Anti-parasitoid defense explains host-plant selection by a
polyphagous caterpillar.
Ecological as well as physiological factors may determine host-plant selection by phytophagous insects. Experimentally separating these factors
can be difficult with specialist herbivores that gain ecological and physiological benefits from a single host. In this study, we use the individually
polyphagous caterpillar Grammia geneura (Lepidoptera: Arctiidae) to separate performance and defensive benefits from different host-plant species.
Previous work showed that certain single host species diets (e.g. Malva
parviflora) enhanced performance and that other single host species diets
(e.g. Ambrosia confertiflora) enhanced resistance against parasitoids. Here,
we specifically hypothesized that this caterpillars natural habit of feeding
on a mixture of toxic plant species provides resistance against parasitoids
at a performance cost. We collected late instar caterpillars from the field,
and, assuming some proportion was already parasitized, randomly assigned
them among two diet treatments for the duration of development in the
laboratory. Caterpillars reared on Malva suffered 46% mortality from parasitoids, while those reared on a mixture of Malva, Ambrosia, and Senecio
longilobus suffered only 26% mortality from parasitoids. More importantly,
the probability of mortality from parasitoids was negatively associated with
the proportion of toxic plants (Ambrosia and Senecio) in the diet of individual caterpillars.
SKELLY, DAVID K.* Yale University, New Haven, CT. Microgeographic
divergence among amphibian populations.
Prior studies have shown that macrogeographic gradients in temperature
associated with latitude and altitude can lead to countergradient patterns of
variation in a number of taxa; individuals from colder environments are
known to grow or develop faster than their conspecifics from warmer environments when placed in a common setting. In this study, I hypothesized
that countergradient variation also is important at microgeographic scales.
The wood frog, Rana sylvatica, breeds in open-canopied, temporary wet-

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309

lands as well as those heavily shaded by vegetation. Shading leads to cooler

thermal environments that are associated with embryonic development
rates as much as 50% slower than those in unshaded wetlands. Wetlands
with contrasting canopy environments are often found within tens or hundreds of meters of each other. In a common garden experiment, embryos
from nearby natural wetlands displayed countergradient variation: individuals collected from shaded wetlands developed up to 12% faster than those
collected from relatively unshaded wetlands. The results of this study suggest that the concept of countergradient variation may be extended to small
scales of space. In addition, the rate and scale of vegetation dynamics (the
agent of wetland shading) imply that divergence in development among
residents of nearby wetlands may be relatively rapid, on the order of decades.

and Alabama are scale-dependent and differ with changing spatial resolution and community extent. Fishes and macroinvertebrates were sampled
three times from May to September 2000 and sampling effort among channel types was equal. We examined trends of mean taxa richness, evenness,
and abundance among channel types and how they varied among three
spatial resolutions (5 m patches, 30 m sections, and 90 m sites) and three
community extents (fish, macroinvertebrate, and combined fish and macroinvertebrate communities). Trends of community structure among channel types differed with changing spatial resolution and community extent.
The results supported our hypothesis and suggested that the scale of assessment is important when evaluating impacts of stream alteration projects
on stream communities.

SLONE, DANIEL H.,1,* SUSAN V. GRUNER2 and JON C. ALLEN.3

1
USDA Forest Service, Southern Research Station, Pineville, LA; 2 University of Florida, Gainesville, FL; 3 University of California, Santa Barbara, Santa Barbara, CA. Estimating time-of-death with a community
model of maggot growth and development.

SMITH, ANNEMARIE,* RALPH E. BOERNER and JENNIFER A.

BRINKMAN. Ohio State University, Columbus, OH, 43210. Effects of
forest ecosystem restoration treatments on soil chemical and biochemical properties.

A model of maggot growth and develoment is presented that estimates the

post-mortem interval (PMI) for criminal homicides or other suspicious
deaths. Building on a previous, simpler model, we have expanded it to
include communities of several maggot species. Validation and error rates
of the model are presented via comparison to field trials of pig carcasses
where the PMI for each sample was precisely known.
SMEE, DELBERT L.* and MARC J. WEISSBURG. School of Biology
Georgia Institute of Technology, Atlanta, GA, USA. Claming Up: Mercenaria Reduce Apparancy to Whelk Predators Via Waterborne
Chemical Cues.
Immobile organisms minimize predation risk by reducing their apparency
from, or defending themselves against, consumers. Hard clams, Mercenaria
mercenaria, are heavily preyed upon by both crabs and gastropods, and
Mercenaria predators locate their prey by following waterborne odor
plumes released by clams while they feed. Previous experiments have
shown that clams grow more slowly in habitats where predation levels are
high, suggesting that hard clams may reduce their apparency to predators
by ceasing to feed. Clam feeding (pumping) behavior was measured in the
presence and absence of blue crab Callinectes sapidus and knobbed whelk
Busycon carica predators in multiple flow regimes. Our results indicate
that clam pumping behavior was significantly reduced in the presence of
both blue crabs and knobbed whelks. Pumping was suppressed to the same
degree when clams were exposed to predator exudates compared to exposure to the predators themselves, which indicates that clams are detecting
their predators via a waterborne chemical cue. The effects of blue crab
presence on clam pumping were reduced in turbulent flows when the crabs
were placed 1 m away, but clam responses to whelks remained similar in
all flow conditions. Clam perceptual distance of crabs was reduced by
turbulence, and clam feeding was suppressed less by crab exudates than
whelk exudates. The stronger suppression of feeding by whelks may be
indicative of the greater threat this predator poses to clams. Whelks are
able to follow odor plumes in turbulent flow conditions whereas blue crabs
are not. Therefore, a blue crab 1 m away poses less threat to a clam in a
turbulent flow. Since crabs move much faster than whelks, brief interruptions in pumping would be sufficient to disguise clams from crabs.
SMILEY, PETER C.* and ERIC D. DIBBLE. Mississippi State University,
Mississippi State, Mississippi, USA. Influence of spatial resolution and
community extent on patterns of community structure within a channelized warmwater stream in Mississippi and Alabama.
It is a central concept in ecology that patterns documented by research are
scale dependent. Yet, scale (extent and resolution) is frequently overlooked
in applied ecology and natural resource management. Studies assessing the
influence of stream alteration projects on stream communities typically examine patterns of community structure among coarse-grain habitat categories without considering if the observed patterns are scale-dependent. We
investigated the hypothesis that patterns of community structure among
three channel types within a channelized warmwater stream in Mississippi

310

Abstracts

Structural restoration based on thinning from below to pretreatment tree

density and basal area, functional restoration based on reintroduction of
dormant season fire and a combination of the two were each applied to
forested areas of 15-30 ha each in three study sites within the oak-hickory
(Quercus-Carya) region of southern Ohio. Soils sampled from permanent
sampling plots were taken in the summers of 2000 (prior to treatment) and
2001 (after winter thinning and spring burning). Nonmetric multidimensional scaling (NMS) of 26 soil chemical and biochemical parameters in
pre- and post-treatment samples revealed two major gradients of variation:
one which described differences among the three study sites and one which
was based on the differences between pre- and post-treatment years. The
separation along this second gradient appeared generally similar in the untreated controls and in the various treatment plots, suggesting that actual
treatment effects were somewhat confounded by year-to-year variation.
Analysis of variance confirmed the importance of among-sites and between-years variance components, and also revealed significant effects of
fire on N mineralization rate and acid phosphatase activity. Thinning affected available Al and phenol oxidase activity, and the combination of
thinning and burning affected soil [C], soil [N], and N mineralization rate.
Ordinations such as NMS that are tailored to data with lognormal distributions (as is the case with many soil properties) can reveal subtle shifts
in soil conditions which may be ecologically meaningful.
SMITH, BRUCE N.,1,* THIMMAPPA ANEKONDA,2 CLAYTON A.
JONES,1 EMILY A. KELLER1 and LEE D. HANSEN.1 1 Brigham Young
University, Provo, Utah, USA; 2 Oregon State Univeresity, Corvallis,
Oregon, USA. Genetic variation in carbon isotope discrimination, respiratory metabolism, growth, bud phenology, and their interrelationships among seedlings of two Douglas-fir varieties and their F2 hybrid
progeny.
In a common garden study, seedlings of Psuedotsuga menziessii (Douglasfir) var. menziessii (coastal) from Lacomb, Oregon and var. glauca (interior)
from Clearwater National Forest, Idaho and their F2 hybrids were grown
in nursery beds in order to understand the genetic variability in growth and
phenology traits of segregating hybrids relative to their parents. Height,
stem diameter, and bud burst percent were measured in May 1999. Metabolic heat rate and respiration rate were measured on the apical meristems
at 30, 35, and 40oC. The tissue was then dried overnight in a vacuum oven,
ground, combusted and analyzed for carbon isotopic ratios. The coastal
variety was from an elevation of 245 m with a mean annual rainfall of
1,400 mm. The interior variety was from an elevation of 871 m and a mean
annual rainfall of 600 mm. The two varieties differed from one another in
growth traits, bud burst, carbon isotopic values and respiration measurements. The F2 hybrid progeny had similar isotopic values to the interior
variety and differed significantly from the coastal variety. However respiration measurements of the hybrids were similar to the coastal variety. Few
trait differences between families were noted. The fast growing coastal
variety showed the least carbon isotope discrimination relative to the slow
growing and presumably more stressed interior variety.

SMITH, CURTIS A.* and WILLIAM G. WILSON. Duke University, Durham, North Carolina. Evolution of dispersal distance in a predator-prey
system: Living on the edge of criticality.
This study explores the dynamics associated with a spatial predator-prey
model in which the dispersal distance of the predator is allowed to evolve
from an initial value significantly greater than the dispersal distance of the
prey. Spatial instabilities quickly lead to a patchy landscape, soon followed
by a steady decrease in mean predator dispersal distance to values just
greater than the dispersal distance of the prey. Depending on mutation rate
and the size of the landscape, the predator persists on the edge of criticality
as a metapopulation. Local extinctions of the predator are followed by the
unimpeded growth of prey patches until the recolonization of predators
results in the large patch either shrinking in size or splitting into numerous
smaller patches. The growth and subsequent splitting of patches provides
a potential mechanism for the recolonization of areas long void of predators, while the occasional merging of large patches prior to recolonization
allows for gene flow between prey populations long separated.
SMITH, DAVID L.,1,2 JEREMY J. BURDON3 and LARS ERICSON.4
1
University of Maryland School of Medicine, Baltimore, MD; 2 National
Institutes of Health, Bethesda, MD; 3 CSIRO Plant Industry, Centre for
Plant Biodiversity Research, Canberra, Australia; 4 Umea University,
Umea, Sweden. When to stop a study: Analysis from an 11-year study
of a parasite metapopulation.
Decisions to continue or terminate studies that have lasted more than a
decade must be based on many factors. One critical factor is the expectation
that the effort will produce additional insights and knowledge. We recently
completed analysis of data from an 11-year study of a parasite metapopulation. The parasite is a macrocyclic rust Triphragmium ulmariae, a foliar
pathogen of a perennial plant Filipendula ulmaria. The host plants are
distributed in 186 island shore populations in the Skeppsvik Archipelago,
northern Sweden. We developed a set of models based on demographic
and environmental factors, and used maximum likelihood and model selection criteria to select a most parsimonious model. We found evidence
for epidemiological patterns at multiple spatial scales. In this study, we
repeated the analysis for subsets of the data to ask what we would have
found if the study had been shorter. We extrapolate from the data to speculate about what will be gained if the study continues. We discuss the
benefits and costs of continuing against the possible value of alternative
courses of action. Possibilities include sampling less frequently, using the
study to establish baseline rates as a precursor to a quasi-experiment, or
abandoning the study.
SMITH, DENA M.* CU Museum and Department of Geological Sciences,
UCB 265, University of Colorado, Boulder, CO, USA. Eocene Insect Herbivory and Climate Change.
Do changes in levels of insect herbivory correspond to changes in climate?
Modern latitudinal trends in insect herbivory and patterns from the early
Cenozoic suggest that climate may be an important factor mediating plantinsect associations. I compare the herbivore damage on fossil leaves from
both the 46Ma (Middle Eocene) Green River Formation at Douglas Pass,
Colorado and the 34Ma (Late Eocene) Florissant Flora of Colorado. These
lacustrine fossil localities occur during the Eocene - Oligocene cooling
trend. The intensity of herbivore damage associated with these fossil assemblages was measured by examining the percentage of leaf area removed
by insect herbivores, the number of leaves damaged in the floras, the number of feeding guilds on individual leaves in the floras, and the overall
feeding guild structure. Levels of herbivory declined as temperature declined during the Eocene - Oligocene cooling trend. There was not a significant change in the number of feeding guilds found attacking leaves or
in the overall feeding guild structure, but there was a slight increase in the
proportion of insect galls from the middle to the late Eocene. There is also
evidence of long-lived plant-herbivore associations that span this time interval.

SMITH, DIXIE L.,1 ROXANE FAGAN2,* and LORETTA C. JOHNSON.2

Pittsburg State University, Pittsburg, Kansas, USA; 2 Kansas State University, Manhattan, Kansas, USA. Changes in soil organic carbon cycling
as juniper forests invade prairie: A d13C isotope approach.
1

Woody plant expansion into grasslands has been occurring across the Great
Plains. The purpose of this study was to investigate changes in d13C dynamics in bulk soils, soil particle size-fractions, and in CO2 from soils
collected from 40-60 year old closed-canopy juniper (Juniperus virginiana)
forest and paired C4 grasslands. We hypothesized that a lifeform shift to
juniper will alter quality, quantity, and distribution of SOC. We quantified
the vegetation origin and cycling of new forest C and the vegetation substrate for microbial respiration by analyzing the d13C of soils and respired
CO2. d13C of SOC in forest soils indicates a large input of juniper-derived
C to 10cm. At 0-2.5cm, 38% of SOC of the bulk soil is juniper in origin.
In contrast, the majority of SOC in bulk soil below 10cm is prairie-derived,
residual carbon. Forest C is present in all size fractions but prevails in the
.212micron fraction. In grassland soils, d13C of respired CO2 generally
reflects the isotopic composition of bulk soil. In contrast, the d13C of respired CO2 from 0-10cm depth in forest soils initially reflected the microbial utilization of new juniper humus (-25) from bulk soil. Thus, microbes preferentially utilized recent, C3-derived juniper SOC. In summary,
d13C data from bulk soils, particle size fractions, and microbial incubations
confirm that fundamental vegetation shifts are altering the quality, and distribution of SOC.
SMITH, ELIZABETH A.,1,* MIQUEL A. GONZALEZ-MELER,1 ROSER
MATAMALA,2 JULIE JASTROW2 and RICHARD M. MILLER.2 1 University of Illinois @ Chicago, Chicago, IL, USA; 2 Argonne National Laboratory, Argonne, IL, USA. Carbon dynamics of a restored tallgrass
prairie chronosequence assessed with stable isotopes.
Restoration of agricultural fields back to native vegetation has the potential
to serve as a sink for atmospheric carbon and therefore mitigate the effects
of elevated CO2. At FermiLab (Batavia, IL), tallgrass prairies have been
sequentially planted since the mid 1970s establishing a chronosequence in
which soil carbon dynamics can be studied. Plant communities consist of
forbs, shrubs, and C3 and C4 grasses, and the relative dominance of vegetation type depends on time of the season and restoration age. The different isotopic 13C signatures of C3 and C4 plant tissues are used to determine the C3 and C4 origin of belowground carbon dynamics. Soil respiration, root composition, and soil organic matter were measured for their
carbon content as well as their d13C signatures for three restored tallgrass
prairies that vary in restoration age from 3 to 25 years of age and compared
to a C3 pasture and a remnant prairie (not cultivated for over 500 yrs).
Throughout the chronosequence, root biomass and soil respiration increased
coinciding with increased plant cover and C4 vegetation dominance in
older plots. Fractions of SOM show an increase in C3 carbon in the stable
clay fraction. Our results suggest that in soils of a restored tallgrass prairie,
carbon of C3 origin has more potential for carbon sequestration than carbon
of C4 origin. Implications for carbon sequestration are discussed.
SMITH, FREDERICK W.,1 LEIGH B. LENTILE1 and WAYNE SHEPPERD.2 1 Colorado State University, Ft. Collins, CO; 2 USDA Forest Service, Rocky Mountain Research Station, Ft. Collins, CO. Direct Fire Effects following the Jasper Fire, Black Hills National Forest, SD.
In late August 2000, the Jasper Fire burned 33,000 ha of the Black Hills
in South Dakota. The Jasper Fire burned under a variety of vegetative,
topographic, and meteorological conditions creating a mosaic of vegetative
mortality in patches of varying size and extent. 27, 48, and 25 % of the
fire burned under high, moderate, and low severity conditions. Within the
context of a larger monitoring design, we provide an initial characterization
of severity by quantifying direct fire effects on the overstory and the forest
floor. We observed an increase in the proportion of crown, bole, and forest
floor affected by fire along a severity gradient from low to high. In low
and moderate severity treatments, 21-70% of the crown was scorched. In
high severity treatments, 92% of the crown was consumed. We indexed
fire severity as the product of the proportion of the ground area charred
and the severity of char scaled from low (1) to high (3). Fire severity was
120 for low and 245 for high treatments on a scale from 100-300. Forest

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311

floor litter depths decreased from 2.54 cm in unburned stands to 0.15 cm

in high severity treatments. Litter mass decreased from 1265 g m2 in unburned areas to 82 g m2 in high intensity treatments. Tree mortality due to
direct fire effects increased from low to high fire severity. Fire selectively
killed smaller trees in low and moderate severity treatments. However, no
trees, independent of diameter, survived in high severity treatments. This
observation suggests that a threshold may exist between diameter and fire
survivability in low and moderate severity fires in ponderosa pine systems
in the Black Hills.
SMITH, JAMES L.* and J DANIEL HARE. University of California Riverside, Riverside, CA. Gas exchange and water use efficiency of
glandular and non-glandular trichome types in Datura wrightii (Solanaceae).
The maintenance of a trichome dimorphism in Datura wrightii in light of
the apparent fitness cost of producing glandular trichomes suggests that
there could be one or more trait-associated life history characteristics that
are beneficial. Sticky plants (glandular) occur more frequently than velvety
plants (non-glandular when mature) in populations located in more mesic
habitats. This suggests that water use and availability may play an important part in determining the relative distribution of sticky and velvety
plants. Leaf hairs have been shown in other species to significantly reduce
absorbed solar radiation, temperature, transpiration rate (E), and photosynthetic CO2 assimilation rate (A). Gas exchange parameters of sticky and
velvety plants grown in a common garden were measured during midday
on several dates in late summer and early fall of 2002. Water use efficiency
(WUE; A/E) was 11.2% higher on average in sticky plants, however there
was an interaction between type and date. On hot, sunny days there was
no difference in WUE, but on cooler, less sunny days WUE was up to
25.0% higher in sticky plants. This difference in WUE cannot be attributed
to stomatal conductance or E, which did not differ significantly between
types; rather, it appears to be due to fluctuations in A, which varied up to
22.1% higher in sticky plants than in velvety plants. Higher WUE in sticky
plants at lower temperature and light intensity is most likely due to physiological or anatomical differences associated with trichome type rather
than any direct effect (shading/evaporation) of the trichomes themselves.
In relatively mesic habitats with lower temperatures and less intense solar
radiation, sticky plants may have a physiological advantage that could
translate into more rapid growth and competitive ability.
SMITH, JAPHIA* and ROGER C. ANDERSON. Illinois State University,
Normal, IL. Historical change and future predictions of vegetation
structure in a central Illinois woodland.
Historically, many Illinois upland woodlands were dominated by shadeintolerant oak and hickory species that required periodic fires for their
maintenance. Fire cessation associated with European settlement permitted
aggressive invasion of these forests by fire sensitive but shade-tolerant sugar maples, which have displaced other native tree species. This vegetation
change is displayed by our study site, the 13 ha Thaddeus Stubblefield
Illinois Nature Preserve in Funks Grove, IL. Changes on the site are documented by historic tree data from Government Land Office (GLO) Survey
Records, and tree and sapling data collected in 1977, 1984, 1994, and 2001.
Based on GLO records, the leading trees in the original grove were white
and red oak, ash, and elm. In 1977, the leading tree species was American
elm, which was replaced by sugar maple as the leading species in the 1984
sample and in all subsequent samples. Tree diversity as measured by the
Shannon-Wiener Index has progressively declined from historic times. A
comparison between the diversity of the tree stratum and the sapling stratum over time indicates increasing similarity between strata while diversity
decreases overall. We also examined the impact of loss of the elms on the
diversity of the forest. STANDSIM, a stochastic forest model, was used to
make predictions about future vegetation change based on current and historical data.
SMITH, MARIAN,1,* HAL CASWELL2 and PAIGE METTLER.3 1 Southern Illinois University, Edwardsville, IL; 2 Woods Hole Oceanographic Institution, Woods Hole, MA; 3 Southern Illinois University, Carbondale, IL.
A stochastic model of the threatened plant Boltonia decurrens in a
dynamic environment.
Boltonia decurrens, a threatened plant that is restricted to the Illinois River
Valley, has a complex life cycle that evolved in response to the dynamics

312

Abstracts

of the historic flood regime. In the past 100 years, hydrological changes
accompanying the regulation of the river have resulted in a dramatic decline
in abundance. We developed deterministic and stochastic demographic
models to explore the causes of this decline. Our analysis indicates that
hydrology affects vital rates and population growth. Elasticity analysis
shows that changes in the timing of spring floods effect a change from an
annual life cycle to biennial or perennial cycles. On a year-to-year basis,
flood timing fluctuates randomly, and our stochastic models show that the
frequency of late-receding floods dramatically reduces the stochastic
growth rate log ls. Flood timing interacts with precipitation levels during
the growing season, and we developed a stochastic model including both
factors to document their effects on log ls. Historical data from 1895-2000
indicate that prior to the 1930s, when levees and navigation dams were
completed, the frequency of late-receding floods was much lower than it
is now. Stochastic analysis, characterizing the environment by pre- and
post-1930s flood and precipitation regimes, links the effects of changes in
the historical regime to population demographics and life cycle characteristics. Life for B. decurrens has become more uncertain: log l s declines,
s2 increases dramatically and the probability of quasi-extinction rises as
the disturbance regime changes after 1900. This is the first time that a
stochastic model has been coupled with historical data on a stochastic environment. Undoubtedly, many plant and animal species evolved in concert
with dynamic habitats and are now threatened by anthropogenic changes
in those dynamics. The data and analyses used in this study can be applied
to management and development strategies to preserve other dynamic systems.
SMITH, MELINDA D.1,* and SCOTT L. COLLINS.2 1 National Center for
Ecological Analysis & Synthesis, Santa Barbara, CA; 2 University of New
Mexico, Albuquerque, NM. Scale-dependence of the effects of fire and
grazing on spatial and temporal heterogeneity in tallgrass prairie.
Natural disturbances affect spatial and temporal heterogeneity in plant communities, but these effects may depend on the type of disturbance or spatial
scale of analysis. In this study, we examined the effects of fire frequency
(1-, 4- and 20-yr intervals) and grazing by bison on spatial and temporal
heterogeneity in tallgrass prairie plant communities. At three spatial scales
(10-, 50-, and 200-m2), we used Euclidean Distance between plots and
between all time steps (1993-2001) for each plot as measures of spatial
and temporal heterogeneity, respectively. At the 10-m2 and 50-m2 scales,
spatial heterogeneity was lowest on annually burned sites and highest on
unburned (20-yr) sites. Fire frequency had no significant effects on heterogeneity at the largest spatial scale. In contrast, grazing significantly reduced
spatial heterogeneity at 200-m2, but had no effect on spatial heterogeneity
at smaller spatial scales. The rate of community change over time (i.e, slope
of temporal heterogeneity vs. time lag) decreased with increasing fire frequency at all scales of analysis, but this pattern was only significant at 50m2. Temporal heterogeneity was not affected by grazing. Our results show
that fire and grazing have differential effects on spatial and temporal heterogeneity and that these effects are scale-dependent. These patterns reflect
the relatively uniform impact of fire at all spatial scales, and the patchy
impact of grazing in both burned and unburned prairie at larger spatial
scales.
SMITH, RACHEL N.* and LEE A. VIERLING. South Dakota School of
Mines and Technology, Rapid City, South Dakota. Improved ground- and
satellite-based methods for deriving LAI in a burned ponderosa pine
ecosystem.
The purpose of our study is to conduct a series of field measurements and
data analyses to investigate the accuracy and various potential improvements of leaf are index (LAI) derivations from satellite sensors over a
ponderosa pine dominated forest. Permanent field sites were established
within and around the 33,800 hectare Jasper fire (burned September 2000),
in the southwestern Black Hills of South Dakota. Spatial variation in fire
severity resulted in a heterogeneous mosaic of canopy LAI and therefore
sites were selected to represent a continuum of LAI values. To account for
the nonrandom nature of ponderosa pine stands and for the effects of
clumping at various scales, correction factors were applied to the effective
LAI values obtained from a LAI-2000 Plant Canopy Analyzer and hemispherical photos. Clumping indices from the needle and shoot scales were

applied from previous LAI studies conducted on ponderosa pine in Oregon.

A Tracing Radiation and Architecture (TRAC) instrument was used to correct for clumping at scales larger than shoot. Corrected and uncorrected
LAI values were then regressed against spectral vegetation indices (SVIs)
derived from IKONOS, Landsat ETM+, and MODIS data. Uncorrected
effective LAI values were better correlated with SVIs (e.g. r2 5 0.51 for
IKONOS NDVI comparisons) than after the LAI values were corrected for
clumping (r2 5 0.18), indicating the need for site specific clumping correction indices. To address this issue, we explored the use of ground-based
Light Detection and Ranging (LiDAR) data as a substitute for destructive
sampling to derive clumping indices at the needle and shoot scales.

bees pollinated sunflowers in fields with varying abundance and diversity

of native bees. We found that when native bee abundance and diversity
were low, honey bees were poor pollinators, with a single visit yielding
only, on average, about 2 seeds. However, when native bees were highly
abundant and diverse, honey bee pollinating ability increased an order of
magnitude, to about 20 seeds per visit.

SMITH, RICHARD G.1,2,* and KATHERINE L. GROSS.1,2 1 W. K. Kellogg

Biological Station, Hickory Corners, Michigan, USA; 2 Michigan State
University, East Lansing, Michigan, USA. Multivariate analysis of weed
community response to a long-term management and rotation experiment.

The extensive floating periphyton mat and its associated invertebrate community are an integral part of the Everglades food web. No published
studies have directly sampled or characterized this community, so its distribution and response to variations in the physical environment are poorly
understood. In this study we sought to describe the response of both the
floating periphyton mat and benthic macroinvertebrate communities to nutrient enrichment and variation in hydroperiod. Here we sampled sites at
two levels of phosphorus enrichment (ambient and enriched) and three
hydroperiods ("short": ,200 days since last dry-down; "long": .400 days
since last dry-down; and "very long": .4000 days since last dry-down) in
a factorial design. This was replicated in two different geographic areas of
the Everglades separated by water control structures ("very long" sites were
only available in one area). We found that community composition varied
with both enrichment and variation of hydroperiod, and that these effects
were different in floating mat and benthic communities. We noted almost
a three-fold increase in the total density of invertebrates in the floating mat
with enrichment. We also observed a significant interaction between hydroperiod and enrichment. When short hydroperiod sites were enriched,
they experienced almost a six-fold increase in the total number of invertebrates, with more than twice as many invertebrates as any other enriched
sites. We feel that this dramatic increase in invertebrates at short hydroperiod, enriched sites may be the result of compounding top-down and
bottom-up trophic interactions. Understanding this interaction and gaining
a deeper understanding of the associated community dynamics will enable
researchers to better identify trophic interactions and determine the impact
of environmental stressors on this system.

Weed communities under annual row crop agricultural systems are the
product of interactions between agronomic management techniques and
environmental factors that act as filters on the potential local species pool.
Organic management and crop rotation are two alternative agricultural
strategies aimed at decreasing reliance on external chemical inputs such as
synthetic fertilizers, pesticides, and herbicides. These management strategies may also result in predictable shifts in weed community composition
and structure by differentially affecting the recruitment and survival of
species from the potential species pool. In this study we examined how the
soil seed bank in the tenth year, and emergent weed community structure
in the tenth and eleventh years differed across treatments of a long-term
crop rotation and management experiment at the Kellogg Biological Station
(KBS) in Southwest Michigan. We used non-metric multidimensional scaling (NMS) and multi-response permutation procedures (MRPP) to identify
weed species groups associated with corn grown under two management
(organic vs. conventional high input) and rotation (continuous vs. corncorn-soybean-wheat) treatments. Indicator species analysis was employed
to identify significant associations between particular weed species and
treatments. Weed community composition and structure were sensitive to
treatment in both years and resulted in significant grouping of treatments
in species space. Indicator species analysis on seed bank densities and
aboveground biomass showed Poa compressa was strongly associated with
high-input continuous corn, in both the seed bank and emergent community. Three other grasses, Panicum dichotomiflorum, Setaria viridis, and
Digitaria sanguinalis, were strongly associated with high-input corn when
grown in a three crop rotation. In contrast, an annual forb, Chenopodium
album was associated with organic continuous corn treatments.
SMITH, SARAH A.* and CLAIRE KREMEN. Princeton University,
Princeton, NJ. Native, free-living bees increase sunflower production by
providing direct pollination and assisting honey bees.
Pollination services is an important ecosystem service provided by native,
free-living bees. Sunflower growers rent honey bees, often ignoring the
benefits they may receive from nearby natural areas and farm management
practices that support native bees. Our research was conducted in Yolo
County, CA, where sunflowers are grown commercially for hybrid seed.
We determined which bees visit sunflower, how they contribute to sunflower pollination, and which factors affect their distribution and abundance.
To determine which bees visit sunflowers and how they are distributed
across the landscape, in our 3-year project, we documented more than
30,000 bee visits on over 15 fields across the landscape. We found bees
from at least 25 species visiting sunflower. We used multiple regression
analyses with bee distribution as the independent variable and 3 dependent
variables: isolation from natural habitat, pesticide use, and history of sunflowers being planted nearby. We found that some species appear to rely
on areas of nearby natural habitat while other species were linked to sunflowers being planted in the same area for many consecutive years. We
examined the direct contribution of native bees to sunflower pollination
using single visit studies, in which we allowed one bee to visit one sunflower head and excluded all other pollinators. We found high variation in
the pollinating abilities of native bees, with single visits from some bee
species resulting in over 20 seeds. We also investigated how well honey

SMITH, SHAWN E.L.* and JOEL C. TREXLER. Department of Biological Sciences, Florida International University, Miami, Florida, USA. Macroinvertebrates in the Florida Everglades: Community-level observations of the effects of hydroperiod and nutrient enrichment.

SMITH, WINSTON P.,* SCOTT M. GENDE and JEFFREY V. NICHOLS.

Ecological correlates of flying squirrel microhabitat use and density in
temperate rainforests of southeastern Alaska.
The northern flying squirrel was proposed as an indicator of biological
diversity in coniferous forests of the Pacific Northwest. We studied habitat
relations of the Prince of Wales flying squirrel (Glaucomys sabrinus griseifrons), an endemic of temperate, coniferous rainforest of southeastern
Alaska, because of concerns over biological diversity from extensive clearcut logging in the region. We used stepwise logistic regression (SLR) to
examine relationships between microhabitat use and individual habitat features (measured variables) and habitat factors, which were independent linear combinations of individual habitat variables identified in factor analysis.
Seasonal (spring, autumn) models were created for upland, old-growth (upland-OG) western hemlock (Tsuga heterophylla)/Sitka spruce (Picea sitchensis) and peatland mixed-conifer forests (peatland-MC). Density of trees
with diameter-at-breast-height (dbh) .74 cm and abundance of Vaccinium
were positively correlated with microhabitat use in peatland-MC during
both seasons with the odds of capturing a flying squirrel increasing by
factors of 2.7 and 16.9 during spring and autumn, respectively, with an
increase in density of 10 trees/ha. Microhabitat use of upland-OG during
autumn was related to Vaccinium ground cover and density of snags with
a dbh of 50-74 cm; microhabitat use during spring was inversely correlated
with percent surface cover of water. SLR models with multivariate factors
performed poorer in predicting capture sites than models of individual habitat variables, indicating that habitat factors were not more thorough in
explaining variation in habitat use than individual habitat elements within
the scope of conditions examined. At the stand level, large (.74 cm) trees
explained 65% of the variation in density between habitats; percent cover
of moss and downed wood in decay classes I-IV explained 70% and 6377% of the variation, respectively. Our results corroborate some general
patterns reported for western coniferous forests, but suggest that northern

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313

flying squirrels in temperate rainforests of southeastern Alaska differ ecologically from populations in the Pacific Northwest in important ways that
likely influence their suitability as management indicators.

SNETSINGER, CAROL* and CAROL BREWER. Division of Biological

Science, University of Montana, Missoula, MT, USA. Integrating computer technology into biology instruction.

Recent technological advances have provided us with tools and techniques

that are essential to the way we do biology today. Clearly, our professional
cultures are tool-rich and have been profoundly affected by these technologies. But has this technological revolution transformed our courses as
well? Computer-based technologies (IT) can be powerful pedagogical tools,
and turn the passive recipient of information into an active participant in
the learning. This requires buy-in by both students and faculty. Through a
4-yr biology curriculum innovation program at the University of Montana,
we have examined the extent to which faculty use IT in their teaching.
Today, most biology faculty surveyed have course web pages (90%) and
use power point presentations (62%). Interestingly, the majority of faculty
(.60%) felt that IT can create opportunities for students learning that
would be impossible without it, but fewer than 15% had used technology
to promote interaction amongst students or for real-time assessment to receive immediate feedback on student understanding. All of the respondents
agreed that using computers proficiently is essential to being a successful
professional biologist, but many fewer (41%) agreed that these skills currently are an integral part of the courses they teach. Faculty and students
had different views on student preparation. While 76% of introductory
biology students were very or totally confident they had the basic computer
skills necessary to be successful in a biology course, only 15% of faculty
agreed that incoming freshman had the computer skills they needed. To
make better use of computers in instruction, our results suggest that faculty
need time, opportunities for professional development, and reliable hardware and software in the classroom. Finally, choosing the content to teach
is the easy part; figuring out how to use computer technology to help
students understand and master the content is the time intensive challenge.

SNYDER, KEIRITH A.,1,* TRAVIS E. HUXMAN,2 DAVID D. BRESHEARS,3 BRADFORD P. WILCOX,4 ERIC E. SMALL,5 RUSSELL L.
SCOTT,6 ROB B. JACKSON7 and KEVIN R. HULTINE.2 1 USDA-ARS
Jornada Experimental Range, Las Cruces, NM; 2 University of Arizona,
Tucson, AZ; 3 Los Alamos National Laboratory, Los Alamos, NM; 4 Texas
A&M University, College Station, TX; 5 University of Colorado - Boulder,
Boulder, CO; 6 USDA-ARS Southwest Watershed Research Center, Tucson,
AZ; 7 Duke University, Durham, NC. An ecohydrological framework for
determining the landscape effects of woody plant encroachment.
Woody plant encroachment into semiarid and arid systems is a global phenomenon with large potential hydrological and biogeochemical consequences. We considered systems in the southwestern United States (pinyonjuniper, mesquite and creosote), where dramatic changes in physiognomy
create potential feedbacks between woody plants and hydrology at the landscape scale. The outcome is a new conceptual framework useful in directing
future research and determining broad scale potential ecological and biogeochemical consequences across a range of ecosystems. Shrub removal
has been proposed as mechanism for increasing water yields, but increased
water yields are likely only in systems with shallow bedrock, riparian systems, or mesic systems where precipitation exceeds potential evapotranspiration. In semiarid and arid systems abiotic controls on evaporation overwhelm biotic control of streamflow. In these systems, the predominant effects of woody plant encroachment that promote ecohydrological feedbacks
are changes in: infiltration, depth of plant water removal, generation of
transient overland flow and near-ground energy budgets that affect the ratio
of evaporation (E) to transpiration (T). In these more xeric areas, the ratio
of E:T is critical because it indicates the biological use of water which
influences (through feed-forward or feed-back effects) landscape structure.
Increasing woody plant cover is associated with larger open interspaces, at
the landscape scale the differential contribution of canopy versus interspaces may change landscape E:T. The alterations of spatial structure and
ecohydrology likely influence carbon cycling. At the landscape scale, total
respiration depends on the magnitude of canopy/interspace respiration flux
scaled by the spatial extent of interspaces and canopy space. Canopy respiration is correlated to T, while interspace respiration is largely driven by
soil water availability and therefore is driven by factors that control E.
Resolving the partitioning of E:T should help resolve the net effect of
woody plant encroachment on other biogeochemical cycles at the landscape
scale.

SNOVER, MELISSA L.1,2,* and GEORGE WATTERS.1 1 Center for Stock

Assessment Research, University of California - Santa Cruz, Santa Cruz,
CA, USA; 2 NOAA/NMFS/Pacific Fisheries Environmental Lab, Pacific
Grove, CA, 93950. Age at maturity and maximum reproductive fitness
in male Coho salmon (Onchorhynchus kisutch).

SNYDER, ROBIN E.* and PETER L. CHESSON. Section of Evolution

and Ecology, University of California, Davis, CA, USA. How the spatial
scales of dispersal, competition, and environmental heterogeneity interact to alter species coexistence.

Male coho salmon adopt one of two mating strategies, fighting or sneaking.
Fighters defend a nest from fertilization by other males while sneakers hide
in proximity to a females nest and sneak fertilizations. Typically, sneakers
mature early, spending only six months at sea while fighters mature a year
later. We have developed an individual growth model for coho salmon. We
used a form of the von Bertalanffy curve where net linear growth rates are
described by E-kL: E relates to the rate of anabolism, k is the growth
coefficient and relates to the rate of catabolism and L is length. Making
the assumption that k is set at an early point in an individuals development
and remains constant throughout life, faster growth in freshwater, or higher
k, results in larger smolts, For seawater growth, we assume that all smolts
from the same source experience approximately the same ocean conditions
and food availability, therefore the E term is constant. Hence, net linear
seawater growth rates decrease with larger values of k. Larger smolts have
a decreased capacity for growth and cannot attain large sizes, which we
consider an ultimate causation of early maturation. We then developed fitness functions for two possible ages at maturity, two and three years. The
fitness functions assume that increasing size conveys a higher likelihood
of mating success for fighters, with the opposite being true for sneakers.
The fitness functions demonstrate that large smolts maximize reproductive
fitness by maturing early and using the sneaking strategy. All other smolt
sizes maximize fitness by maturing at 3 years and using the fighting strategy. This work points to mechanisms that offer ultimate and proximate
explanations for early maturation in coho salmon.

Spatial mechanisms for coexistence depend on biological and physical processes occurring at a number of spatial scales, and coexistence may be
promoted or suppressed as the relative sizes of these scales shift. We use
a model of plant competition in a spatially varying environment to show
how shifting scales of dispersal, competition, and environmental heterogeneity affect coexistence. Spatial coexistence mechanisms are partitioned
into three types: the storage effect, nonlinear competition, and growthdensity covariance. We then follow a two-tiered approach, first expressing
the strength of each of these mechanisms in terms of covariances between
population densities and environmental quality and then explaining how
changes in the scales of dispersal, competition, and environmental heterogeneity should affect the covariances. For example, nonlinear competition
is related to the variances of resident density and resident environment and
their covariance. It promotes coexistence more strongly as the competitively dominant species disperses over shorter distances and as members
of the competitively dominant species compete with themselves over longer
distances and with inferior competitors over shorter distances. Our approach allows us to quantify how important different scales of environmental heterogeneity are for coexistence. We also enable investigations at
two different levels: researchers may measure the relevant covariances directly (in computer simulations or in field studies) and look for their effects
on coexistence or they may investigate how the spatial scales of biological
and physical processes determine these covariances. Having the key components of our theory measurable both theoretically and empirically facilitates testing at both of these levels.

314

Abstracts

SOJA, AMBER J.,1,2,* HERMAN H. SHUGART,1 PAUL W. STACKHOUSE,2 ANATOLY SUKHININ3 and SUSAN CONARD.4 1 University
of Virginia, Charlottesville, Va, USA; 2 NASA Langley, Hampton, Va; 3 Sukachev Institute of Forestry, Krasnoyarsk, Siberia, Russia; 4 USDA Forest
Service, Arlington, Va. Investigating potential climate-induced change
in fire return intervals for 1999, 2000, and 2001 in boreal Siberia.
Evidence of fire-induced, climate-related change is investigated in boreal
Siberia by comparing calculated fire return intervals with expected fire
return intervals. Boreal regions are particularly significant because these
are the regions that will initially experience climate change and this is
where the largest reservoir of terrestrial carbon resides. Under current climate change scenarios, fire season length and fire weather severity are
expected to increase. Because temperatures have already increased across
Siberia in the last decades, it follows that the interval between fire occurrence at particular sites is expected to decrease. The hypothesis to be tested
is fire return intervals will decrease. Satellite-based data is used to estimate
area burned, which is overlaid on an ecoregion map to calculate area burned
in each ecoregion across Siberia. Data from 1999, 2000, and 2001 are used
to calculate average fire return intervals for each ecosystem, as well as an
average boreal forest fire return interval. Results from this investigation
show that fire return intervals are either equivalent to or greater than expected fire return intervals, which was not expected. However, both an
underestimation in area burned and the minimal number of years analyzed
could result in anomalous fire return intervals. Interestingly, the expected
percentage of young stands, which is calculated based on the estimated
average boreal forest fire return interval (159 years), is similar to published
estimates of the percentage of existing young stands (6.5%).
SOLOMESHCH, AYZIK I. and MICHAEL G. BARBOUR. * University of California, Davis, CA, USA; 2 Institute of Biology Russian Academy
of Science, Ufa, Russia. Reconstruction of California native grasslands.
1,2

1,

California interior grassland covering about 25% of the state is dominated

by non-native annuals and represents a dramatic example of large-scale
species replacement. The change over the past 250 years occurred so rapidly, and so early, that no descriptions exist for their pre-contact botanical
composition. Range of alternative opinions includes chaparral, broadleaved
forest, annual grasses, and rhizomatous grasses. Nevertheless a widely accepted paradigm is that they were originally dominated by perennial bunchgrasses Nassella cernua and N. pulchra, associated with half a dozen other
perennial grasses (Clements 1920). But because of the pervasive extent of
exotics, the composition of pre-contact Californian grasslands has never
been reconstructed with any degree of assurance (Hamilton 1997). A resolution of this uncertainty would have enormous restoration and conservation importance. We assume it has remained so long unsolved because
of a past focus on dominant species, which have been completely replaced
and may not be used as a strong evidence for any of expressed opinions.
We test the bunchgrass hypothesis analyzing the complete floristic assemblages of 55 modern grasslands sampled in Sacramento area, giving particular emphasis to native species that are now typically neither abundant
nor dominant. It was shown, that despite the low abundance of native
species, their contribution to a-diversity is much greater (r50.87) than the
statistically insignificant contribution of exotics (r50.11). Both detrended
correspondence analysis and classification of stands according to BraunBlanquet approach confirms that the five groups of plots were floristically
distinct. Ratio between native and exotic species and life form spectra differ
from stand to stand, indicating degree of species replacement. Nassella
pulchra among others native species exhibited significant positive correlations (r50.320) with total cover of invasive species. Bunchgrass paradigm does not explain variety of native species assemblages in modern
grasslands. Our results support the alternative hypothesis, that it was a
heterogeneous and complex mosaic, with floristically different communities
occurring on an array of different microhabitats at a relatively fine scale.
SOMMERKORN, MARTIN,1,* KNUTE J. NADELHOFFER,1 LORETTA
C. JOHNSON,2 GEORGE W. KLING3 and EDWARD B. RASTETTER.1
1
The Macaulay Institute, Craigiebuckler, Aberdeen, UK; 2 Kansas State
University, Division of Biology, Manhattan, KS, USA; 3 University of
Michigan, Dept. of Ecology & Evolutionary Biology, Ann Arbor, MI, USA.
Carbon and nitrogen allocation of tundra plants assessed through an
ecosystem scale dual tracer experiment.
Plant allocation determines the location, timing, and magnitude of inputs
into ecosystem carbon (C) and nitrogen (N) pathways. In tundra systems,

plant belowground production generally exceeds aboveground production.

The fate of aboveground production differs greatly from that of belowground production. Plant allocation patterns can therefore have strong influence on the regulation of ecosystem biogeochemistry. To examine allocation patterns and controls on the fate of carbon allocated belowground,
we labeled 1m2, paired control and chronically fertilized wet sedge tundra
plots with 15N and 14C on three dates during the growing season. On the
fertilized plots, nine years of chronic N+P additions had altered the relative
abundances of species and had increased aboveground plant biomass by a
factor of 3, belowground biomass by a factor of 2. Through isotope analysis
of harvested plant material and of respired CO2, we found that 7 days after
labeling, control plots had lost 60-75%, fertilized plots 30-40%, of the
assimilated C as CO2, indicating higher growth efficiency of fertilized plots.
Root production ranged from 0.1-0.5 gCm-2d-1 and was higher in control
plots than in fertilized plots. Root production peaked mid season in fertilized plots and late season in control plots, whereas leaf production of fertilized plots lagged behind that of control plots. At mid season, fertilized
plants allocated 8 times more C to leaves and 20 times more to rhizomes
compared to control plants. Fertilized plants were able to take up 5-8 times
more N compared to control plants. They also allocated 7-9 times more N
to rhizomes and up to 24 times more N to leaves compared to control
plants. We conclude that chronically increased nitrogen availability shifts
allocations patterns away from nutrient acquiring tissues and toward C
acquiring tissues. The increase in N availability also enhances allocation
toward storage and vegetative reproduction.
SOMMERVILLE, GERICKE L.* and MICHAEL B. COUGENOUR. Colorado State University, gericke@nrel.colostate.edu, mikec@nrel.colstate.
edu, Fort Collins, CO, USA. Spatial patterning of elephant impact and
woodland dynamics in an East African savanna.
East African savannas are unique, dynamic ecosystems, which tend to shift
in woody cover and composition over time. The major factors influencing
these dynamics are still under investigation, and elephants are considered
to be a keystone species for restructuring woody habitat. In Serengeti National Park, we employed a spatially-explicit sampling design to assess the
characteristics of elephant impact in relation to patch quality (species composition, stand structure, patch size, etc) across the landscape. Elephant
impact was assessed by measuring the cross-sectional area of broken
branches and then converting to biomass. Impact, or biomass removal,
exhibited two scales of pattern with stand structure at the small scale (85m,
I 5 0.1319) and species type at the large scale (1 km, I 5 -0.1454). A
spatial autoregressive model explained approximatley 85% of the variation
across the landscape by regressing impact iteratively for each scale of pattern (species composition at the 1km scale, R2 5 0.6193, p50 and tree
density at the 85m scale, R2 5 0.6021, p50). Additionally, tree density
showed a feedback correlation with species composition, the strength of
which was affected by the magnitude of elephant impact. The SAVANNA
model was used to model these dynamics and the interaction with fire.
Preliminary results indicate that elephant impact suppresses preferred, fireintolerant species into the flame zone whereas avoided species tend to be
fire-tolerant and survive to replace the canopy.
SONG, BO,1,* HARBIN LI2,* and WILLIAM E. FULL.3 1 Baruch Institute
of Coastal Ecology and Forest Science, P.O. Box 596 Clemson Unviersity,
Georgetown, SC 29442-0596, USA; 2 Center for Forested Wetlands Research, USDA Forest Service Southern Research Station, 2730 Savannah
Highway, Charleston, SC 29414, USA; 3 Baruch Institute of Coastal Ecology and Forest Science, P.O. Box 596 Clemson Unviersity, Georgetown,
SC 29442-0596, USA. Effects of forest management on bird distribution and dynamics in the Hobcaw Barony of South Carolina.
Hobcaw Barony is located at the southern tip of the Waccamaw Peninsula
near Georgetown, South Carolina. Hobcaw Barony was set up as the
17,500-acre wildlife refuge, where the main tree species include longleaf
pine, loblolly pine, cypress, sweetgum, and the main bird species include
white turkey, and red cocked woodpecker. The forest management includes
repeated logging and prescribed burning. Coupling the forest dynamics
under forest management with the survey of the Audubons Bird Count, a
landscape model, LEEMATH was used to estimate the bird dynamics and
their distribution patterns under forest management across the Hobcaw Bar-

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315

ony landscape. The major inputs include: a) GIS Vegetation map, road map,
and stream network; b) Forest inventory data; and c)Habitat matrix. The
simulation result has shown that the spatial habitat for red cocked woodpeckers decreased, which may have resulted from Hurricane Hugo in 1989.
SORRIE, BRUCE A. and ALAN S. WEAKLEY.* University of North
Carolina Herbarium / NC Botanical Garden, Chapel Hill, NC, USA. Developing a blueprint for conservation of the longleaf ecosystem based
on centers of endemism.
In recent decades there has been a surge in interest in the conservation of
longleaf pine savannas and associated fire-maintained ecosystems of the
Southeastern United States Coastal Plain, as their extent and the condition
of remaining examples has continued to decline. There has also been an
increased appreciation of the high community and species diversity of these
systems, and the large number of endemic species associated with them.
Conservation strategies must take into account the landscape integrity of
potential conservation areas and the concomitant likelihood that land managers will be able to maintain natural or semi-natural fire regimes. Conservation strategies must also take into account the distribution patterns of
the many narrowly endemic species if the goal of conserving the systems
and all of their component species is to be met. We present an analysis of
geographic patterns of plant endemism in the Southeastern Coastal Plain.
Of over 1600 vascular plant taxa endemic to the Southeastern Coastal Plain,
about 900 are directly associated with longleaf pine savannas. These endemics are distributed non-randomly, and distinct centers of endemism can
be identified and mapped. Among the best developed of these are 1. the
Apalachicola Lowlands of Florida and nearby portions of the the Florida
Panhandle and adjacent Georgia and Alabama, 2. various parts of the Florida Peninsula, 3. the Cape Fear Arch area of southeastern North Carolina
and adjacent South Carolina, and 4. the West Gulf Coastal Plain of Louisiana and eastern Texas, though other areas have substantial numbers of
narrow endemics as well. A comparison of these areas of endemism with
conserved longleaf pine ecosystems on public and private lands suggests
that conservation will need to rely on existing and new conservation lands,
and that a successful strategy will require small (but manageable) preserves
as well as larger landscape-scale conservation areas.
SOSNOSKIE, LYNN M.,* CATHERINE P. HERMS and JOHN CARDINA. The Ohio State University, OARDC, Wooster, OH. Species composition of the weed seedbank with varying crop and soil management.
We characterized weed density, species diversity, and community composition in the soil seedbank in a long-term study with three crop sequences
(continuous corn, corn-soybean, corn-oat-hay) and three tillage systems
(conventional-, minimum- and no-tillage). We identified and counted germinable seeds in the top 10 cm of soil in early spring (1997-1999) to
calculate seed density, species diversity indices, and a synthetic relative
importance index for each species. Repeated measures ANOVA showed
that total seed density differed with crop sequence and tillage system, with
an interaction among these factors and years. Seed density was higher in
the continuous corn than the other crop sequences (two of three years), and
higher in no-tillage than other tillage systems. There were more species in
the corn-oat-hay sequence than in corn-soybean or continuous corn, and
species diversity declined with increasing soil disturbance. Canonical discriminant analysis showed that the first axis explained 40 to 60% of the
within-subjects variation for species composition and was strongly associated with crop sequence. Canonical scores for the plots planted to cornoat-hay clustered separately from those in continuous corn and corn-soybean. These trends were statistically significant according to Mahalanobis
squared distances and were confirmed with cluster analysis. Tillage systems
did not show a similar degree of separation, suggesting that crop rotation
was more important in influencing community composition. After one cycle
of crop rotation using glyphosate-tolerant corn and soybean varieties, these
patterns of community composition were no longer apparent, suggesting
that the seedbank responded quickly to altered management. Weed control
and other cultural practices in the corn-oat-hay system favored species with
life-history characteristics that differ from species more commonly associated with corn and soybean systems.

316

Abstracts

SOTKA, ERIK* and STEPHEN PALUMBI. Hopkins Marine Station Stanford University, Pacific Grove, CA. Wont you be my "neighbor"?
Genetic estimates of dispersal in a rocky intertidal barnacle.
The assumption that marine populations are interconnected over enormous
spatial scales is based in part on the potential of long-lived pelagic larvae
to disperse 100s of kilometers when entrained in streaming boundary currents (e.g., California Current). However, the spatial spread of pelagic larvae will be far less extensive if small-scale features of nearshore oceanography (e.g., eddies) prevent larvae from entering boundary currents. Using DNA sequence data from mitochondrial and nuclear loci, we inferred
patterns of dispersal among populations of the acorn barnacle Balanus
glandula along exposed coastlines of Washington, Oregon and California.
In some regions, alongshore dispersal distances are smaller than commonly
assumed by one to several orders of magnitude. Thus, the replenishment
of local populations of this and perhaps other marine spawners on exposed
coastlines may sometimes depend on the dynamics of populations within
restricted geographic distances. Continental coastlines are not completely
"open" systems; rather, in some places, the coastline is split into "neighborhoods" that barnacles and their offspring rarely vacate.
SOUTER, NICHOLAS J.,1,* C. MICHAEL BULL1 and MARK N.
HUTCHINSON.2 1 Flinders University of South Australia, Adelaide, South
Australia, Australia; 2 South Australian Museum, Adelaide, South Australia, Australia. Adding burrows to enhance a population of the endangered pygmy blue tongue lizard, Tiliqua adelaidensis.
The endangered pygmy blue tongue lizard, Tiliqua adelaidensis, occupies
spider burrows in a population near Burra, South Australia. In each of 12
20 x 20 m plots at that population, we added 36 artificial burrows, providing about a ten fold increase in suitable burrows for lizards. Over three
surveys during the spring and summer of 2001 02 there were significant
increases in lizard numbers in the experimental plots relative to the controls, both for adult lizards and for new recruits after clutches were produced. This local increase in population density may be due to lizards
locating suitable burrows more easily where burrow numbers were supplemented. The increased availability of high quality burrows may also reduce
mortality among lizards searching for suitable burrows. Additional burrows
also led to an increase in local density of a burrow dwelling centipede
that is a potential predator of lizards, but there was no evidence of predation, and any negative predation impact was outweighed by the beneficial
effect of providing more burrows.
SOUTHER, REBECCA F.,* RICHARD A. GOYER and GERALD J. LENHARD. Louisiana State University, Department of Entomology, Baton
Rouge, LA, USA. The interactive effects of nutrient augmentation and
insect herbivory in Louisiana swamps.
Insect-tree health relationships in Louisiana swamps were investigated in
the context of current field situations and future management tactics. Currently, many swamps are in decline, and reconstruction of the natural hydrologic regime (diverting fresh, nutrient-rich waters from the Mississippi
River) into degraded wetlands is planned or underway. A field study across
different forest densities was designed to evaluate the interactive effects of
nutrients and lepidopteran herbivory by the baldcypress leafroller, Archips
goyerana, and the forest tent caterpillar, Malacosoma disstria on baldcypress (Taxodium disstichum) and water tupelo (Nyssa aquatica), respectively. The second year of nutrient augmentation to mature trees in swamps
showed that fertilizer application (mimicking loading rates of the Mississippi River) increased nitrogen levels in the leaves and resulted in higher
basal area growth than unfertilized trees. Growth of baldcypress was higher
than tupelo in the intermediate and sparsely forested sites (the more degraded sites), but not at the highest density site (the healthiest of the three
sites), where growth was similar between the species. Nutrient content of
leaves (phosphorous and nitrogen) was positively correlated with tree density among the sites. Tupelo had a higher leaf nitrogen content than baldcypress. The nutrient content of BCLR insect droppings (frass) contained
as much as six times more nitrates than the FTC. Fertilizer application also
increased total nitrogen and phosphorous in the leaves and total nitrogen
content of frass. However, frass nitrate and ammonia (nitrogenous forms
causing eutrification) did not increase when trees were fertilized. Wetland

tree-insect complexes are proving to be sinks for these eutrifying compounds, a major focus of river diversions. Nitrogen loading from diverted,
Mississippi River water may enhance productivity of these two ecologically
important tree species.

SPILLER, DAVID A.1 and ANURAG A. AGRAWAL.2 1 University of California, Davis, Davis, CA; 2 University of Toronto, Toronto, ON, Canada.
Intense disturbance enhances plant susceptibility to herbivory: Natural
and experimental evidence.

SPENCE CHERUVELIL, KENDRA,1,* NANCY A. NATE,1,2 PATRICIA

A. SORANNO1 and MARY T. BREMIGAN.1 1 Michigan State University,
East Lansing, MI, spencek1@msu.edu; 2 Wisconsin Department of Natural
Resources, Bureau of Fisheries Management and Habitat Protection, Madison, WI. A field-test of the unimodal relationship between fish growth
and macrophyte cover in lakes.

Following Hurricane Lili, which passed directly over the site of our ongoing study in Great Exuma, Bahamas, herbivory increased on devastated
islands exposed to the storm surge but not on protected islands. Observations suggested that foliage sprouting on severely damaged shrubs was
more susceptible to herbivores than new foliage on undamaged shrubs. To
test this hypothesis we conducted a controlled field experiment: hurricane
damage was simulated by pruning shrubs on replicated islands. Seven
months after the manipulation herbivory was 68% higher on pruned shrubs
than on controls. Leaf size and percent nitrogen were higher and leaf toughness and trichome density were lower on pruned shrubs than on controls.
The experimental results indicate that enhanced herbivory on exposed islands following Hurricane Lili was caused, at least in part, by increased
susceptibility of the sprouted foliage to herbivorous arthropods. Since
sprouting occurs on many woody plants following natural or anthropogenic
disturbances, the findings in this study may have broad implications.

Macrophytes serve a dual role for fish in lakes. They provide a substrate
for macroinvertebrate populations, which are the primary food source for
many young fish, and they provide refuge from predators. Therefore, many
foodweb interactions among fish are mediated by macrophytes. Since the
influential paper by Crowder and Cooper in 1982, the idea accepted by
both researchers and fisheries managers is that there is an optimal intermediate macrophyte cover for fish growth and foraging in lakes. Some
small-scale experimental studies of largemouth bass and bluegill have
found that bass foraging success is poor at high levels of macrophyte cover.
In addition, although many southern U.S. reservoir studies have shown that
macrophyte cover .10% promotes young-of-year bass abundance, moderate cover levels (40-60%) have been correlated with poor growth and
low piscivory, and overall variability has been quite high. This high variability, and the fact that few whole-lake tests of this relationship have been
conducted in north temperature lakes, has resulted in no clearly defined
optimal range of macrophyte cover for bass and bluegill growth. Therefore,
we performed a field test of this idea using 39 lakes in Michigan, U.S. We
sampled macrophyte cover at the whole-lake scale in 2001 and 2002 (range
of cover 5 18-84%), and we assessed fish growth by calculating length at
age for largemouth bass and bluegill using existing datasets from the MI
Department of Natural Resources collected in the 1990s. Relationships between fish growth for each age class and macrophyte cover were analyzed
for the 39 lakes using nonlinear and linear regression. We did not find
strong evidence to support the idea of an optimal intermediate macrophyte
cover for largemouth bass or bluegill growth rates. However, we did find
that for bluegill ages 5-8, there were significant negative relationships between growth and macrophyte cover.
SPICER, RACHEL* and NOEL M. HOLBROOK. Harvard University,
Cambridge, MA. Effects of within-stem gas concentrations on sapwood
physiology: Implications for sapwood aging and senescence.
The gas composition within woody stems is significantly lower in O2 and
higher in CO2 than the ambient atmosphere, but it is not known how these
native concentrations affect respiration of living xylem tissue (i.e., parenchyma cells), or might contribute to the process of sapwood senescence.
We designed a series of experiments to determine (a) the pattern of O2
contents at different radial depths within woody stems, (b) rates of cellular
respiration under the reduced O2 found in stems, and (c) the potential for
elevated CO2 within stems to inhibit respiration. Species studied included
two conifers (Tsuga canadensis, Pinus strobus), two ring-porous angiosperms (Quercus rubra, Fraxinus americana) and one diffuse-porous angiosperm (Acer rubrum). Oxygen concentrations were not consistent with
a hypothesis of parenchyma death by prolonged anoxia: in most species,
O2 contents in the innermost sapwood (adjacent to the heartwood, in which
all parenchyma are dead) were typically between 10% and 12% (mole
fraction), and rarely as low as 5%. When equilibrated to an atmosphere of
10% O2, rates of sapwood respiration were similar across species, with the
innermost sapwood often, but not always, respiring at a significantly lower
rate than the outer sapwood. In contrast, when respiration was expressed
per live tissue (i.e., parenchyma) volume, conifers had higher rates of respiration than angiosperms, and showed no difference between inner and
outer sapwood. Tissue age was not a good predictor of sapwood respiration,
with tissue in the innermost sapwood respiring at the same rate across
several species despite differences in age of more than 30 years. In contrast,
a build-up of CO2 in the sapwood may inhibit respiration: in Pinus strobus,
equilibration to an atmosphere of 10% CO2 caused a 25% reduction in
respiration. Future work will focus on seasonal changes in both O2 and
CO2 concentrations within stems.

SPINELLI, GIANFRANCO1,* and MONICA MEINI.2 1 University of Eastern Piedmont, Novara, NO, Italy; 2 University of Florence, Firenze, FI,
Italy. Human pressure in fragile environments: The case of the European Mediterranean coast.
The European Mediterranean coast has always been a strongly humanized
area, thanks to the well known processes of demographic, political and
economic growth which have involved it along the centuries. During the
last century this process has also implied phenomena of change in land use
and population distribution, with special effects on coastal belts and islands,
which are particularly fragile environments. Besides usual driving forces
(industrialisation, urbanisation, traffic growth) tourism has become in the
last few decades the major factor of change and human pressure. The main
objective of our research is to assess the contribution of these driving forces
and elaborate a synthetic index of human pressure. As regards the methods
used, four main elements have been considered (demographic dynamics,
industrial development, tourism and accessibility), of which the most suitable statistical indicators have been identified and properly weighed, in
order to build the synthetic index. The application has concerned the coastal
provinces of the Republic of Croatia, which seemed particularly interesting
because of the long coastline (5790 km) and the great number of islands.
The highest values were found, besides in the main urban areas, also in
many islands of the archipelago. The paper is aimed at showing the results
of the methodology used, which should be suitable for application to other
cases.
SPONSELLER, RYAN A.,* JILL R. WELTER and STUART G. FISHER.
Arizona State University, Tempe, AZ. Soil organic matter storage in a
Sonoran Desert landscape: Patterns according to scale.
We describe patterns in soil organic matter (OM) storage in the upper
Sonoran Desert, and ask how the processes responsible for patterns change
with scale. In particular, we seek to identify the scales at which stream
features become important predictors for observed patterns in soil properties. To this end, soil OM was quantified at 3 spatial-scales: (1) a 50 x 100
m upland plot, (2) a subcatchment, with sites organized along a gradient
of stream intermittency, and (3) the entire drainage basin. Soils were sampled at two depths (0-2, and 2-12 cm), from beneath dominant plant species
and from inter-plant spaces. At the plot-scale, OM storage (at 0-2 cm) was
greater under plants than in bare locations, though significant differences
in storage were observed among plant species. In some cases, storage under
plants corresponded closely to plant size. For example, canopy volume
explained 75% of the variation in soil OM under Mesquite (Prosopis velutina). No differences in OM storage at 2-12 cm depth were found among
vegetated and bare locations. At the subcatchment-scale, soil OM storage
increased with stream permanence. Within sites, however, soil OM decreased laterally from streamside terraces to ridge-tops; the magnitude of
this lateral change varied with landscape position. Within the subcatchment,
patterns in soil OM corresponded to changes in vegetation size and production. Finally, at the basin-scale, patterns in soil OM storage related to

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317

variation in subcatchment size and geomorphic structure. In summary, for

the upper Sonoran Desert, fertility-island models adequately describe patterns in soil OM at local-scales. At broad-scales, however, patterns in soil
OM are the consequence of stream networks and geomorphic structures
that direct the movement of water across the landscape, and govern spatial
and temporal patterns in water availability.
SPRINGER, CLINT J.,1,* EVAN H. DELUCIA2 and RICHARD B.
THOMAS.1 1 West Virginia University, Morgantown, WV; 2 University of
Illinois at Urbana-Champaign, Urbana, IL. Stimulation of GPP by CO2
enrichment in the Duke Forest FACE Experiment estimated using
PnET-Day.
The objective of the Duke Forest Free-Air-CO2-Enrichment (FACE) experiment is to predict the response of forest ecosystems to future concentrations of atmospheric carbon dioxide. The Duke Forest FACE site, located
in a loblolly pine forest near Chapel Hill, NC, consists of three control
FACE rings (ambient) and three treatment FACE rings that are fumigated
with CO2 enriched air (ambient+200 ml l-1 CO2). Here we estimate annual
gross primary productivity (GPP) of ambient and elevated CO2 rings over
the 6 years of CO2 treatment using a process based model, PnET-Day.
PnET-Day is a daily time step model which uses the relationship between
foliar N concentration and maximum photosynthesis (Amax-N relationships)
to calculate canopy photosynthesis. We parameterized the PnET-Day model
with (Amax-N relationships measured through the canopy of both ambient
and elevated CO2 rings at the Duke Forest FACE site. The model was also
parameterized with site-specific climate and vegetation variables to adjust
canopy photosynthesis to estimate GPP. Mean annual GPP of the ambient
grown forest was 2274 g C m-2 yr-1 while annual GPP of the elevated forest
was 2586 g C m-2 yr-1. This represents a 14% stimulation of mean annual
GPP by elevated CO2 over the 6 years of CO2 treatment and is in close
agreement with the 18% stimulation of annual GPP by elevated CO2 determined using biometric techniques at the Duke Forest FACE site.
SPROTT, PATRICIA L.* LTER Network Office, University of New Mexico, Albuquerque, NM, USA. From research to classroom: How LTER
integrates research into education.
Over time and across disciplines and ecosystems, scientists and educators
involved in the Long Term Ecological Research Network (LTER) have
developed many innovative tools and techniques for integrating research
with learning activities for students and teachers at all levels. This poster
will feature specific examples of how research and teaching come together
through LTER. In these broad based, multi-faceted programs, scientists
interface with graduate and undergraduate students, and educators and students from elementary, middle and high schools to provide deeper knowledge of how science is done, and how hands-on science is best integrated
in their schools. This poster will illustrate the details of how educators in
the LTER Network, work across sites, integrating field and laboratory techniques with education.
SRIVASTAVA, DIANE S.,1,* THOMAS C. BELL1,2 and BARBARA
1
University
of
British
Columbia,
sriRICHARDSON.3
vast@zoology.ubc.ca, Vancouver, BC; 2 Oxford University, Oxford, UK;
3
Braid Road, Edinburgh, UK. Bromeliad food webs as model systems
for biodiversity research.
Biodiversity research requires precise control over species composition in
food webs. This has traditionally been difficult for most animal communities, due to dispersal. One solution is to use food webs in natural container communities, such as bromeliads. We have used the aquatic food
webs in bromeliads to examine processes occurring at individual, population, landscape and biogeographic levels. At the population level, we demonstrate indirect extinction cascades between these two very different
branches of the bromeliad food web: loss of insect diversity leads to loss
of ciliate diversity. The mechanism for this pattern may involve detrital
processing chains, an individual-level effect between detritivore insect species. Bromeliad surveys reveal landscape-level effects on larval abundances, and demonstrate clear patterns in the effects of habitat size and complexity on trophic and functional diversity in food webs. Finally, bromeliad
food webs offer a replicable system for detailed biogeographic compari-

318

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sons, and we show surprising similarities in structure between diverse regions of the world.
ST.CLAIR, SAMUEL B.* and JONATHAN P. LYNCH. Penn State University, University Park, PA, USA. Photochemical responses of sugar
maple and red maple seedlings to excess manganese in contrasting light
environments.
Photosynthetic sensitivity to nutrient imbalances appears to be an important
factor contributing to forest decline in the northeastern US. Tree species
growing on acidic, non-glaciated soils prevalent in the eastern deciduous
forest, accumulate foliar manganese (Mn) to levels that are several times
greater than the critical toxicity threshold in most crop species. We hypothesized that in susceptible species excess foliar manganese decreases
photosynthesis in response to high light, through photo-oxidative stress. To
test this hypothesis, we assessed photosynthetic responses (C02 exchange
and chlorophyll fluorescence) of sugar maple and red maple seedlings at
variable Mn and light levels in greenhouse conditions. Rubisco appeared
to be a key target of Mn toxicity, as indicated by decreased carboxylation
efficiency in seedlings treated with excess Mn. Photochemical sensitivity
to the interactive effects of Mn and light varied with leaf development.
Mature sugar maple leaves were sensitive to excess Mn in high and low
light. High light induced Mn toxicity in young leaves of sugar maple and
mature leaves of red maple. Young red maple leaves were tolerant to Mn
toxicity regardless of the light treatment. These results indicated that under
most treatment conditions, sugar maple was photochemically more sensitive to Mn toxicity than red maple. Excess Mn and high light increased
the PhiPSII /PhiCO2 of mature sugar maple leaves, a physiological marker
indicating susceptibility to photo-oxidation. These results suggest that observed correlations between excess foliar Mn and sugar maple decline
symptoms on the eastern deciduous forest have the potential to be mediated
by Mn toxicity of the photosynthetic apparatus.
STAFFORD, NATHANIEL B.* and SUSAN S. BELL. University of South
Florida, Tampa, FL, USA. Biological correlates of hydrodynamic regime
in a seagrass landscape.
Disturbance events play a significant ecological role in vegetated habitats
by removing plant structure and dramatically altering the biotic and abiotic
parameters of the habitat. An understanding of the link between disturbance
and seagrass landscapes may benefit from an estimation of the level of
physical exposure or wind/wave intensity within an area. Such forces represent chronic disturbances in that they are constant, although their intensity
may vary. A Relative Exposure Index (REI) based on depth, exceedance
wind speeds and fetch was calculated for five seagrass landscapes in Tampa
Bay, FL. At each site seagrass (Thalassia testudinum) rhizomes were collected by excavating three 0.5 x 0.5m areas of seagrass. Internodal distance
was measured on multiple rhizome segments to evaluate the effect of physical disturbance regime on plastic structural characteristics of the seagrass.
Results indicate that seagrass rhizome internodal distances are longer in
landscapes exposed to greater hydrodynamic energies as modeled with the
REI. These results suggest differences in growth patterns as a result of
wind and wave exposure. Further work is underway to evaluate the recovery rate of seagrass following small scale experimental disturbances (0.5 x
0.5m gaps) under different energy regimes, as well as using aerial photography to characterize long-term, large scale gap dynamics in relation to
physical setting. A series of archived photographs spanning a 16-year period have been scanned and the gaps are currently being manually digitized
on-screen. Resulting spatial data will be analyzed using a Geographic Information System (G.I.S.) to correlate gap frequency, size, and expansion/
closure rates, as well as various landscape metrics (e.g., complexity of gap
edges, gap location and arrangement) to general hydrodynamic regime.
STANLEY, THOMAS R.* USGS Fort Collins Science Center, 2150 Centre
Ave., Bldg. C, Fort Collins, Colorado, USA. Deconvolving avian nesting
data using Fourier transforms to recover the distribution of nest initiation times.
Nest initiation in birds is thought to be influenced by factors like temperature, precipitation, predation, and even global warming, and there is much
ecological interest in the influence of nest initiation times on factors like

clutch size, nesting success, and nestling growth and survival. The most
common method for reconstructing distributions of nest initiation times
involves aging the eggs or young in an active nest, then backdating to
determine when the nest was initiated. However, when nests are not found
on the day they are initiated and nest survival rates are less than one, this
method is biased and can lead to erroneous conclusions or inferences. Nest
initiation in a population can be conceptualized as a temporal signal that
passes through a filter, in this case a survival process, to yield an observable
signal n(t) that is the number of active nests in the population at time t. In
this talk, I show mathematically that this filtering process is the convolution
of the distribution of nest initiation times and the survival function for
nests. Because n(t) is observable, and because the survival function can be
modeled and parameterized (e.g., using the Mayfield estimator), it is therefore possible to deconvolve n(t) to recover the unobserved distribution of
nest initiation times. I demonstrate deconvolution of n(t) using the Convolution Theorem and Fourier transforms when n(t) is sampled at regular
or irregular intervals.
STARK, JOHN M. Department of Biology, Utah State University, Logan,
UT, USA. Plant carbon inputs regulate timing of nitrogen pulses in
semiarid soils.
Little is known about the timing and magnitude of nutrient pulses in semiarid ecosystems, and even less is known about the factors that regulate
these pulses. We studied a big sagebrush community in southern Idaho to
answer the questions: Are pulses of water and nitrogen correlated with one
another? Do these pulses coincide with plant demand? And what are the
mechanisms regulating these pulses? We monitored soil moisture, temperature, inorganic N, net N mineralization rates, gross N mineralization and
immobilization rates, and CO2 flux in soils beneath sagebrush and crested
wheatgrass plants at 2-wk intervals throughout the growing seasons of two
years. Interestingly, water pulses did not always cause a pulse of N mineralization, as is usually assumed. During mid- to late summer of both
years, water addition actually stimulated a phase of net N immobilization.
A mass-balance model linking C and N dynamics suggests that this immobilization phase results from the microbial community shifting from low
to high C:N substrates during late July. The shift in substrate C:N appears
to coincide with senescence of plant foliage and possibly near surface roots,
and thus may result from a large release of plant biomass into the labile
soil organic matter pool. During the subsequent few months, the C:N of
this pool gradually declines, presumably due to microbial degradation and
CO2 release, and by the time the fall rains stimulate plant growth, a new
phase of mineralization has begun. Therefore, while water and nitrogen
pulses are not always correlated, water does appear to stimulate N mineralization during seasons when plants are actively taking up N.
STARK, SCOTT C., DANIEL E. BUNKER and WALTER P. CARSON.
University of Pittsburgh, Pittsburgh, PA. Predicting plant invasion in
North American ecoregions: A macroecological approach.
Over three thousand exotic (i.e., non-native) plants currently dwell in the
United States and Canada. Biological invasion (i.e., the influx of successful
exotics) is implicated in the decline and extinction of native organisms.
Ecologists have correlated species traits with invasion success and ecosystem properties with invasibility. For instance, the diversity-invasibility
hypothesis predicts that diverse native ecosystems should resist invasion
and thus exotic richness should decline with increasing native richness.
However, recent macroecological work has shown that these comparisons
can be confounded if the effects of area are not considered. Here we compare species-area curves between exotic and native plants across the major
ecological units (e.g., boreal forest, grassland) of the United States and
Canada. If the slope of species-area curves differ between native and exotic
species, then processes structuring these communities are likely to differ.
The dependence of species richness on area (i.e., slope of the species-area
curve) differs between ecoregions but is strikingly similar for exotic and
native species within individual ecoregions. This suggests that exotic fraction (exotic richness / native richness: a common metric of invasion) is an
area-independent metric that may be employed in site comparisons within
certain ecosystems. However, some ecosystems may not follow this generalization (e.g., grasslands). After controlling for area dependence we tested the diversity-invasibility hypothesis at large scales. Within U.S. states

and Canadian provinces, area-corrected native and exotic richness have a

strong positive correlation (r250.42). In agreement with the findings of
other recent studies, the diversity invasibility hypothesis is not supported
at large scales.
STARNES, JOHN H.,1,* JENNIFER E. HEWITT,2 MICHELE M. WEBBER,1,2 ALBERT J. MEIER1 and MITCHELL B. CRUZAN.3 1 Western
Kentucky University, Bowling Green, KY; 2 Mammoth Cave National Park,
Mammoth Cave, KY; 3 Portland State University, Portland, OR. Response
of Eggerts sunflower (Helianthus eggertii) to disturbance at Mammoth
Cave and Coffee County.
Eggerts sunflower (Helianthus eggertii) is a federal and state-listed threatened plant that occurs from central Kentucky and Tennessee to northern
Georgia. The largest studied occurrences are in Coffee County, Tennessee.
Twenty-two small occurrences are in and around Mammoth Cave National
Park, Kentucky. Although efforts are currently in place to protect known
populations and to increase the total number of individuals throughout both
sites; several life history responses to disturbance have not been studied.
At MACA we studied the effects of substrate disturbance on this threatened
species and to determine whether relocation is a viable conservation method. The roadside population was transplanted to two similar sites within
the park. We measured the disturbed population, 5 natural undisturbed populations, and 5 groups of transplanted individuals during the 2002 growing
season. We found that the original roadside group increased from 1505
stems before the relocation event to 2480 stems. We found that severe
substrate disturbance increases stem height and density over one growing
season. At AEDC we looked at several different factors that could effect
the growth of H. eggertii in three occurrences. There were several differences in these populations such as fire history, management schemes, and
individual site characteristics. Site 1 was a heavily disturbed site in an open
field. Site 2 was a roadside population at the forest edge, and site three
was a forest site with frequent fires. We found that several life history traits
were significantly different in the sites such as height, diameter of stems,
nodes, flowers, reproduction, and leaves (P,.0001).
STARNES, SARAH M.* and MATTHEW D. MORAN. Hendrix College,
Conway, AR. Response of individual plants to fire in a relict grassland.
We tested the role of a spring fire on growth rates, survivorship, and seed
production of individual plants in a relict grassland. We performed a prescribed burn on April 1, 2002 on one half of our field sites and compared
the response of the plants to unburned control sites. We examined the
response of six plants species including three grasses (Schizachyrium scoparius, Sporobolus sp., and Panicum virgatum) and three forbs (Coreopsis
lanceolata, Specularia perfoliata, and Echinacea pallida), the six most
common species at our site. The two annual forbs, C. lanceolata and S.
perfoliata showed reduced growth rates and high mortality in the burned
plots while the perennial forb, E. pallida, exhibited no response. Among
the grasses, P. virgatum exhibited increased growth and seed set in burned
plots while seed set was much reduced in Sporobolus. Surprisingly, S.
scoparius, the dominant plant on our field site, showed no response to
burning. These results show that there is a varied response to burning by
the dominant plant species, and therefore it is likely that periodic fire will
change the community structure of this grassland.
STARR, GREGORY,1 TIMOTHY A. MARTIN,1 MICHAEL W. BINFORD,2 HENRY L. GHOLZ1,3 and LEVENT GENC.2,* 1 School of Forest
Resources and Conservation, University of Florida, Gainesville, FL, USA;
2
Department of Geography, University of Florida, Gainesville, FL, USA;
3
National Science Foundation, Arlington, VA, USA. Integration of carbon dynamics from leaf to landscape in Florida pine forest.
In recent years Florida AmeriFlux studies have shown that managed plantations of Pinus elliottii have the capacity to rapidly sequester carbon (maximum annual NEE approximately 800 g C m-2 year-1). However, from 1999
to 2001 a severe drought reduced rainfall in the north central Florida region
by 60%. Maximum canopy leaf area index remained relatively stable over
the drought with minor changes in temporal display occurring due to early
leaf senescence. Midday stomatal limitations reduced leaf-level photosynthetic capacity of pines by 30%. At the ecosystem level, daytime net eco-

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319

system exchange (NEE) of CO2 showed an approximately 30% reduction,

with similar reductions in night ecosystem respiration rates as the drought
persisted. Because of these compensating effects of decreased GEP and
decreased ecosystem respiration, NEE on longer time scales in our P. elliottii flatwoods systems appears to be relatively stable in the face of extreme soil water deficits. A collaborative effort between the Florida
AmeriFlux project and a UF NASA-funded project is currently attempting
to estimate integrated NEE over time for a 15 km x 15 km landscape in
north-central Florida. This effort combines eddy-covariance and mensurational measurements of NEE with land-use and stand-developmental classifications made from Landsat images. Preliminary results suggest that NEE
of this forested landscape remains relatively stable over decadal time scales.
At this same scale NDVI decreases over the severe drought, indicating a
lack of relationship between NDVI and NEE. A more relative relationship
appears to exist between GEP and NDVI at the landscape level within these
coniferous forests. The study of these relationships between NDVI, NEE,
and GEP are currently being expanded beyond the conifer forests of the
regions and into riparian and agricultural areas with the intentions of developing a better understanding of the regions carbon dynamics.
STEELE, MICHAEL A.,1,* TOM A. CONTRERAS,1 AMY B. MCEUEN,1
MICHAEL SHERICK,1 SETH REED1 and PETER SMALLWOOD.2
1
Wilkes University, Wilkes-Barre, PA; 2 University of Richmond, Richmond, VA. Differential responses of oak species to acorn masting: an
evaluation of the predator satiation hypothesis.
The predator satiation hypothesis suggests that masting evolved in response
to seed predators and that seed survival and seedling establishment will
only occur during high mast years. Oak species may, however, differ in the
conditions required to satiate predators because of specific acorn-hoarding
decisions of small mammals. Specifically, small mammals in temperate oak
forests are known to selectively disperse and cache acorns of the red oak
subgenus (Eythrobalanus) in contrast to acorns from the white oak subgenus (Quercus) which are preferentially eaten. We therefore predicted that
oak species would show differential responses to oak masting depending
on the degree of synchrony of acorn production across species and subgenera. We hypothesized that caching and recruitment of red oaks will
occur in all years of high acorn production and that white oaks should
establish only when they mast alone. We tested this prediction by monitoring acorn production and following the fate of tagged acorns (9000/year)
at three sites in NE Pennsylvania over four years. As predicted, caching
and establishment of northern red oak (Quercus rubra) increased in a year
of high red oak and white oak production. White oaks (Quercus alba) failed
to mast without red oak in the years of our study and, as predicted, caching
for this species was extremely low across all years and sites. We suggest
that the conditions necessary for establishment of white oak species are far
more limited than for species of red oak. We also argue that an understanding of the behavioral decisions of small mammals may be key to explaining
patterns of oak recruitment within temperate forests.
STEETS, JANETTE A.* and TIA-LYNN ASHMAN. Department of Biological Sciences, University of Pittsburgh, Pittsburgh, PA. Vegetative herbivory affects the mating system of Impatiens capensis.
Vegetative herbivory can affect plant mating system directly, by lowering
plant resources, or indirectly, by influencing pollinator visitation to damaged plants. Using multiple experimental approaches, we analyzed the
mechanisms by which vegetative herbivory affects the mating system of
Impatiens capensis, a species with an environmentally-determined mixedmating system (producing both cleistogamous, obligately selfing flowers,
and chasmogamous, facultatively outcrossing flowers). We found that herbivory affected the expression of mating system directly by increasing the
proportion of flowers and seeds that were cleistogamous and by altering
the relative quality of cleistogamous and chasmogamous progeny. The
overall direct effect of herbivory on mating system was to increase the
proportion of fitness achieved through cleistogamous progeny. In addition,
vegetative herbivory affected characteristics associated with the mating system including chasmogamous floral display size and floral attraction traits,
resulting in a decrease in pollinator visitation to damaged plants. The significant decreases in flower number, flower shape, and pollinator visitation,
may lead to changes in the outcrossing rate of chasmogamous flowers. Our

320

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findings are among the first to demonstrate that vegetative herbivory has
direct consequences on plant mating system and should be considered a
factor shaping mating system evolution.
STEGEN, JAMES C.* and ROSS BLACK. Eastern Washington University,
Cheney, Washington, USA. The predator-prey relationship between water mites (Piona sp.) and the zooplankton of small ponds.
Relative to other invertebrate predators, the importance of water mites to
zooplankton community dynamics is poorly understood. We examined the
in situ impact of natural mite densities (Piona sp.) on the ambient zooplankton community in 20L enclosures over 5 days. Relative to a control,
we observed a 60% and 63% drop in Daphnia pulex biomass in the 0.1
and 0.2 mites/L treatments, respectively. This pattern suggests interference
among predators. An estimate of community importance was calculated at
256, and the per capita interaction strength was calculated to be 20.28.
Both values suggest water mites as potential keystone predators. A second
experiment showed that the effect of mite predation on D. pulex does not
change with changing D. pulex density. The interactive effect of notonectids and mites on the zooplankton community was investigated using 378L
mesocosms. We found these predators to interact additively in their effect
upon D. pulex. In addition, these predators resulted in a positive indirect
effect on the copepod Diaptomus leptopus. Since D. leptopus density was
negatively correlated with D. pulex denisty, this indirect effect may have
been mediated through the reduction in D. pulex density.
STEINAKER, DIEGO F.* and SCOTT D. WILSON. University of Regina,
Canada, Regina, Saskatchewan, Canada. Above- and belowground contributions to nitrogen cycling in aspen forest and northern prairie.
Studies of nitrogen dynamics typically focus on aboveground production,
and report greater N inputs to the soil from primary production (NPP) in
forest than in prairie. But belowground NPP is often greater than aboveground, and the true values of N input from primary production, therefore,
may be different than previously appreciated. Our main goal was to quantify the contributions of above- and belowground biomass to nitrogen cycling in adjacent aspen (Populus tremuloides) forest and mixed-grass prairie habitats, at the northern edge of Great Plains, in Saskatchewan, Canada.
Aboveground NPP and N input to the soil (N input 5 NPP x [N]) was
determined from herbaceous plants litter production in both habitats. Belowground NPP and N input was measured using a combination of observation using minirhizotron images and destructive sampling. NPP and N
inputs from aboveground biomass were three times greater in forest than
prairie, but productivity and N inputs from aboveground were small compared with those from belowground. Root production accounted for about
75 % and 90 % of total primary productivity in forest and prairie respectively. Root length and mass was greater in forest than in prairie, but length
and mass production did not differ significantly between the habitats. Although nitrogen concentration was higher in forest than in prairie roots,
annual N input to the soil from fine roots was not significantly different
between habitats. Our results show that biomass production and nitrogen
inputs via fine roots greatly exceeded those from foliar litter. As a result,
N input from total primary productivity in prairie was at least comparable
to forest.
STEINAUER, MICHELLE L. and JAMES E. PARHAM.* University of
Nebraska, Lincoln, NE. Geographic distribution of host and habitat use
of an acanthocephalan parasite, Leptorhynchoides thecatus.
Leptorhynchoides thecatus, a parasite of freshwater fishes, is variable in
host use, development, and habitat use throughout its range. To examine
the extent and pattern of variation, a meta-analysis was performed. Fish of
more than 40 genera were reported definitive hosts for L. thecatus; however,
many of these fishes infrequently serve as hosts for this species. Common
hosts were determined by calculating prevalence ranks for each fish species
at each site from which this parasite was reported. Species of black bass
and rock bass were the most common hosts used. Development of L. thecatus within the fish host is another variable trait because fish can serve
as either definitive or paratenic hosts. Maturity ratios based on the relative
prevalence of adults and cystacanths were compared across watershed regions for 3 fish species. Smallmouth bass rarely harbored cystacanths at

any location. Most of the time, largemouth bass harbored adults in the
central United States and upper Great Lakes region, but harbored cystacanths more often than adults in the southeastern U.S. Adult L. thecatus
were more prevalent than cystacanths in bluegill sunfish in the north central
U.S., but cystacanths were more prevalent in the Great Lakes region and
south central U.S. Habitat use of L. thecatus within the definitive host also
differs and worms can inhabit the ceca, intestine, or both regions. Analysis
of this trait was difficult because most papers did not report specific location of the parasite. However, L. thecatus occurred in the intestine of
sunfishes in the southeastern U.S., in the ceca in fish of all species in the
central U.S., and in both ceca and intestine in fish of all species in the
Great Lakes area. The results of this meta-analysis raise further questions
concerning the cause of variation in L. thecatus. Differences in life history
traits may be due to ecological factors that differ across geography or to
genetic differences, which may indicate that L. thecatus comprises multiple
cryptic species.
STEINER, CHRISTOPHER F.1,* and MATHEW A. LEIBOLD.2 1 Rutgers
University, New Brunswick, NJ, U.S.A.; 2 University of Chicago, Chicago,
IL, U.S.A. Community assembly and the generation of scale-dependent
productivity-diversity relationships.
The relationship between species diversity and productivity can be strongly
scale-dependent. At the local scale of population dynamics the relationship
is commonly unimodal or humped, but it is frequently monotonically increasing at larger spatial scales. Such contrasting patterns can occur if beta
diversity (compositional dissimilarity among sites) also increases with productivity, but mechanistic reasons for this remain clouded. Here we use
models to explore the influence of ecological assembly on the diversity of
multitrophic communities along gradients of productivity. We show that
assembly dynamics can generate scale-dependent patterns consistent with
those observed in nature; while unimodal productivity-diversity patterns
are produced at the scale of local communities, positive monotonic patterns
emerge when diversity is measured at larger spatial scales (among communities). This occurs due to increases in beta diversity with productivity.
Our results suggest that increases in beta diversity, and resultant scaledependency, depend on three vital and interactive elements: the presence
of more than one consumer trophic level, a greater propensity for cyclic
compositional change at high productivities, and the stochastic nature of
species dispersal and invasion history.
STEINER, SUSAN M.* and BRIAN D. KLOEPPEL. University of Georgia, Otto, NC, USA. Activities, experiences, and products of the Coweeta Schoolyard LTER Program, 1998 to 2003.
This program provides formal instruction, field research experiences, and
data summary and analysis experiences to K-16 students and instructors
using Coweeta Long-term Ecological Research (LTER) projects as a model.
Coweeta LTER scientists and staff have been providing middle school, high
school, and community college students "hands on" field and laboratory
research experience for the last five years. The types of activities include
assisting scientists with current research projects such as the leaf productivity measurement on the terrestrial gradient plots, overland water flow
measurement for the riparian zone restoration project, and tree stem respiration measurements for the carbon flux component of the LTER regionalization project. Other activities have been custom designed to minimize
the "bigfoot" effect on our long-term research sites. These activities provided students and teachers with optimal research experience emphasizing
data collection and analysis techniques that dovetail into current projects.
These custom activities include a remeasurement of tree size at a longterm research site in the Joyce Kilmer Old Growth Forest in western North
Carolina. During a week long intersession of the Rabun Gap Nacoochee
School in Rabun Gap, Georgia, students and teachers assisted scientists
remeasuring 350 trees. Three study sites were established in 1995 that
included cove, mid-slope, and ridge habitats. The participants learned techniques of plot layout, field identification of trees, and tree measurement.
Teachers and leaders from the school also included interdisciplinary curricula such as journal writing, historical literature, math, and statistical
analysis skills. This multidisciplinary approach partnered Coweeta scientists with the Rabun Gap Nacoochee School Environmental Stewardship
Program for a week of productive science activities for both groups. The

Coweeta Schoolyard LTER Program acknowledges funding since 1998

from annual supplement grants from the National Science Foundation.
STEPHENSON, ANDREW G., JORGE I. MENA-ALI* and STEVEN E.
TRAVERS. Dept of Biology, The Pennsylvania State University, University Park, PA, USA. Effect of genotype and floral age on growth of selfpollen tubes in styles of Solanum carolinense.
Self incompatibility (SI) is a widespread character, present in many plant
species. Because SI is regulated by S-alleles, it is often considered or treated as a qualitative trait, species being either self-incompatible or self-compatible. In contrast, numerous studies have found that the strength of SI
varies between populations within a species, and even between individuals
within populations. Several factors have been suggested to explain the degree of variation in SI, but many of these factors have not been examined
until very recently. Furthermore, there is a lack of information regarding
how this variation may affect the genetic makeup and evolutionary history
of plant populations over time. Recently, we have begun to investigate the
behavior of self-pollen tubes in Solanum carolinense, a self-incompatible
perennial weed. A series of controlled pollinations were performed in order
to determine how the age of the flowers and the SI genotype affect the
growth of self-pollen tubes. We found that the growth of self pollen tubes
is significantly influenced by the genotype of the plant, the age of flower
and their interaction. We are currently examining whether these differences
among genotypes are due to specific S-alleles (and closely linked modifiers)
or whether they are due to unlinked modifiers of SI.
STEPHENSON, ROBERT E.* and STEVEN A. JULIANO.* Illinois State
University, Normal, IL, USA. Declining food quality and development
of a juvenile insect: Adaptive response or physiological constraint?
The Wilbur-Collins model of growth and development postulates continuous flexibility of development rate in response to changing growth rate. In
contrast, the Travis model postulates that development becomes unresponsive to changing growth rate late in the life cycle. We tested these models
by determining the effects of a decline in food quality on metamorphosis
to the adult stage in males of the grasshopper Romalea microptera. Newly
eclosed 5th instar R. microptera were raised on six different diet regimes,
including high (5%) protein food, low (1%) protein food, and a series of
high-to-low switches that occurred on days 4, 7, 10, and 13 of the 5th
instar. Males in the low-fed (LL) and switch day 4 (HL4) groups took a
significantly longer time to reach the adult molt than did males in the highfed (HH), switch day 10 (HL10), and switch day 13 (HL13) groups. Means
for HH, HL13, HL10, and HL7 were not significantly different. These
results are consistent with the hypothesis that life history transitions such
as the molt to the adult stage become inflexible in response to environmental fluctuations such as declining food quality, thus suggesting a physiologically determined period of developmental canalization late in the juvenile life of this grasshopper. These results refute the hypothesis of adaptive acceleration of development in response to a deteriorating environment. These results suggest that juvenile development in this grasshopper
follows a pattern consistent with the Travis model of growth and development, and inconsistent with the Wilbur-Collins model of growth and
development.
STERNER, ROBERT W.,* JAMES M. HOOD and HEATHER HENDRIXSON. Ecology, Evolution and Behavior, University of Minnesota, St.
Paul, MN, USA. Organismsal stoichiometric patterns: intersections
with temperature and body size.
Two studies that integrate body size, biophyics and biological stoichiometry will be discussed. In the first study, growth rate of Daphnia herbivores
as a function of temperature and stoichiometry was examined. Animal P
content may relate to growth by way of P-rich nucleic acids, particularly
RNA. We found strong growth-P-RNA couplings under conditions of high
food quantity. However, under low food quantity, stoichiometric couplings
broke down. Temperature effects were well described by the Bolztman
relationship and were independent of nutritoinal state, allowing us to write
a single predictive equation for growth that includes temperature and stoichiometry. In the second study, we examined the stoichiometry of several
elements (C,N, P and some trace elements) in freshwater fish species from

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321

Minnesota. Interspecific variation in element content was large. Ca and P

were tightly correlated with a ratio equal to the Ca:P in fish bone. Hence,
previous suggestions that body P in vertebrates is tied to skeletal investment
were supported. The different elements exhibited differing allometric relationships.
STEVENS, GLEN N.* and ROBERT H. JONES. Virginia Polytechnic Institute and State University, Blacksburg, VA. Root proliferation and herbivory in nutrient rich soil patches: A tradeoff for plants?
Root growth is typically stimulated by nutrient rich soil patches, but rates
of root herbivory may be too, which could result in important tradeoffs for
plants. We investigated net fine root biomass production in response to
microsite fertility and fine root herbivores in an aggrading (7 years since
clearcut harvest) upland forested stand in the coastal plain of South Carolina. Two hundred randomly located ingrowth cores were established in
the stand in March 2002 by first removing a 10 cm diameter x 30 cm deep
core of soil, removing the roots, amending the soil according to treatment,
and then repacking the soil to approximate the original bulk density. Treatments were randomly assigned to ingrowth core sites (50 per treatment)
from a 2 x 2 factorial combination of +/- fertilizer (Osmocote slow-release
15-9-12) and +/- pesticide (granular chlorpyrifos, Lorsban 15G). Cores
were harvested in October and November of 2002 (100 cores each month).
We observed no significant effects of harvest date on net ingrowth core
root biomass (P 5 0.22). Both pesticide applications and fertilizer additions
led to significantly greater root biomass (both P , 0.001); additionally, we
saw a significant interaction between pesticide and fertilizer (P , 0.05).
The significant pesticide effect suggests that root-feeding insects are having
a strong impact on root biomass production. More interestingly, the significant interaction between herbivores and fertility showed stronger herbivore impacts on more fertile microsites. Thus, while a plants tendency
to proliferate roots in nutrient-rich patches may lead to increased nutrient
uptake, increased vulnerability to root herbivory is a potentially significant
cost of such root foraging.
STEVENS, MARTIN HENRY H.1,* and WALTER P. CARSON.2,* 1 Ecology Graduate Program, Department of Botany, Miami University, Oxford,
OH, USA; 2 Program in Ecology and Evolutionary Biology, Department of
Biological Sciences, University of Pittsburgh, Pittsburgh, PA, USA. Resource ratio tradeoffs exist, but fail to explain effects of nitrogen on
plant species richness.
Tradeoffs in plant traits associated with resource ratio specialization and
response nonlinearity should explain variation in plant species richness
along productivity gradients. Unfortunately, few experiments have documented the presence of such tradeoffs in general, nor have any experiments
tested whether such tradeoffs can explain changes in relative abundance
associated with the loss of diversity. We manipulated light and soil nitrogen
in field experiments that tested for the presence of these tradeoffs, and
tested whether these tradeoffs explain the loss of richness with increasing
soil nitrogen. Common species appeared to occupy significantly different
light and nitrogen niches that were consistent with tradeoffs assumed by
resource ratio specialization and response nonlinearity. Tradeoffs existed
between species success in low light versus species sensitivity to increasing
light (r 5 -0.90), and between species sensitivity to increasing light versus
sensitivity to increasing soil nitrogen (r 5 -0.66). These results are consistent with both resource ratio and response nonlinearity as general mechanisms of coexistence in plant communities that are especially relevant to
resource gradients. These tradeoffs, however, failed to explain the change
in species abundances along the resource gradient. Thus, while these tradeoffs appear to exist, they do not explain the loss of species richness along
this soil resource gradient.
STEVENS, MICHAEL T.* and RICHARD L. LINDROTH. University of
Wisconsin-Madison, Madison, WI. Cost of tolerance to herbivory in aspen.
Plant defense against herbivores often comes at a cost to plant fitness.
Numerous studies have addressed costs of chemical resistance, but few
have addressed costs of tolerance to herbivory, especially in woody plants.
Aspen (Populus tremuloides) is an early-successional tree subject to dam-

322

Abstracts

age by a wide variety of herbivores. Aspen is defended chemically, but can

also counteract the negative effects of herbivore damage through regrowth.
To determine if this form of defense is costly, we utilized a common garden
of 12 aspen genotypes grown under two nutrient conditions and subjected
to either 0% or 75% defoliation. The diameter (d) and height (h) of each
aspen ramet was measured initially in October 2001 and again in August
2002. We used d2h as an index for growth since it is highly correlated with
total biomass based on a harvest of a subset of ramets (R2 5 0.99, p ,
0.05). The growth of ramets in our experiment was significantly affected
by nutrients, defoliation, genotype, and treatment interactions (p , 0.05).
Defoliated ramets grew 23% less than undefoliated ramets. Within a genotype, tolerance is defined as the difference in growth between damaged
and undamaged plants. A cost of tolerance is indicated by a negative relationship between growth when undamaged and tolerance. After one season of growth, we found such a relationship under low nutrient conditions
(r 5 -0.681; p , 0.05), but not under high nutrient conditions. Under low
nutrient conditions, the most tolerant genotypes grew the least when undamaged. This cost may be a result of lost opportunities for photosynthesis
due to allocation to roots instead of leaves. Stored reserves in the roots are
an effective means for rebounding after herbivory, but genetically-determined allocation to roots rather than leaves may come at a cost to growth
when herbivores are few or absent.
STEVENS, MICHELLE L.,1,* SUZIE ALWASH,1 AZZAM ALWASH,1
THOMAS L. CRISMAN,2 GEORGE ZALIDIS,3 HASSAN PARTOW,4
MIKE EVANS5 and BRIAN COAD.6 1 Eden Again Project, Long Beach,
CA, USA; 2 Director, Center for Wetlands, University of Florida, Gainesville, Florida, USA; 3 Greek Biotope Wetland Centre, Aristotle University,
Thesaliniki, Greece; 4 UNEP-GRID Geneva, Division of Early Warning and
Assessment, CH1219 Chatalaine, Geneva; 5 Birdlife International, Wellbrook Court, Girton Rd., Cambridge, CB30NA; 6 Canadian Museum of
Nature, Ottawa, Canada. Eden again: A conceptual restoration plan for
the Mesopotamian marshlands of southern Iraq.
The Eden Again Project is developing scientific guidelines for a conceptual
restoration plan for the Mesopotamian Marhshlands of southern Iraq, at the
confluence of the Tigris and Euphrates river basins. An international Technical Advisory Panel was convened to develop scientific guidelines for
restoring these marshes, which historically covered 15,000-20,000 km2 and
are now drained and dessicated to less than 90% of their original area.
Regional adverse impacts include deterimental impacts to Perisan Gulf fisheries, possible extinciton of 21 globally endangered or endemic species,
major reductions in the West Siberian-Caspian-Nile flyway, regional climate change, and loss of culturally significant resources. Restoration planning is based on a hydrologic model developed with satellite imagery,
available biological data primarily gathered prior to 1980, interviews with
exiled southern Iraqi people, and analogs from adjacent areas. The primary
goal is to utilize available water resources to demonstrate a mosaic of
opportunities for restoration as one strategy to be considered by the people
of Iraq.
STEVENS, RICHARD D.* National Center for Ecologial Analysis and
Synthesis, University of California, Santa Barbara, Santa Barbara, CA.
Latitudinal gradients in the phylogenetic diversity of New World bat
communities.
Although described for many taxa, latitudinal gradients in biodiversity have
typically focused on the number and relative abundances of species. Nonetheless, other measures of biodiversity, especially measures of functional,
morphological, and phylogenetic diversity may exhibit complementary yet
more informative geographic gradients. I investigated latitudinal gradients
in the phylogenetic diversity of 32 New World bat communities. I used a
supertree of all extant bat species generated by Jones et al. (2002) to characterize the phylogenetic relationships among taxa. I then quantified four
aspects of the phylogenetic diversity of communities: 1) average pair-wise
nodal distance among species, 2) average nearest-neighbor nodal distance,
3) variance of pair-wise nodal distances among species, and 4) variance of
nearest-neighbor nodal distances. Significant latitudinal gradients exist regarding phylogenetic diversity. Nonetheless, these gradients could result
from the underlying latitudinal gradient in species richness. I performed a
suite of randomization analyses to characterize latitudinal gradients in phy-

logenetic diversity that would be expected given the observed gradient in

species richness. A number of phylogenetic-diversity gradients were different from simulated expectations. In general, phylogenetic diversity was
lower and decreased with increasing latitude at a faster rate in the observed
communities than in simulated ones. To the extent that phylogeny reflects
the evolutionary and ecological attributes of organisms, measures of phylogenetic diversity should characterize emergent properties of communities
that reflect meaningful variation among species. Although taxonomic and
phylogenetic diversity gradients are complementary, it is likely that mechanisms other than those that affect taxonomic diversity affect the phylogenetic composition of species within communities.
STEVENSON, BLAKE K.* and MATTHEW D. MORAN. Hendrix College, Conway, AR. Effect of spring burning on an arthropod community in a relict grassland.
We tested the role of a spring fire on the arthropod community inhabiting
a relict grassland in central Arkansas. Because spring is the time of rapid
population growth in grassland arthropods, we hypothesized that fire would
cause significant reductions in these organisms. We performed a prescribed
burn on April 1, 2002 on one-half of the plots, and compared the response
of the arthropod assemblage to unburned control plots. The total number
of arthropods was reduced in burned plots. However, the response was not
uniform between trophic levels. Herbivores showed a strong decline while
carnivores and detritivores showed no significant responses. Amongst the
herbivores, both sap-feeding and grazing arthropods declined in abundance.
The response of herbivores in this experiment was stronger than previous
experiments that utilized a fall burn. Spring is not the time of natural fires
in central Arkansas because of high rainfall and moderate temperatures.
We suggest that many herbivorous arthropods are not adapted to fire in this
season. These results have implications for the management and restoration
of these grasslands as most prescribed burning is currently performed in
the spring at these field sites.
STILES, ARTHUR1,* and SAMUEL SCHEINER.2 1 Department of Plant
Biology, Arizona State University, Tempe, AZ; 2 Division of Environmental
Biology, National Science Foundation, Arlington, VA. Effects of habitat
fragmentation on remnant Sonoran desert plant communities in the
CAP-LTER, Phoenix, Arizona.
The Phoenix metropolitan area has experienced rapid rates of population
growth in the second half of the 20th century. This has been accompanied
by the large-scale conversion of Sonoran desert scrubland to urban land
covers. In the Salt River Valley, many desert mountains and buttes were
preserved as parkland as development marched over the plains, resulting
in a network of remnant patches, usually dominated by an isolated or range
of peaks, separated from each other by stretches of residential and commercial land. This urban matrix represents a barrier to free flow of desert
plant propagules across the landscape. In order to assess the impact of
habitat fragmentation on remnant woody vegetation, which includes an
assortment of tree, shrub, and cactus species, field sampling was conducted
in twenty remnant patches arrayed across metropolitan Phoenix. These data
were used to generate estimates of small-scale species density as well as
large-scale estimates of patch richness. Sampling was stratified by habitat
type and diversity measures were calculated as weighted averages based
on the distribution of major patch habitats, including slope exposures, flatlands, and washes. Relationships between species diversity and patch characteristics (e.g. topographical area, isolation, habitat heterogeneity) are explored with regression analysis.
STILING, PETER* and DANIEL MOON. Department of Biology, University of South Florida, Tampa, FL, USA. Disentangling the effects of
host plant quality and quantity on trophic interactions in a salt marsh
community.
Productivity has been argued to be one of the most important factors influencing trophic dynamics. A difficulty inherent in the studies that manipulate productivity by changing nutrient availability is that plant quality and
plant biomass are influenced simultaneously. In this study we disentangled
the potentially confounding effects of plant quality and quantity on trophic
dynamics by separately manipulating nutrients and plant biomass, while

simultaneously reducing pressure from the most common natural enemies

of two focal herbivores, in a fully factorial design. Plant quality of the sea
oxeye daisy, Borrichia frutescens, a common coastal species in Florida,
was manipulated by adding nitrogen fertilizer and adding sugar to decrease
available nitrogen. We manipulated plant biomass by pulling by hand 25%
or 50% of Borrichia stems on each plot. Parasitism of our focal herbivores,
Pissonotus quadripustulatus and Asphondylia borrichiae was reduced with
yellow sticky traps that catch hymenopteran parasitoids. Plant quality significantly affected both herbivores, with fertilization increasing, and sugar
decreasing the densities of the two species. Plant quality also significantly
affected other common herbivores on Borrichia, as the density of a stem
boring lepidopteran and the frequency of leaf chewing varied directly with
foliar nitrogen levels. Biomass manipulations had no significant effects on
any herbivore species. Parasitoid removal treatments significantly increased
the densities of both focal herbivores, and the impact of top-down manipulations was nearly equivalent to the impact of plant quality manipulations
on herbivore densities. These top-down manipulations resulted in a trophic
cascade, as the density of Borrichia stems decreased significantly on parasitoid removal plots. In this study we found that plant quality and not
plant quantity influences trophic interactions. Thus, the utility of predictive
trophic theories incorporating productivity may depend upon our ability to
distinguish between the effects of plant quality and those of plant quantity
and architecture.
STILLWELL, CRAIG* and CHARLES W. FOX. University of Kentucky,
Lexington, KY. Thermal evolution of life history traits: The effect of
temperature during development and egg laying.
In many ectothermic animals, changes in temperature experienced during
the lifetime of an organism result in phenotypic plasticity of several life
history traits. However, few studies consider the separate and interactive
effects of temperature during development and temperature during egg laying on these traits. We investigate the effects of temperature during development on adult body size and development time in two populations of
the desert seed beetle, Stator limbatus, occurring at different elevations and
examine the interactive effects of temperature experienced during development and during egg laying on age at first reproduction, egg size, fecundity and egg mortality. For body size and development time, we observed significant population by temperature interactions suggesting that
these traits may be targets of thermal selection (although the effect was
small for development time). At low temperature the high elevation population was larger in body size whereas at high temperature the low elevation population was larger in body size. There were strong interactions
between the effects of developmental and egg laying temperatures on age
at first reproduction, fecundity, egg size and egg mortality. We also found
that the typical egg size/fecundity trade-off is influenced by temperature:
The shape or magnitude of the trade-off depends on both developmental
and egg laying temperature of females. This study demonstrates that life
history trait responses to temperature are complex and dependent upon the
interactive effects of temperature experienced during development and egg
laying.
STIMMLER, BRYAN* and CHERYL SWIFT.* Whittier College, Whittier,
CA. The effect of disturbance on Alnus rhombifolia in Southern California riparian communities.
Alders, Alnus rhombifolia, are a dominant species in riparian woodlands
in Southern California. Alder stems appear to grow in linear clumps suggesting that resprouting from fallen stems is a primary mechansim for
reestablishment following disturbance resulting from increased discharge
during storm events. Discharge, the amount of water moving through the
stream, is proportional to drainage area, and flood frequency and intensity
are directly proportional to discharge. The frequency with which stems fall
as a result of high discharges influences the structure of the community as
the number of stems increase. The purpose of this study was to determine
the effect of the probability of flooding on the number and size of alders
in two reaches draining different sized watersheds with different discharges.
The results of this study show a shift from fewer larger stems to more,
smaller stems with an increase in discharge. This shift is the result of
increased numbers of fallen trees due to increases in flood frequency and
intensity. This leads to a mix of large and small stems as older branches

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323

establish vertical stems with younger branches, and as new vertical stems
are established by trees deposited from upstream by floods. The reach
draining a smaller watershed supported fewer larger stems, and stems of
different ages were not clumped together. Higher densities of smaller stems
increased sediment trapping within the reach sampled. Bank topography
also affected the structure of the alder community. Within a reach steeper
banks supported fewer larger stems as a result of decreased overbank flow
leading to decreased disturbance and fewer fallen trees.
STIMSON, HUGH C.,1,* DAVID D. BRESHEARS,2 SHAWN C. KEFAUVER1 and SUSAN L. USTIN.1 1 University of California, Davis, Davis,
CA; 2 Los Alamos National Laboratory, Los Alamos, NM. Spectral sensing of foliar water conditions in two conifer species.
Despite the fundamental importance of plant water stress in determining
properties and dynamics at ecosystem and landscape-scales, approaches for
assessing plant water stress at these scales are largely lacking. We evaluated
the relationships between foliar water content and spectral changes in two
coniferous speciesPinus edulis and Juniperus monospermawhich are
codominants of extensive pinon-juniper woodlands in North America. Two
indices of foliar water condition, plant water content (% mass) and plant
water potential, were compared to four spectral analyses: continuum removal of the 1200 and 970 nm water absorption features, the Normalized
Difference Water Index (NDWI), and the "red edge" position. For P. edulis,
plant water content was significantly correlated with all four indices, red
edge position (R25 0.45) and 1200 nm continuum removal (R250.64) were
exceeded by the NDWI (R250.91) and the 970 nm continuum removal
(R250.92). Although the correlations were weaker than for plant water
content, plant water potential for P. edulis was significantly correlated with
red edge position (R250.31), 970nm (R250.41) and NDWI (R250.48). The
relationships were weaker for J. monosperma: water content was significantly related to the 970 nm and 1200 nm continuum removal (R250.35
and R250.37 respectively) and the 1200 nm continuum removal was related
to water potential (R250.40).
STINSON, KRISTINA A.* and F. A. BAZZAZ. Harvard University, Cambridge, MA. Elevated CO2 reduces size asymmetry in competing stands
of Ambrosia artemsiiafolia.
The predicted doubling of atmospheric carbon dioxide over the next century is likely to affect population, community and ecosystem processes by
altering competition for resources within and among plant species. We investigated how CO2 enriched environments would influence light competition in the allergenic species, common ragweed(Ambrosia artemisiifolia)
using an open top chamber design in which competing stands were grown
at either ambient (360 ppm) or twice ambient (720 ppm) levels of atmospheric CO2. We tested whether the variance in size, photosynthetic rate
and architecture between subordinate and dominant individuals was reduced at elevated CO2. We also generated vertical light profiles to determine whether predicted changes in plant architecture and size hierarchies
altered the availability of light within competing stands. Initially, ambient
chambers received less light than elevated chambers. Over time, however,
ambient chambers received more light than elevated chambers. This difference in light availability was apparently due to shading caused by architectural changes in the plants. Stands grown at elevated C02 had lower
variances in total branch length between subordinate and dominant plants
than stands grown at ambient conditions. Photosynthetic rates and biomass
of both dominants and subordinates were reduced at elevated CO2. However, total branch length of subordinates increased significantly at elevated
CO2 while ambient grown stands maintained greater size asymmetry for
this trait. We conclude that subordinates are able to catch up to dominant
plants in high CO2 conditions via changes in canopy architecture. Reduced
size asymmetry between dominants and subordinates may also be facilitated by reduction in performance of dominants. Reduced competition for
light may lead to greater population sizes of Ambrosia artemissiifolia in
the near future.
STODOLA, KIRK W.,1 ERIC LINDER,1 DAVID BUEHLER2 and DAN
KIM.2 1 Mississippi State Universtiy, Mississippi State, Mississippi; 2 University of Tennessee, Knoxville, Tennessee. Variation in nesting success
along an elevational gradient for three late-successional hardwood
songbird species.
In the Southern Appalachians three abundant songbird species, the Ovenbird (Seiurus aurocapillus), Black-throated Blue Warbler (Dendroica caeru-

324

Abstracts

lescens), and the Wood Thrush (Hylocichla mustelina) were all surveyed
for nesting success along an elevational gradient. All three species show a
preference for late-successional Oak-Hickory stands and show similar
range of distributions with respect to elevation. However, within this range
each species shows a preference for elevation with the Wood Thrush preferring low elevations, Ovenbirds preferring mid-elevations, and Blackthroated Blue Warblers preferring high elevations. Preliminary nest success
data in this region appears to shed light on this distribution. Three 15 ha
study plots located along an elevational gradient ranging from 900m to
1300m and were surveyed May through July of 2002 to determine nesting
success of these three songbird species. Mayfield estimates yielded differing patterns in nesting success with respect to each species and elevation.
The Wood Thrush showed greatest nesting success, 62%, at 900m in elevation, with nest success decreasing with increasing elevation, 46% at
1100m, and 35% at 1300m. The Ovenbird peaks in nesting success at midelevations with nest success of 40% at 900m, 62% at 1100m, and 35% at
1300m. Finally the Black-throated Blue Warbler exhibits highest success
at upper elevational plots, 20% at 1100m, and 31% at 1300m. This data
suggests that when implementing conservation programs it may be important to take into account the elevational gradient of the habitat.
STOHLGREN, THOMAS J.,1,* DAVID T. BARNETT,2 CURTIS H.
FLATHER,3 PAM FULLER4 and JOHN KARTESZ.5 1 USGS Fort Collins
Science Center, Fort Collins, CO; 2 Natural Resource Ecology Lab, Colorado State University, Fort Collins, CO; 3 USDA Forest Service, Fort Collins, CO; 4 USGS Caribbean Science Center, Gainesville, FL; 5 Biota of
North America Program, University of North Carolina, Chapel Hill, NC.
Species diversity and the patterns of invasion of multiple biological
groups in the United States.
We document the broad-scale patterns of invasion of multiple biological
groups to begin to set priorities for prevention, early detection, containment, and restoration. Very strong positive correlations exist between native
and non-native plant species richness in 48 of 50 states using county-level
data. At statewide scales, non-indigenous fishes, and snails and mollusks
had similar invasion patterns and were significantly positively cross-correlated with native and non-native plant species richness (r values .0.4, P
.0.05). County level data showed that the proportional abundance of nonindigenous birds was associated with crop production and human populations. We show how native species richness declines with latitude and elevation, while non-native species more closely track human populations
(r.0.56, p,0.001).
STOKES, THOMAS A.,1,* LISA J. SAMUELSON,1 MARK COLEMAN,2
JOHN STANTURF3 and TED LEININGER.4 1 Auburn University, stoketa@auburn.edu, Auburn, AL, USA; 2 USDA Forest Service, Aiken, SC,
USA; 3 USDA Forest Service, Athens, GA, USA; 4 USDA Forest Service,
Stoneville, MS, USA. Water relations of Populus deltoides in response
to varying nutrient and water availability.
We tested the hypothesis that increased nutrient and water availability
would increase total plant hydraulic conductance in three-year-old Populus
deltoides clones. Sap flux density and leaf water potential were studied in
response to resource availability and clonal source (MS clone and east TX
clone) at two different research installations: one in Aiken, SC and the
other in Stoneville, MS. Resource availability was manipulated by irrigation and fertilization and the same clones were planted at each site. Aboveground primary productivity and LAI were increased by increasing resource availability at both sites, and production and LAI were two times
greater at the MS site than the SC site. Height and diameter averaged 5.5
m and 5.75 cm, respectfully, for the SC site while for the MS site average
height and diameter were 10 m and 9.4 cm, respectfully. Average maximum
seasonal LAI was 2 m2 m-2 at SC and 4 m2 m-2 at MS. Although stand
development differed between sites, sap flux density by clone and treatment
were similar between sites and ranged between 60 and 100 g m-2 s-1. Predawn and midday leaf water potentials were similar between clones, treatments and sites, and midday water potential averaged -1.5 MPa. Cumulative stand-level transpiration was 225 mm in SC stands versus 500 mm in
MS stands. Total plant hydraulic conductance did not vary with resource
availability and was on average 3 mol m-2 s-1 MPa-1 in SC trees and 7 mol
m-2 s-1 MPa-1 in MS trees. These results indicate that resource availability

had a greater influence on growth than whole tree water relations and that
total plant hydraulic conductance may increase with increasing plant size
in Populus deltoides.
STONE, PETER A.,1,* MICHAEL S. ROSS,2 PABLO L. RUIZ,2 DAVID
L. REED2 and GAIL L. CHMURA.3 1 South Carolina Dept. Health and
Environmental Control, stonepa@dhec.sc.gov, Columbia, SC; 2 Florida International University, Miami, FL; 3 McGill University, Montreal, Quebec,
Canada. Organic-sediment stratigraphy of elongated tree-islands, Everglades National Park.
Larger tree-islands in freshwater peat marsh of the southern Everglades are
distinctly elongated parallel to sluggish overland flow, occupy slight ridges
of organic sediment (peat at the top), and are usually focused at their
upstream end on a sediment-buried bedrock mound. Six tree-islands were
cored to limestone bedrock to examine stratigraphy of accreted organic
sediments and reveal the depositional history. Archeologically modified
organic soil and debris comprised the sediments present on the bedrock
mound. Stratigraphies beneath the long downflow "tails" were much more
complex than the deeper peats typically found in elongated tree-islands of
the northeastern Everglades. Southern Everglades profiles commonly included inter-layering of peat (fibrous organics), muck (fine grained organicand-mineral mud), and in places apparent ash from organic-soil fires, sometimes all lying above basal marsh marl (calcareous silt precipitated by algae
in seasonally flooded marshes). Establishment of organic-sediment ridges
thus appears to post-date and succeed the development of the Everglades
marsh itself in these specific geographic positions. The abundance of finetextured siliceous mineral matter of the muck was not anticipated, given
the lack of an obvious source, the very low current velocities, and the rarity
of similar constituents in the surrounding marsh peats. The muck and
muck-peat mixtures found at mid-levels in these profiles suggest that physical transport of solid material has been important in the formation of ridges, and thus probably in the wider Everglades system. An alternative mechanism, i.e., in which ridge development depends exclusively on enhanced
plant production and peat sedimentation resulting from nutrient delivery
(perhaps dissolved) from the upstream "head" into a low-nutrient-status
marsh, thus seems insufficient to explain the low, forested ridges of southern Everglades tree-islands. Mineral, chemical, and pollen content of the
muck constrain its likely origin.
STOVER, DANIEL B.,1,* FRANK P. DAY,1 JOHN J. DILUSTRO2 and
BERT G. DRAKE.3 1 Old Dominion University, Norfolk, VA; 2 Savannah
River Ecological Laboratory, Aiken, SC; 3 Smithsonian Environmental Research Center, Edgewater, MD. Effects of elevated CO2 on root turnover
and mortality in a Florida oak-scrub ecosystem.
Fine root dynamics, such as mortality and turnover, influence sequestration
of carbon in the soil. Measurements of the effects of elevated CO2 on fine
root dynamics were conducted in an oak-scrub ecosystem at Kennedy
Space Center, Florida. Minirhizotron tubes were utilized within ambient
(350 ppm) and elevated (700 ppm) open top chambers along with reference
controls. Fine root productivity, mortality and turnover were analyzed and
compared to results five years earlier. In the second year of the study, CO2
enrichment resulted in higher fine root turnover (0.0266 compared to
0.0175 fractional root loss/day in the ambient), higher root production
(0.0939 compared to 0.0524 mm/cm/day in the ambient) and higher mortality (0.0574 compared to 0.0231 mm/cm/day in the ambient). The effect
of elevated CO2 on fine root abundance was reduced to non-significant
levels after seven years of treatment. Preliminary evaluations of mortality
and turnover in the seventh year of the study have yielded inconclusive
results to date with regard to CO2 treatment effect. Disappearance of all
treatment effects would suggest convergence of fine root systems among
treatments, but similar root abundance in ambient and elevated could be
masking higher turnover rates of fine roots in elevated chambers.
STOY, PAUL C.,1,* GABRIEL G. KATUL,1,2 RAM OREN,1 MARIO B.S.
SIQUEIRA1,2 and JEHN-YIH JUANG.1 1 Nicholas School of the Environment and Earth Sciences, Duke University, Durham, NC, USA; 2 Department of Civil and Environmental Engineering, Duke University, Durham,
NC, USA. Carbon dioxide and water vapor exchange in three adjacent
ecosystems for two drought-impacted years.
Understanding the impacts of vegetation cover on biosphere/atmosphere
exchanges of carbon and water is a research priority for ecosystem ecology

and management given the close coupling of terrestrial ecosystems and

climate. We explored differential impacts of vegetation on CO2 and H2O
dynamics with two years of eddy-covariance measured CO2 and H2O fluxes
in three adjacent ecosystems in the Duke Forest, NC: a loblolly pine (Pinus
taeda) plantation, an oak-hickory dominated hardwood forest, and a grass
field. All three sites experience identical climatic and edaphic conditions.
We examined three hypotheses: 1) land cover has a greater impact than
environmental drivers on net ecosystem carbon exchange (NEE), 2) environmental drivers have greater impact on evapotranspiration (ET) than land
cover, and 3) more diverse ecosystems are more resistant to environmental
perturbations. Our primary argument is that H2O fluxes are primarily controlled by energy supply while CO2 fluxes are much more dependent on
species-specific structure and function, including drought resistance. The
carbon and water dynamics of the hardwood forest showed clear functional
resistance to mild and severe drought, presenting a management option that
is robust to the various environmental perturbations encountered during the
measurement period (and for future climate projections for the Southeast).
The pine plantation was sensitive to drought and severe weather, but assimilated carbon at a higher rate under drought-free conditions. The grass
field represented a net source of CO2 to the atmosphere during both years
of measurement. These results further highlight the dramatic differences in
ecosystem function across heterogeneous landscapes, and pose important
challenges for understanding ecosystem response to projected global changes.
STRAND, ALLAN E.,* DAVID M. COUILLARD and KATHRYN J.
NILES. College of Charleston, Charleston, SC. Using genetics to differentiate among models of plant movement:lessons from the beach annual Cakile edentula.
A central component to understanding changing landscape-level demographic processes is dispersal. It is required to augment sizes of existing
populations and for colonization of new or extirpated habitat. Fully understanding the role that dispersal plays at the landscape level requires an
understanding of both the rate and pattern of movement. Unfortunately,
characterizing dispersal ecologically is logistically difficult. Genetic markers provide an alternative approach to characterizing both the rate and pattern of dispersal. To demonstrate this genetic approach in a Southeastern
plant species, we characterized chloroplast DNA-based genetic variation in
14 populations of the beach annual Cakile edentula var. Harperi. Chloroplasts typically are inherited maternally and provide a means to characterize
seed dispersal. Based upon patterns of chloroplast DNA variation, Cakile
populations in this region appear to be exchanging seeds at rates previously
unreported in naturally dispersed plants. Furthermore, there is evidence that
a stepping-stone model of dispersal best fits these data. Because it occupies
essentially continuous habitat and disperses well, changes in climate are
unlikely to affect colonization ability in this species. Estimates of nearfuture sea-level rise, however, suggest that suitable habitat will disappear
and reduce overall numbers of populations.
STRANGE, JAMES P.,1,* LIONEL GARNERY2 and WALTER S. SHEPPARD.1 1 Department of Entomology, Washington State University, Pullman, WA, USA; 2 CNRS Laboratoire Populations Genetiques et Evolution,
Gif-sur-Yvette, Ile-de-France, France. Genetic characterization of honey
bee (Apis mellifera L.) drone congregation areas above the forest canopy in southwest France.
The honey bee (Apis mellifera L.) mates in spatially and temporally defined
areas remote from the colony. Male honey bees gather in these areas (drone
congregation areas or DCAs) and fly about waiting for a virgin queen bee
to enter the DCA. Because these areas can occur up to 30 meters above
the ground, they have been difficult to study. In fact, previous studies have
focused primarily on DCAs that occur at very accessible locations (fields
or forest clearings) and the existence of A. mellifera mating areas above
forest canopies has been largely unknown and, therefore, ignored. Here we
report the presence of two DCAs located above dense pine forest canopy
in southwest France and contrast the genetic composition of these mating
sites with nearby DCAs which occur in fields and forest clearings. Genetic
analysis of drone honey bees trapped at these sites using mitochondrial
DNA shows that honey bees from two subspecies lineages (one native and
one introduced) utilize these sites. Preliminary results show a partial tem-

Abstracts

325

poral partitioning of the DCAs between the two groups. Implications for
past and future studies are discussed.
STRATFORD, JEFFREY A.,1 W. DOUGLAS ROBINSON1,2 and PHILIP
CHANEY.1 1 Center for Forest Sustainability, Auburn University, Auburn,
AL; 2 Oregon State University, Corvallis, OR. Distribution of bird diversity across a gradient of urbanization: The West Georgia project.
Spread of human cities into rural areas modifies the distribution and abundance of major vegetation groups. This shift in habitat availability modifies
the landscape-level characteristics of avian communities. We investigated
how bird diversity varied along a gradient from an urban center, Columbus,
Georgia, north into rural areas of the piedmont physiographic region.
Breeding bird survey-style point counts (n5550) were used to enumerate
species richness and diversity. We used landsat imagery to characterize
major habitat configurations at each point and evaluated how habitat and
bird diversity changed across the gradient. Three species of conservation
concern have been nearly eliminated, occurring only in small remnant populations far from urban areas. Neotropical migratory bird richness changed
as a complex function of multiple variables, but was most influenced by
total forest cover (positive effect) and amount of edge (negative effect)
within 200 m of census points. Urban and suburbanizing landscapes were
dominated by a few, widely distributed species and by three non-native
species. Although bird species richness is strongly influenced by habitat
changes as a function of urbanization processes, total richness and diversity
in this landscape is only 65% of predicted pre-European settlement values.
Large-scale landscape modification has already taken place, leading to a
drop in regional bird diversity. A subsequent major decline occurs when
landscapes become urbanized. Without major changes in how urban landscapes are designed avian species richness will be reduced by more than
40%, largely through the local extinction of habitat specialists such as many
Neotropical migrant species.
STRECKER, ANGELA L.* and SHELLEY E. ARNOTT. (strecker@
biology.queensu.ca) Queens University, Kingston, Ontario. The effects of
an invasive invertebrate predator, Bythotrephes longimanus, on zooplankton communities recovering from acidification.
Freshwater ecosystems worldwide are endangered by the stresses imposed
by human activity. During the past century, sulphur dioxide emissions resulted in the acidification and biological impoverishment of thousands of
lakes in North America. International agreements have resulted in emission
reductions, improvements in lake pH, and subsequent biological recovery
of some lakes. Biological recovery, however, may be influenced by additional stresses, such as the invasion of exotic species. Bythotrephes longimanus is a large predatory cladoceran that invaded the Great Lakes in the
1980s from Eurasia. It has since spread into many Canadian and American
inland lakes. A field experiment was conducted to test the effects of the
invasion of Bythotrephes on recovering zooplankton communities in Killarney Wilderness Park near Sudbury, Ontario. Experimental enclosures
were deployed in Kakakise Lake and consisted of 1-m diameter, 8-m deep
clear plastic bags, suspended from a wooden frame at the lake surface. The
experiment had two treatments: Bythotrephes (presence, absence) and zooplankton community (recovered, non-recovered). Total zooplankton abundance decreased in both the recovered and non-recovered enclosures that
received the Bythotrephes treatment. Several acid-tolerant and acid-sensitive species were differentially affected by the invasion of Bythotrephes,
depending on what stage of recovery the zooplankton community was in.
This suggests that Bythotrephes may impede recolonization of some acidsensitive species and may decrease some acid-tolerant species which sustain zooplankton community abundance in acidic conditions. These results
will have important implications for the management of recovering lakes
and invasive species by providing insight on the effects of an invasion as
an additional obstacle to biological recovery.
RAN ENGLUND1 and LARS ERICSTRENGBOM, JOACHIM,1,2,* GO
SON.1 1 Department of Ecology and Environmental Science, Umea University, SE-90187, Umea, Sweden, Sweden; 2 Present address: Department
of Forest Resources, University of Minnesota, 1530 Cleveland Ave N, MN
55108, St. Paul, Minnesota, USA. Experimental scale and precipitation
modify effects of N-deposition on a plant pathogen.
When logistical and financial factors limit the spatial scale used in ecological experiments, it is important to know how such simplifications affect

326

Abstracts

the generality of the results. Increased disease incidence on bilberry (Vaccinium myrtillus) of the parasitic fungus Valdensia heterodoxa has previously been found to play a key role for N-induced changes in species
composition changes in boreal forests in Sweden. The fungus mediates
vegetation changes by defoliating the dominant species V. myrtillus. This
enables the grass Deschampsia flexuosa to increase in abundance. We studied the potential effect of a spatial scale-dependent interaction between the
parasite and V. myrtillus in a N fertilization experiment in northern Sweden.
Plot sizes included in the experiment were 1, 10, 100 and 1000 m2. Disease
incidence by the fungus was, on average, higher in large plots with high
N treatment than in small plots with low N treatment. The difference in
disease incidence between small and large plots increased between 1998
and 2000. Potential mechanisms that may cause the observed scale dependence include a) scale-dependent dispersal of the fungus and b) a combination of scale-dependent heterogeneity in soil moisture and a nonlinear
growth function of the fungus. Disease incidence during 1996-2000 showed
a strong positive correlation to summer precipitation and high precipitation
enhanced the effect of nitrogen. Climate models suggest that global warming will lead to increased summer precipitation in Northern Scandinavia.
Thus an important implication of our result is that global climate change
may enhance positive effects of N-deposition on the incidence of the pathogen and, indirectly, on plant species composition. Most certainly, spatial
scale-dependence also exists in a large number of other experiments and a
greater awareness of this issue may be needed in order to enable accurate
predictions.
STRONG, DONALD R.,1,* HEATHER G. DAVIS,2 DEBRA AYRES,1
CAZ TAYLOR,3 JANIE CIVILLE3 and JOHN LAMBRINOS.4 1 Section
of Ecology and Evolution, University of California, Davis, Davis, CA,
95616; 2 Population Biology Graduate Group, University of California, Davis, Davis, 95616, CA; 3 Ecology Graduate Group, University of California,
Davis, Davis, CA, 95616; 4 Dept. of Environmental Science and Policy,
University of California, Davis, Davis, CA, 95616. Ecological and Evolutionary Misadventures of Alien Spartina alterniflora in the Pacific.
This is a tale of two estuaries being invaded by smooth cordgrass, Spartina
alterniflora, a native of Atlantic saltmarshes that substantially affects organisms, hydrology, and the commercial uses of Pacific estuaries where it
invades. With no seedbank, it disperses almost exclusively by seed-of-theyear that floats on the tide. In Willapa Bay, WA, far north of native cordgrass, S. alterniflora colonizes mudflats that previously had no emergent
vegetation, converting them to cordgrass meadow and resulting in the loss
of habitat for native species, fishing, and mariculture. Spread of this invasive has been roughly exponential since its introduction 100 years ago,
and it is now present throughout the 43 km long Bay. The spread has been
attenuated by an Allee effect in which colonies at the leading edges of the
invasion set little viable seed, probably for want of outcross pollen from
too-distant neighbors. Recruitment in some years is low. In San Francisco
Bay, S. alterniflora has hybridized in the last 25 years since its introduction
with the native California cordgrass, S. foliosa. Extensive hybrid swarms
are sweeping parts of the Bay, and both the native and alien parents have
virtually disappeared from these areas. Hybridization is reciprocal, and both
parental species have been seed parents to hybrids. Pollen swamping of
natives by hybrids probably drives spread of the hybrid swarm. Hybrids
have higher fitness than either parent, and accelerating hybridization threatens the very existence of California cordgrass.
STUART, GLENN,2 CORINNA GRIES1 and DIANE HOPE.1,* 1 Center
for Environmental Studies, Arizona State University, Tempe, AZ, USA;
2
Department of Anthropology, Arizona State University, Tempe, AZ, USA.
The relationship between pollen and extant vegetation across an arid
urban ecosystem and surrounding desert.
A probability-based sample was used to determine the relationship between
pollen counts in surface soil and extant perennial vegetation, across the
Central Arizona - Phoenix region, including the urban area, surrounding
agricultural and native Sonoran desert land. We asked whether taxa behaved as expected from known pollination characteristics (e.g. anemophilous versus zoophilous) and whether the pollen:plant relationship differed
between the undeveloped desert and agro-urban regions. We mapped pollen:plant abundance across the study area and applied two statistical ap-

proaches to examine the data quantitatively. Firstly, the sum of total pollen
concentration divided by total cover computed for each taxon, was used to
compare pollen productivity of different taxa using a cluster analysis (K
means, normalized). Secondly PC-Ord was used on a cross matrix containing the presence/absence of both pollen and plants for each taxon at
each site. Both techniques produced the same broad groupings. Group I
contained wind-dispersing taxa with high pollen counts relative to the
amount of vegetative cover (e.g. Pinus, Cupressacea). Groups II and III
were comprised of largely native taxa for which both pollen and plants
were typically found at the same sites, the former having a more widespread abundance (e.g. Asteraceae, Ambrosia) than the latter (e.g. Larrea,
Parkinsonia, Prosopis). Groups IV and V were comprised of taxa with
relatively low pollen and plant cover across the region, taxa in Group V
(e.g. Citrus) having a higher overall plant abundance than in group IV (e.g.
Fraxinus, Acacia). Group VI consisting of zoophilous taxa with modest
plant cover but very little pollen in samples and included most of the
Cactaceae. These finding have important implications for allergy-related
pollen forecasting techniques, in particular that significant variation can
occur in pollen rain across a city, even for taxa with very mobile pollen.
STYRING, ALISON R.* Louisiana State University Museum of Natural
Science, Baton Rouge, Louisiana. Patterns of ecological and morphological diversity in temperate and tropical woodpeckers.
Ecological and morphological patterns of diversity were investigated in
woodpeckers from tropical sites in Peninsular Malaysia (Pasoh Forest Reserve and Sungai Lalang Forest Reserve) and Guatemala, and two temperate sites in North America. Multivariate analyses separated species into
two ecomorphs: conventional - species that excavated frequently and had
large bills, long bracing tails, and relatively short toes, and novel - species
that used a variety of microhabitats (e.g. bamboo, ant nests, and broadleaf
foliage) and had relatively short tails, short, flattened bills, and long toes.
Both temperate and tropical species were classified as the first ecomorph,
but the novel ecomorph comprised primarily tropical species. These
woodpeckers used tropical resources not available year-round in temperate
forests such as arboreal ant and termite nests, bamboo, and broadleaf foliage. These novel resources may explain, in part, the maintenance of high
woodpecker diversity in tropical rainforests.
STYRSKY, JOHN D.* and MICKY D. EUBANKS. Auburn University,
Auburn, AL. An investigation of factors that influence the status of the
red imported fire ant as a keystone predator.
Despite considerable evidence for the existence and broad impacts of keystone species in biotic communities, the ecological factors that determine
how and why a species functions as a keystone, particularly in terrestrial
systems, are not well known. The Red Imported Fire Ant (Solenopsis invicta Buren) is an abundant predator with strong and pervasive effects in
agroecosystems throughout the southeastern United States. Recent evidence, however, suggests that the community-level impact of fire ants
varies in different crops (higher in cotton, lower in soybean), probably
because fire ants are more abundant in the canopy of cotton plants than
they are in soybean plants. We discuss two hypotheses to account for this
pattern: 1) fire ants are attracted to honeydew-producing aphids in cotton
whereas no aphid pest is present in soybeans to draw ants onto the plants
(yet), and 2) dense trichomes on soybean plants inhibit fire ant activity
whereas no glandular trichomes are present on cotton plants. While waiting
for the recently detected soybean aphid to invade the Southeast from the
Upper Midwest to test the aphid hypothesis, we conducted a field experiment with different varieties of soybeans that ranged in trichome density
(glabrous, intermediate, and dense) in the presence and absence of fire ants
to test the trichome hypothesis. Neither trichome presence nor density affected fire ant abundance on soybean plants, but the presence of fire ants
did slightly reduce the abundance of pests (lepidopteran larvae) and natural
enemies (big-eyed bugs, damsel bugs). Lepidopteran larvae and big-eyed
bugs were more abundant on soybean varieties with trichomes, regardless
of whether fire ants were present or not, suggesting a variety preference
by caterpillars and a numerical response by bugs. Other natural enemies
(spiders) were unaffected by fire ants or trichomes. Our results, therefore,
are inconsistent with the hypothesis that the inhibition of fire ant activity
on plants by trichomes prevents strong impacts by fire ants on soybean
pests and natural enemies.

SU, BO,* SHIQIANG WAN, ASFAW BELAY, DAFENG HUI and YIQI
LUO. University of Oklahoma, 770 Van Vleet Oval, Norman, OK, USA.
Reducing carbon substrate affects nitrogen availability in relation to
soil respiration in a tallgrass prairie.
Carbon (C) substrate is the major energy source for microorganisms mediating C and nitrogen (N) cycles. However, little is known about how
reducing C substrate affects N cycling and C-N interactions in tallgrass
prairie. By reducing C substrate input via shading, clipping and shading+clipping, we examined C reduction effect on soil inorganic N concentrations, net N mineralization and nitrification rates in relation to soil respiration in a tallgrass priaire ecosystem from July 2001 to July 2002. Results showed that shading/clipping decreased soil respiration, but increased
or tended to increase inorganic N concentrations on most sampling dates.
Shading and shading+clipping tended to increase net N mineralization and
nitrification during most incubation periods, but clipping tended to reduce
net N mineralization and nitrification during the last four incubation periods. Consequently, shading and shading+clipping increased but clipping
reduced annual net N mineralization and nitrification. The increased N
availability under shading and shading+cliping treatments is primarily due
to reduced plant N uptake, N immobilization and increased microbial turnover caused by the more favorable microclimates under shading. Seasonally, inorganic N concentrations and net N mineralization and nitrification
rates were positively related to microbial biomass C/N ratio, but negatively
related to soil respiration. Multiple regression analysis showed that without
soil respiration included, soil moisture and temperature altogether explained
18% and 36% of the seasonal variations in NH4+-N and NO3--N concentrations, respectively, but could not explain any seasonal variations in N transformation rates among treatments. With soil respiration included, however,
45%, 37%, 37% and 38% of the seasonal variations in NH4+-N, NO3--N,
net N mineralization and nitrification were explained, respectively. These
results suggest that although microclimates more or less affected inorganic
N concentrations, both inorganic N (esp., NH4+-N) concentrations and N
cycling rates were mainly determined by C substrate input as reflected by
soil respiration in tallgrass prairie.
SUAZO, ALEXIS A.,1,2,* ANGELIQUE T. DELONG,2 ALICE M. BARD1
and DONNA M. ODDY.3 1 Department of Environmental Protection, Apopka, Florida, USA; 2 University of Central Florida, Orlando, Florida, USA;
3
Dynamac Corporation, Titusville, Florida, USA. Weight loss in species
of the Old Field Mouse in inland and coastal Florida.
The purpose of this study is to investigate weight loss based on repeated
trapping of Peromyscus polionotus sp. and to determine if the weight loss
is of statistical significance. As a consequence of the trapping experience,
it has been hypothesized that small mammals, which are repeatedly captured within a short interval, will lose body weight. This tendency has been
observed in Sigmodon hispidus, Microtus ochrogaster, Microtus pennsylvanicus, and four species of small mammals from Europe and Africa. The
Anastasia Island Beach mouse (Peromyscus polionotus phasma,) at Anastasia State Park, St. Augustine, FL was trapped for two days every three
months for seven years. Mean weight loss was analyzed using a two-tailed
t-test (p,0.001). The Southeastern Beach mouse (P. p. niveiventris) at
Cape Canaveral, Titusville, FL was trapped for three days every 3 months
for two years. Analysis of variance (p50.0124) supports our contention
that repeatedly captured mice will generally lose a small amount of their
body weight. The Old Field mouse (P. p. rhoadsi) at Lake Louisa State
Park, Clermont, FL was trapped for three days every week for five months.
Preliminary results show a similar negative trend in weight loss as observed
in the coastal subspecies of beach mice. Analysis of variance will be used
to determine if this trend is of statistical significance.
SUBERVIELLE, STACY and JEROME J. HOWARD.* Department of Biological Sciences, University of New Orleans, New Orleans, LA, USA.
Light and moisture limitation of Sapium sebiferum seedlings in bottomland hardwood forest.
The invasive tree species Sapium sebiferum (Chinese tallow) is common
throughout the southeastern United States, although most work to date has
occurred in high light environments such as coastal prairie. We investigated
light and moisture limitation of germination and survival of seedling tallow

Abstracts

327

328

Abstracts

Blackbrush (Coleogyne ramosissima Torr.) is a desert shrub that grows in

a defined mid-elevational band in the southwestern United States. Seeds
gathered from populations at varying elevations in the Mojave Desert and
Colorado Plateau were germinated and grown at 158C. The grown tissue
was placed in calorimeter ampules, and the metabolic heat rate (Rq) was
measured at several temperatures. Then a vial containing NaOH was placed
in each ampule to measure the rate of CO2 evolution (RCO2) at each temperature. Rates were measured at temperatures from 5 to 358C. The measurements were used to calculate metabolic efficiency and to predict specific growth rates. Preliminary results indicate optimum temperatures for
metabolism and growth range from 10 to 358C, showing differential adaptation among populations.

SUN, OSBERT J.,* JOHN CAMPBELL, BEVERLEY E. LAW and VERNON WOLF. Department of Forest Science, Oregon State University, Corvallis, Oregon, USA. Factors controlling soil carbon storage in the coniferous forests of Oregon, USA.

A
W

Residential development is increasing in many rural landscapes throughout

the US, yet the impacts of such development are poorly understood. Residential development is often concentrated near aquatic ecosystems, and
understanding terrestrial-aquatic interactions may be critical for assessing
ecological effects of development. Introduction of wood from riparian forests into the littoral zone, where it provides important habitat for fish, is a
key terrestrial-aquatic interaction that is not well understood. Increasing
development could alter the amount and quality of littoral habitat through
direct removal of coarse woody habitat (CWH) by humans and through
reduced input rates if homeowners thin riparian forests. We sampled the
shorelines of 30 lakes in Vilas County, Wisconsin, USA, that ranged in
building density from zero to 38 houses/km of shoreline. At eight sites per
lake, we measured the intensity of land use; the abundance of littoral CWH
along a 50-m transect at 0.5-m water depth; and the composition, structure
and amount of living trees and CWH in three 100-m2 plots of riparian

SUMMERS, HEIDI, BRUCE SMITH and LEE HANSEN. Brigham Young

University, Provo, UT. Metabolic response to temperature in blackbrush (Coleogyne ramosissima).

We investigated soil carbon storage across a steep gradient in precipitation

in Oregon, USA, in chronosequences of the Sitka spruce (Picea sitchensis
(Bong.) Carriere) forest type near the coast, the montane Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) forest type of the western Cascade
Range, and the ponderosa pine (Pinus ponderosa var. Laws.) forest type
typical of central Oregon. Soil carbon storage varied from 36 kg C m-2 in
Sitka spruce, 14 kg C m-2 in Douglas-fir, and 7 kg C m-2 in the ponderosa

SUGDEN-NEWBERY, ANNA E.,* MONICA G. TURNER, TIMOTHY

K. KRATZ and STEPHEN R. CARPENTER. University of WisconsinMadison, Madison, WI. Effect of lakeshore development on riparian
forest and littoral woody habitat.

Fort Stewart is an 113,089-hectare U.S. Army Installation located on the

Lower Coastal Plain of Georgia. The Canoochee and Ogeechee Rivers
(Ogeechee River Basin) flow through the installation, and thousands of
hectares of wetlands within their watersheds dominate the Fort Stewart
landscape. The waters of the installation have never been systematically
surveyed for freshwater mollusks. During the summers of 2001 and 2002,
79 sites across the installation were surveyed revealing eleven species of
native freshwater Unionids. Two additional Unionid species were observed
immediately adjacent to Fort Stewart waters. In addition, the exotic Asiatic
clam, Corbicula fluminea, and eight species of freshwater gastropods were
also observed. Timed searches were conducted to assess relative abundance
of mussel populations. Preliminary survey results reveal a sparsely populated mussel community with limited distribution and low diversity. High
Unionid abundance was limited to three locations: a low order stream reach
below the Hinesville/Fort Stewart Regional Water Pollution Control Plant,
a series of old abandoned borrow pits within the floodplain of the Canoochee River, and an oxbow slough on the Ogeechee River. Mussel abundance and diversity appeared to be positively influenced by alkalinity and
pH. Increasing levels of total phosphorus appeared to favor mussel abundance up to 6 mg/l, beyond which mussel counts declined. The Asiatic
clam was observed at all sample sites, except those occurring in a first
order stream segment within the Altamaha River basin, located in the
southwestern end of the installation. Results of this survey will provide
environmental managers with the necessary data to better coordinate military mission activities to protect the aquatic resources of Fort Stewart.

H
D

Relationships between plant traits and competitive ability are assumed to

have important consequences for community structure and function. We
ask whether traits (e.g., seed mass and potential growth rate) are related to
measures of competitive performance (the effect of experimental reductions
of neighbor biomass on demographic parameters such as emergence,
growth, or survival), and then whether these measures of competitive performance are related to species relative abundance patterns. We examined
relationships in desert annual, old-field, and tallgrass prairie communities.
In desert annual communities, seed mass and potential growth rates were
not related to any measure of competitive ability, and the relative effect of
neighbors on emergence and survival (but not on growth), were the measures of competitive ability that best predicted relative abundance. In oldfields, potential growth rates were negatively related to the effect of neighbors on adult growth, but it was the effect of neighbors on germination
and seedling growth (rather than on adult growth or survival) that were the
measures of competitive ability most correlated with abundance. In tallgrass prairie, seed mass was correlated with species relative abundance;
however, it was tolerance to defoliation and drought stresses, rather than
any measure of competitive ability, that best translated seed size to abundance patterns. These results challenge a general assumption that competitive ability, and particularly that competitive ability in terms of the effect
of neighbors on adult growth, should translate variation in species traits to
patterns of community structure. The influence of other types of interactions in addition to competition (e.g., stress tolerance, herbivory) as well
as other components of competitive performance (e.g., the effect of neighbors on seedling emergence and adult survival) may be critical processes
by which individual traits scale to community structure.

SUKKESTAD, KATHRYN E.,1,* THOMAS D. BRYCE1 and EUGENE P.

KEFERL.2 1 Fish & Wildlife Branch, Fort Stewart, Georgia, USA; 2 Coastal
Georgia Community College, Brunswick, Georgia, USA. The Survey of
Freshwater Mollusks of Fort Stewart, Georgia.

IT

SUDING, KATHARINE N.,1,* DEBORAH E. GOLDBERG2 and TIMOTHY G. HOWARD.3 1 University of California, Irvine, CA; 2 University
of Michigan, Ann Arbor, MI; 3 New York Natural Heritage Program, Albany, NY. Linking species traits to competitive ability and the structure
of plant communities.

forest. Littoral and riparian CWH and riparian forest stand density were
highly variable both within and among lakes. Estimates of the riparian
CWH by lake varied from 25 to 550 logs/ha, whereas littoral CWH varied
from 0 to 280 logs/km of shoreline. Whole-lake estimates of riparian forest
stand densities varied from 375 to 1875 stems/ha. Increased building density had a negative relationship with both riparian and littoral CWH (R2 5
0.547, p , 0.001 and R2 5 0.385, p , 0.001) These results suggest that
CWH reduction is an important pathway through which residential development and human activity may alter terrestrial-aquatic interactions in rural
landscapes.

in a bottomland hardwood forest in Louisiana. We sowed seed and transplanted seedlings of standardized size into replicated plots at the edge and
in the interior of bottomland forest tracts. We also studied light limitation
in shadehouse experiments under two levels of moisture availability. In
field studies, survival and performance were positively associated with
moisture availability and negatively associated with light levels. At high
moisture levels survival began to decline, with peak survival occurring at
soil moisture levels of about 25% by weight. In shadehouse experiments,
survival did not differ among light levels from full sun to 6% of full sun.
Performance measures such as height growth showed no systematic, significant relationship to light levels. Allocation patterns varied significantly,
with much higher root allocation in high-light treatments. The results suggest that moisture is more important than light for initial establishment and
recruitment of Sapium seedlings in bottomland forest, and that this species
has significant potential for establishing and persisting in bottomland habitats.

pine types. The range of values in soil carbon paralleled differences in

precipitation, soil nitrogen concentration, clay content, and annual carbon
transfer in litterfall. Soil carbon storage was inversely correlated with the
mean residence time of forest floor litter carbon. Stand-replacing disturbance in terms of harvesting had no significant impact on soil carbon storage within forest types. Spatial variations of soil carbon storage in Oregon
are affected by precipitation and the rate that litter falls and is transformed
into soil particulate organic matter.
SUNDARESHWAR, P V.,1,* J T. MORRIS,2 ERIC KOEPFLER,3 BRANDON FORNWALT4 and C J. RICHARDSON.1 1 Duke University Wetland
Center, Nicholas School of the Environment and Earth Sciences, Duke
University, Durham, NC, USA; 2 Department of Biological Sciences, University of South Carolina, Columbia, SC, USA; 3 Marine Science Department, Coastal Carolina University, Conway, SC, USA; 4 Marine Science
Program, University of South Carolina, Columbia, SC, USA. Phosphorus
limitation of coastal ecosystem processes - biogeochemical implications
for differential nutrient limitations.
In coastal wetlands, primary production is conventionally thought to be
limited by nitrogen. Through bioassays, we show that although, the plant
community in a pristine salt marsh was found to be limited primarily by
nitrogen availability, the bacterial community in the soil was limited by
phosphorus. Phosphorus limitation of the growth of nitrogen transforming
bacteria can affect carbon fixation, storage and release mediated by plants.
Hence, in coastal wetlands individual trophic groups may respond differently to nitrogen and phosphorus loading. It is likely that differential nutrient limitations observed in marshes also occur in other ecosystems and
probably represent a consequence of ecosystem development that maximizes overall resource utilization and conservation.
SUTTER, ROBERT D.1 and ROBERT L. BENDICK.2 1 The Nature Conservancy, Southeast Divsion, Durham, NC, US; 2 The Nature Conservancy,
Southeast Division, Orlando, NC, US. Challenges for Conserving Biodiversity in the Southeast United States.
The Southeast United States is a diverse ecological region. It includes
spruce-fir forests in the highest mountains in eastern North America, rich
hardwood forests of the Southern Appalachians, unique limestone glades
of central Tennessee and Alabama, longleaf pine savannas and sandhills,
extensive bottomlands of the coastal plain, imperiled wetland prairies and
sand pine scrub of Florida, the only coral reef in the continental United
States, and dozens of productive bays and estuaries. This region contains
23 percent of all terrestrial communities identified in the U.S. including
some of the most endangered ones in the country. Four of the six most
species-diverse states occur in this region and the region is especially rich
in fish, mussels, snails and amphibians. The 1,363 species at-risk in the
Southeast constitute 22 percent of all at-risk species identified in the U.S.
National and regional conservation agencies are struggling to protect this
unique natural heritage in the face of a changing human landscape. These
changes include rapid, sprawling growth, the change in the ownership and
management of forestlands, increased demands on water resources, fire
suppression and the spread of invasive, non-native plants, animals and pathogens. The extensive ecological diversity and escalating threats creates a
challenging environment for conservation. We will discuss the diverse challenges to conservation at several different scales. At the largest scales
(global, national, ecoregional, state) the challenges include selecting sites
and cross-cutting strategies (including government policy) that will have
the greatest conservation impact. At the multi-site scale a primary challenge
is to coordinate efforts across different ownership and management agencies to meet specific conservation goals. The site level includes the challenges of conservation planning, adaptive management, and the interplay
of science and land management. A key concept is the integration of conservation across strategies, ecological systems, institutions and time.
SUTTLE, KENWYN B.* and MARY A. SORENSEN. Department of Integrative Biology, University of California, Berkeley, CA, USA. Topdown effects cascade to plants through pollinators.
Pollinators are a diverse group of organisms that serve a vital role in the
functioning and perpetuation of most terrestrial ecosystems. Given the

prevalence of pollen-limitation in flowering plants, predation on pollinators

may affect not only pollinators, but also their host plants. The potential for
predator effects to transmit through pollinators has received little attention,
however, as our understanding of top-down effects on lower trophic levels
is drawn predominantly from food webs in which herbivores compose the
second trophic level. We show that by disrupting plant-pollinator relationships, predators can inhibit pollination and consequently decrease fruit and
seed set in flowering plants. In a riparian system, crab spiders that hunt
from flowers of the invasive plant Leucanthemum vulgare decreased the
plants seed production by reducing the frequency and duration of floral
visits by pollinating insects. Such pollinator-mediated indirect effects may
be a common feature of food chains with pollinators at the second trophic
level. Their existence has implications for research into plant-pollinator
relationships, biotic resistance to invasion, and the breadth of predator influence in ecological communities.
SWAN, CHRIS* and MARGARET PALMER. Department of Entomology,
University of Maryland, College Park, MD, USA. Non-additive effects of
leaf diversity on litter decomposition: The role of compensatory feeding
by detritivores.
Temperate stream food webs rely heavily on the annual input of leaf material from the riparian zone. The availability of this material as food for
detritivores is described by leaf decomposition rate. We have previously
reported that mixed-species leaf litter can decompose slower than what
would be predicted from averaging single-species estimates. Since streams
harbor assemblages of leaf-shredding invertebrates, and these organisms
impart some control on decomposition by feeding differentially among leaf
species of varying quality, we sought to determine if detritivore feeding
rate explained this pattern. We performed a laboratory experiment where
we offered a leaf-shredding isopod (Caecidotea communis) one of eight
leaf litter treatments: four single- and four mixed-species treatments. The
mixed-litter treatments comprised each of the four possible three-species
combinations. We followed decomposition rate of each leaf species alone
and in mixture under the presence/absence of the isopod. In three of the
four mixed-species treatments, decomposition was slower than the average
of each single-species alone. Consumption of the slowest decomposing
species, American Sycamore, was greater when offered to the isopod alone,
than when Sycamore was present in mixtures with more palatable leaf
species. When this organism is offered a mixture of resources, it avoids
refractory leaf species in lieu of more labile ones. Thus, this organism
compensates for poorer resources present as single-species leaf litter by
increasing its feeding rate. The implications for this result include: (1) since
leaf mixtures in stream ecosystems reflect riparian tree diversity, the loss
of plant species from the riparian zone may result in altered in-stream
organic matter dynamics, and (2) filter-feeding consumers rely on the byproducts (particles, feces) of shredder feeding. If shredder feeding rate is
altered by the change in leaf litter species composition, the trophic transfer
of organic matter from the riparian zone to filter-feeders via this feeding
link may also change.
SWANSON, BROOK O.* Northern Arizona University, bos@dana.ucc.
nau.edu, Flagstaff, Arizona. Competition reduction and the maintinance
of polymorphism in a cichlid, Herichthys minckleyi.
Polymorphic species provide a system to study the evolution and maintenance of morphological diversity. At least two distinct prey processing
morphotypes occur in the Cuatro Cienegas cichlid (Herichthys minckleyi).
One morphotype has gracile pharyngeal jaws with papilliform pharyngeal
teeth. The other morphotype has robust pharyngeal jaws and molariform
pharyngeal teeth. These morphologies are thought to be adaptations for
processing different food items. We conducted two field competition experiments to examine competitive interactions at different densities and test
the hypothesis that the morphotypes reduce intraspecific competition. We
also evaluated how density affects the strength of competition. Competitive
interactions were examined within and between the morphotypes using enclosures. High-density and low-density cage treatments containing a single
morphotype were used to quantify within-morph competition. Cages containing both morphotypes were used to quantify between-morph competition. Results indicate that intraspecific competition is reduced but not eliminated by the presence of two morphotypes in the population. Experiments

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329

at lower densities indicate that resource partitioning is more pronounced

under periods of resource limitation.
SWART, CHARLES C.,* BRUCE E. FELGENHAUER and ROBERT G.
JAEGER. Department of Biology, University of Louisiana at Lafayette, 300
E. St. Mary Blvd., Lafayette, LA, 70504. Temperature effects on lifehistory characteristics of the Giant Waterbug, Belostoma lutarium (Hemiptera: Belostomatidae).
Thermal energy is known to cause variation in reaction rates of most biological processes. Environmental temperature can have drastic effects on
life history characteristics including growth rates, body size, fecundity, and
propagule size. In most species, body size can affect clutch size through
limitation of egg production or egg care by the female. In animals with
paternal care, male body size can also limit clutch size. Males of the semiaquatic heteropteran, Belostomatinae brood eggs on their dorsal surface,
thereby limiting clutch size by the area available due to body size. Males
can also affect body size of the offspring by preferentially occupying warm
or cool microhabitats during brooding and egg hatching. Our results indicate that temperature has a significant effect on adult size, time to maturity,
and egg size, but not growth rate, in the Giant Waterbug, Belostoma lutarium. Furthermore we use data from field measurements and field collected
and laboratory reared animals to develop a model predicting seasonal fitness based on body size, temperature, and time to maturity.
SWEMMER, TONY M.,1,* WILLIAM D. STOCK2 and WILLIAM J.
BOND.2 1 Kansas State University, Manhattan, Kansas; 2 University of
Cape Town, Cape Town, Western Cape, South Africa. Nitrogen nutrition
of grazing-tolerant grasses.
Heavy grazing of grass communities is generally considered to increase
rates of nutrient cycling. This has been shown as increased rates of net
nitrogen (N) mineralization for the grazing lawns of the Serengeti ecosystem (east Africa) and a number of North American grasslands. This study
tested the hypothesis that grazing-tolerant grasses which dominate heavilygrazed communities rely on a high supply of N to cope with frequent
herbivory. The species used are dominant on grazing lawns in the mesic
savanna of the Hluhluwe-Umfolozi Park, South Africa. Grazing lawns here
are similar in structure to those described for the Serengeti, and support a
similar density of large grazers during the growing season. An initial sand
culture experiment revealed no significant effects of N source (nitrate, ammonium or a combination) for three grazing lawn species. To test for the
effects of N amount, these species and one additional grazing lawn species,
were grown in a series of solution culture experiments (with biweekly
clipping). N was supplied as ammonium nitrate at a range of concentration
levels. The four species showed considerable variation in relative growth
rate (RGR) and biomass allocation patterns in response to N concentration.
For five of six pairwise, species comparisons significant differences were
found for maximum RGR and RGR response curves. Many biomass allocation variables, including root mass fraction, leaf area ratio and specific
leaf area, also showed significant differences between species. While
growth responses under controlled conditions do not necessarily reflect responses under natural conditions, these results strongly suggest that grazing-tolerant grasses do not share a similar, high demand for inorganic N.
This calls into question the generality of the accepted pattern of increased
N availability with grazing, and the significance of any such increases for
grazing-tolerant grasses.
SWENSON, JANICE E.,* SARAH E. MOONEY and ANTHONY M.
ROSSI. University of North Florida, Jacksonville, FL, USA. Bacteria in
the fluid of four species of pitcher plants (Sarracenia spp.)
Pitcher plants, Sarracenia spp., along with other carnivorous plants, represent a unique group of organisms because they are simultaneously primary producers and predators. While numerous studies have documented
the eukaryotic micro- and meio-fauna that inhabit the pitchers of Sarracenia spp., few have detailed their bacterial community composition. In
addition, previous studies investigating the microbial community structure
of pitcher plants, especially S. purpurea, have generally sampled only open
pitchers in monospecific populations of Sarracenia. In the current study,
we sampled both opened and unopened pitchers from four species of Sar-

330

Abstracts

racenia, which occur sympatrically on the UNF campus. Although our

investigation is incomplete, preliminary results suggest that bacterial community species richness is influenced by plant species. Suprisingly, samples
of fluid extracted from the unopened pitchers of at least one species, the
hooded pitcher plant (S. minor), using a sterile tuberculin syringe contained
an endogenous bacterial community (although it was less species rich than
opened pitchers). Isolation of resident bacteria may aid in understanding
the evolutionary relationships of these symbiotic microbial communities
and their host plants.
TADA, KAZUFUMI,1 TADASHI HIBINO,1 KAZUO MURAKAMI,2 HIDEO MATSUMOTO3 and KIM CHI TRAN.4 1 Depertment of Social and
Environmental Engineering, Graduate School of Engineering, Hiroshima
University, Higashi-Hiroshima, Hiroshima, Japan; 2 Institute for Marine Resources and Environment, Institute for Marine Resources and Environment
Environmental Hydraulics Research Group, National Institute of Advanced
Industrial Science and Technology, Kure, Hiroshima, Japan; 3 Hiroshima
port airport technical investigation office, Chugoku district maintenance
office, Ministry of Land Infrastructure and Transport, Hiroshima, Hiroshima, Japan; 4 School of Policy Studies, Kwansei Gakuin University, Sanda,
Hyogo, Japan. The basic study on modeling variable fluid mud.
By the inflow load from Ota River and Seto Inland Sea, or by death and
excretion of zooplankton, death of phytoplankton, nutrition salt, and organic matter sediment, a fluid mud layer (high turbidity layer) is formed
near a submarine surface. The purpose of this research is to study the
variable characteristics of fluid mud in Hiroshima Bay. Fluid mud is easily
transported and deposited on stagnated ocean space. It is suspected that
Kure Bay is the stagnated ocean space within Hiroshima Bay. Fluid mud
thickness varied seasonally from 3 cm in May to a peak of 8 cm in September. The wet density of mud was lowest in September. The nutrition
salt in mud is highest in September. The fluid mud changes state physically
and chemically from July to September. In one month of continuous observations during the summer of 2002 in Kure Bay, the flow velocities
were 0-3 cm/s, and turbidity measurements were 4-10 ppm at the seabed.
Sedimentation flux increased with depth. Moreover, it was larger in September than August. In mud, insoluble Fe3+ reduces to Fe2+, and will begin
to dissolve in water. The soluble Fe2+ reacts immediately with hydrogen
sulfide, and FeS is generated, which is responsible for the black color of
the mud. Fluid mud is characterized by the amount of sedimentation and
horizontal inflow, and the chemical change in the mud.
TADROS, MOHSEN 1.,1,* FARRG SHARSHIR,1 FAIEZ ABU-ATTIA1
and GHDA ELSHAFEI.2 1 Tanta University,Fac.of Agric,Kafr el-sheikh,
Tanta University,Fac.of Agric, Kafr el-sheikh, Kafr el-sheikh, Egypt; 2 Tanta University,Fac.of Agric,Kafr el-sheikh, Tanta University,Fac.of Agric,
Kafr el-sheikh, Kafr el-sheikh, Egypt. Soil Acarina and Collembola under two vegetable crops grown under plastic tunnels in Kafr El-Sheikh,
Egypt.
Three plastic tunnels were chosen for carrying on the present investigation
situated in Kafr El-Sheikh area. The first one was 270 m2, the second 357
m2, while the third was 410 m2. The first tunnel (CK) was used to plant
cucumber Cucumis sativus L. var. Delta star, and tomatoes Lycopersicon
esculentum, var. strain B-VF 145B, and was never treated with any toxicants during the experiment that lasted for 2 seasons (1999-2000). The
other two tunnels were grown with cucumber and tomato separately and
were treated with 8 different pesticides for the control of white fly, Aphids,
leaf miners, thrips and the red spider mite. Soil samples were taken on a
monthly basis at random in-between the grown plants, at 8:00 A.M. A steel
cylinder of 4" diameter was used to sample 20 cm deep. The procedure
lasted from October 1999 until April 2000, and samples were taken from
both the CK site and the treated ones. Extraction of organisms took place
for 48 hours. Results revealed in recording 30 mite species of 17 families
and 4 suborders, and were found to fall under 4 acarina sub-orders; Oribatida, Gamasida, Actinedida and Acaridida. The recorded springtails were
of 8 species of 5 families. Soil acarina species under cucumber plantations
were found to exceed its numbers 55.21% than those under tomatoes 44.49.
The springtail Lepidocyrtus cyaneus (Tullberg) was considered a dominant
species. The applied toxicants, were found to decrease soil fauna acarina
and collembola; however, it was more effective under cucumber plants than

under tomatoes, and soil acarina was more sensitive to toxicants than soil
collembola.
TAINTER, JOSEPH A.* Rocky Mountain Research Station, 333 Broadway
SE, Albuquerque, New Mexico, U.S.A. Complexity and Sustainability,
Part I: Social Complexity.
While there has been a tradition within ecology to equate complexity with
diversity and diversity with stability, in human systems these relationships
are quite different. In a human social system, efforts to achieve stability or
sustainability often generate complexity. Complexity in human institutions
arises through efforts at problem solving, with problem solving being frequently directed toward sustaining a valued activity or way of life. This
presentation defines sustainability, resiliency, and collapse in human institutions, and shows that each condition may arise from different strategies
of complexity in problem solving. Historical and contemporary case studies
illustrate principles of sustainability that apply in general to problem-solving institutions, including institutions concerned with natural resource management.
TANEVA, LINA,1 ROSER MATAMALA,2 WILLIAM H. SCHLESINGER3 and MIQUEL A. GONZALEZ-MELER.1 1 University of Illinois at
Chicago, Chicago, IL; 2 Argonne National Laboratory, Argonne, IL; 3 Duke
University, Durham, NC. Responses of autotrophic vs. heterotrophic soil
respiration to elevated CO2: Implications for soil C storage.
Soils are the largest active terrestrial carbon pool, with 2.5 times more C
in the top meter of soil than is found in terrestrial vegetation. Through the
process of soil respiration (RS), soils contribute to an annual flux of CO2
to the atmosphere that is 10 times greater than that of fossil fuel combustion. Because of the size of this flux, even small changes in the rate of RS
could have significant impacts on atmospheric CO2 concentrations. Increasing evidence indicates that rising atmospheric CO2 enhances carbon uptake
in most ecosystems, highlighting terrestrial carbon sinks as an important
factor in mitigating greenhouse gas emissions. However, soils have the
potential to store part of the anthropogenic CO2 only if a substantial proportion of the additional C fixed by plants grown under elevated CO2 enters
soil C pools that turn over slowly. If the additional C is allocated to relatively labile soil C pools, little net C storage may occur. We used stable
isotopes to study the response of RS and its components to elevated CO2
in an intact loblolly pine-dominated forest in North Carolina under FACE.
The depleted 13C signature of the fumigation gas is reflected in new photosynthetic tissue and can be followed into the different soil carbon pools.
The d13C of soil-respired CO2 can be used to separate RS into its autotrophic
and heterotrophic components, and its d18O can further partition heterotrophic respiration. Our data indicate that autotrophic respiration rises under
elevated CO2, suggesting that NEP in this forest may be limited at high
CO2 by increased root and rhizosphere activity.
TANG, JIANWU,* YE QI and ALLEN GOLDSTEIN. University of California at Berkeley, Berkeley, CA. Separating root respiration from soil
respiration in a ponderosa pine plantation in the Sierra Nevada.
Partitioning soil respiration into autotrophic and heterotrophic respiration
is of critical importance for building process-based soil carbon models
since these components respond differently to abiotic and biotic drivers and
have different spatial and temporal variations. To remove the influence of
root autotrophic respiration from total soil respiration, we trenched a 3m x
3m plot in a ponderosa pine plantation in the Sierra Nevada. We measured
soil CO2 efflux in the trenched plot as well as two non-trenched plots
between August 2001 and October 2002. We used multivariate regression
with independent variables of soil temperature and moisture to analyze
measurement data of soil respiration and heterotrophic respiration. We estimated root respiration as the difference between total soil respiration and
heterotrophic respiration. In addition to environmental variables, root respiration is affected by plant physiology, phenology, and photosynthesis.
The annual accumulations of total soil respiration, heterotrophic respiration,
and autotrophic respiration between October 1, 2001 and September 30,
2002 were 78.2 mol m22 year21, 52.2 mol m22year21, and 26.0 mol m22
year21, respectively. Total soil respiration, heterotrophic respiration, and
autotrophic respiration peaked in June. The ratio of autotrophic respiration

to total soil respiration (Fa/F) is not a constant seasonally with an annual

average of 0.33. In the growing seasons between May and October Fa/F
averaged 0.37 while in non-growing seasons Fa/F averaged 0.28. We used
aerial photos, image analysis, and GIS to study the spatial variation of soil
respiration. The spatial variation of soil respiration is mainly explained by
root density, and also influenced by soil nitrogen content and soil carbon
content.
TANNER, SHANN C.,* CHRISTINA E. WELLS, GREGORY L. REIGHARD, DANIEL A. KLUEPFEL and MELISSA B. RILEY. Clemson University, stanner@clemson.edu, Clemson, South Carolina. Isolation and
identification of cyanogenic rhizobacteria from the roots of peach [Prunus persica (L.) Batsch].
Peach replant disorder (PRD) occurs when peach trees are replanted to an
orchard previously cropped with peaches: lower growth rates and yields
are observed in the replanted trees. Methyl bromide fumigation adequately
controls PRD, suggesting that the disorder is the result of harmful interaction(s) between the soil faunal community and peach roots. As methyl
bromide fumigation is phased out, the task of determining the specific cause
of PRD is now an important goal. One theory suggests that prunasin (Dmandelonitrile b-D-glucoside), a cyanogenic glycoside found in peach tissues, may play a critical role in replant disorder. Many soil bacteria have
the ability to degrade cyanogenic glycosides and release cyanide, a potent
respiratory poison. Bacteria capable of degrading prunasin may release cyanide when feeding on decomposing roots of old peach trees, thus inhibiting root growth of the new trees. We examined the rhizosphere bacterial
community of peach to determine whether it contained bacteria capable of
evolving cyanide from prunasin. Rhizobacteria were isolated from the fine
roots of 7-year-old peach trees on M9 carbon source utilization media containing 0.05% prunasin as the sole carbon and nitrogen source. Five visually distinct colonies were selected and grown-out in pure culture on trypticase soy broth agar (TSBA) plates. The prunasin-degrading rhizobacteria
were then tested for their cyanogenic potential using a picrate paper assay.
All five prunasin isolates exhibited some cyanogenic potential; two strains
were strongly cyanogenic. The cyanogenic isolates were identified using
fatty acid methyl ester (FAME) analysis, and the two strongly cyanogenic
strains were identified as Arthrobacter globiformis and Cellulomonas cartae. To determine whether cyanogenic bacterial activity can cause growth
reductions in peach trees, we are inoculating peach seedlings with the cynanogenic bacterial isolates in the greenhouse and monitoring their growth
and root system development.
TAUB, FRIEDA B.* University of Washington, taub@u.washington.edu,
Seattle, WA, USA. Aquatic community metabolism: Effects of toxic
chemicals in microcosms.
Just as the oxygen consumption of an animal can be used to estimate energy
expended, community metabolism can be estimated (based on the heat of
combustion of organic material as ca. 14 kJ per gram O2). Net Ecosystem
Production (NEP), the net energy gain (net photosynthesis) minus (night
respiration) reflects the net daily energy change. I demonstrate that NEP is
a more useful measure than P:R ratio for microcosms treated with three
toxic chemicals. Streptomycin, an antibiotic and selective algicide reduced
net photosynthesis for more than a month; yet, P:R ratios were almost
always near 1 because R decreased a similar amount. Malathion, an insecticide, temporarily eliminated Daphnia; net photosynthesis was increased
during the absence of grazers, but P:R ratios were almost always near 1,
since R was high when P was high. CuSO4, both a selective algicide and
a zooplankton toxicant, had a range of effects depending on concentration;
low CuSO4 concentrations eliminated grazers and sensitive algae but allowed insensitive algae to bloom; at higher CuSO4 concentrations, most
algae as well as grazers were eliminated, and net photosynthesis was reduced; still P:R ratios were usually close to 1. NEP and accumulated NEP
were more useful in understanding the trophic interactions and demonstrating statistical differences from controls.
TAYLOR, BARBARA E.,* DAVID E. SCOTT and J. W. GIBBONS. Savannah River Ecology Laboratory, University of Georgia, Aiken, SC. Risk
of reproductive failure and persistence of the Marbled Salamander
(Ambystoma opacum) in a wetland pond.
For amphibians that breed in wetland ponds, disturbance may often be the
most important factor affecting reproductive success. We analyzed a 24-yr

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331

record of breeding adults and juvenile recruits for the marbled salamander,
Ambystoma opacum, at Rainbow Bay, a seasonal wetland pond on the
Savannah River Site in South Carolina. Complete reproductive failure occurred in 6 of 22 yr when the species bred and near complete failure in an
additional 4 yr. These failures were due mainly to late filling or early drying
of the pond. A computer simulation model for population dynamics of A.
opacum, developed previously for this species on the SRS, enabled us to
infer levels of survival in pond and upland habitat necessary to produce
the observed levels of recruitment and to maintain breeding populations.
We then used the model to explore persistence of the populations when
catastrophic mortality is applied to aquatic stages. The model is age-structured. It tracks terrestrial females individually, so that effects of demographic stochasticity on breeding and death are represented. Catastrophic
reproductive failure has a profound effect on population dynamics, and the
probability of extinction is consequently fairly high for plausible values of
survival in the upland habitat. Details of life history, particularly life span,
become important to predicting persistence. The high probability of reproductive failure makes the population extremely susceptible to local extinction, but the high reproductive capacity also renders it amenable to rescue
by recolonization. These results underscore the importance of understanding and managing dynamics of this species at the meta-population level.
TAYLOR, CAZ M.,* HEATHER G. DAVIS, JANIE C. CIVILLE and
ALAN M. HASTINGS. University of California, Davis, Davis, CA. Estimating pollen dispersal in an invasive grass.
This study investigates the manner in which variation in conspecific density
correlates with variation in seed production in a self-incompatible invasive
plant. Reproductive rates of isolated Spartina alterniflora at the leading
edge of an invasion suffer a more than ten fold reduction in reproductive
rate. This wind-pollinated grass, native to the East and Gulf coasts of North
America, has invaded mudflats of a large estuary in Willapa Bay, Washington, USA that are historically uninhabited by any other emergent plant.
The low reproductive rates suggest that isolated plants are limited in their
ability to make viable seed by pollen dispersal distances. This pattern,
compounded by the self-incompatibility of S. alterniflora, has the potential
to strongly affect the dynamics of this invasion. In order to estimate the
shape of the pollen dispersal density function, we measured seed production
in fifty plants growing at different degrees of isolation and used remote
sensing to map the spatial arrangement of conspecifics in the neighborhood
of each plant. We also obtained data describing the wind speed and direction at the study site. We first explored how much of the variation in seed
production could be explained by simple neighborhood density measures
such as distance to nearest neighbor and area occupied within a known
radius. The latter was used to determine the effective neighborhood of
pollen dispersal. We then used maximum likelihood to compare the effectiveness in explaining the variation in seed production of several isotropic
and non-isotropic spatial models of pollen dispersal.
TAYLOR, DAVID I.,* ROBYN A. DUNMORE and DAVID R. SCHIEL.
University of Canterbury, Christchurch, New Zealand, New Zealand. A
hemispheric comparison of grazing and growth of habitat-forming algae across exposure gradients.
Models of intertidal community structure highlight that the relative importance of processes like grazing changes over environmental stress gradients.
A commonly used proxy for environmental stress is wave exposure. Tests
of these models often use mussel and barnacle assemblages, but are these
relevant for habitat-forming seaweeds? We tested the importance of grazing
in the survival of related seaweeds across wave exposure gradients in
southern New Zealand and Oregon. Fucoid algae can dominate the lower
and mid intertidal zones in New Zealand and in the mid and upper zones
in Oregon. We tested grazing effects on early benthic life-stages (germlings) that are critical to establishment of algal populations but are vulnerable to invertebrate grazers. We transplanted germlings (c. 0.15 mm) of
Hormosira banksii and Durvillaea antarctica in New Zealand and Pelvetiopsis limitata and Fucus gardneri in Oregon in a series of experiments
across wave exposure gradients. The grazing effect on germling survival
was often small (, 5% percent within the first 7 days) but varied considerably and accounted for up to 38 % of mortality at sheltered and exposed
sites. However, no clear gradient in grazing effects was found across wave

332

Abstracts

exposures. Overall, germling survival was low, usually , 20 % after 14

days, even when molluscan grazing was eliminated. Across exposures, sedimentation and ephemeral algae cover affected germling mortality. Growth
differed considerably between species with Durvillaea growing up to three
times faster than the other species within 14 days. It appears that during
this critical establishment phase the direct effects of grazing do not change
across wave exposure gradients, but instead factors like plant characteristics, sedimentation, smothering and desiccation are more important. This
work is part of the PISCO/Mellon project.
TAYLOR, JASON,1,* CHARLENE DAVANZO,2 BRUCE W. GRANT3
and DAN UDOVIC.4 1 Ecological Society of America, Washington, DC,
USA; 2 School of Natural Science, Hampshire College, Amherst, MA,
USA; 3 Department of Biology, Widener University, Chester, PA, USA;
4
Department of Biology, University of Oregon, Eugene, OR, USA. Teaching Issues and Experiments in Ecology (TIEE): A web site and CDROM for undergraduate ecological education.
Teaching Issues and Experiments in Ecology (TIEE) is an NSF-funded,
ESA-sponsored, peer-reviewed, and classroom/ field tested collection of
teaching materials (1) to facilitate inquiry-based Experiments to Teach
Ecology in laboratory courses, and (2) that uses active-learning methods
and current controversial Issues to Teach Ecology for lecture classes. The
TIEE web and CD ROM-based resource collection is not merely a static
site of labs or topics for lecturing, it is an interactive set of new materials
that are both scientifically sound and pedagogically innovative. Our goal
is to have each new peer-reviewed unit meet standards-based learning objectives and use innovative strategies for teaching, assessing growth among
the full diversity of student learners, and improving teaching through formative and summative evaluation. In addition, the peer review aspect of
TIEE submissions and ESAs publication of the accepted educational materials will hopefully stimulate greater participation in ecological educational improvement by faculty who teach ecology. The poster will describe
the pedagogical vision and classroom materials, as well as annotated links
offsite, that are directly linked through keywords from the "Experiments"
and "Issues" sections of the site. The resources provide background explanations and references to the pedagogy and enable faculty to use and evaluate new innovations in education in their classrooms and participate in
rigorous research in ecological education to learn what works and why.
TAYLOR, MILTON D.,1,* JENNIFER A. ZETTLER,2 TIMOTHY P. SPIRA1 and CRAIG R. ALLEN.1 1 Clemson University, Clemson, SC; 2 Armstrong Atlantic State University, Savannah, GA. Experimental introductions of fire ants changes native ant populations.
We experimentally introduced polygyne populations of Solenopsis invicta
into previously uninfested sites to determine the consequences of fire ant
invasion on native ant species. From July 1999 to August 2001, we measured ant species richness, diversity, and abundance from five sites in South
Carolina. At each site, we established one plot into which fire ants were
introduced, and we paired that treatment plot with a control plot in which
colonizing fire ants were baited with Amdrot. All plots consisted of a clearcut and an intact, adjacent forest. By the end of the experiment, fire ant
abundance in treatment sites increased dramatically (17-fold), but they
could not be completely excluded from control sites. However, we found
significant treatment effects in both clearcuts and forests for 15 species in
eight genera that included Aphaenogaster, Brachyponera, Camponotus,
Crematogaster, Forelius, Formica, Pheidole, and Trachymyrmex. We also
found significant seasonal variation in ant abundance for 34 species and
significant differences between abundance in forest and clearcut for 15
species. The introduction of this non-native ant causes significant changes
in ant assemblages within just a few years after fire ant establishment.
Long-term studies are needed to determine the effects of invasion by nonindigenous species on native ant species and its consequences on biological
processes.
TEMPLER, PAMELA H.,* MARY K. FIRESTONE and WHENDEE SILVER. University of California Berkeley, ptempler@nature.berkeley.edu,
Berkeley, CA, USA. Plant and Microbial Controls on Nitrogen Retention and Loss in Puerto Rican Forest Soils.
Understanding the mechanisms that lead to nitrogen (N) retention in tropical forests is important because these systems currently have high potential

for N losses, including the greatest nitrous oxide (N2O) emissions globally
for natural ecosystems, and large potential nitrate losses to groundwater.
The majority of research on N biogeochemistry has been conducted in
north temperate ecosystems where N limitation to net primary productivity
(NPP) is common, and where anthropogenic N deposition is altering N
cycling. Much less research has been conducted in the tropics, which are
generally characterized by rapid rates of N cycling and the lack of strong
N limitation to NPP. We conducted a field experiment using 15N tracers and
root ingrowth cores to compare and contrast the relative importance of N
loss pathways (denitrification, N2O production via nitrification, and N
leaching) versus N retention mechanisms (dissimilatory reduction of nitrate
to ammonium (DNRA) and assimilatory N uptake by plants and soil microbes) in a subtropical forest in Puerto Rico. DNRA rates were detectable
and within a range comparable to gross mineralization and gross nitrification. Plant roots at all sites took up approximately five times more N as
ammonium than nitrate and were a relatively strong sink for N as indicated
by the relatively large amount of 15N that they took up. This supports our
hypothesis that DNRA is a N conserving mechanism within tropical forests
because the ammonium that is produced from this process is likely to be
taken up by plants rather than re-nitrified and potentially leached or lost as
N2O gas from the ecosystem.
TENHUMBERG, BRIGITTE,1,2,* ANDREW J. TYRE,1,2 ANTHONY POPLE1 and HUGH P. POSSINGHAM.1 1 University of Queensland, St. Lucia, QLD, Australia; 2 University of Nebraska-Lincoln, Lincoln, NE. Can
harvest refuges counteract the effect of evolutionary change in body
size caused by size selective harvesting?
In wildlife harvesting the selection criteria for targeted animals is often a
large body size. There is increasing evidence that such size selective harvesting can lead to a directional change in body size in heavily harvested
ecosystems. We constructed a stochastic simulation model to explore the
possible evolutionary responses to size selective harvesting, and the effects
of different management strategies. In particular we examined the effects
of harvest refuges and minimum size regulations in kangaroo populations
in eastern Australia. Our modeling results suggest that when an entire population is subject to size-selective harvesting this is likely to result in smaller kangaroos of a given age. The effect of size-selective harvesting increases with increasing harvesting rate and increased minimum size of the harvested animals. In scenarios with moderate dispersal from an un-harvested
population all effects of harvesting on genetic structure are negligible.
TESKEY, ROBERT O.* and MARY ANNE MCGUIRE. University of
Georgia, Athens, GA. A mass balance approach for calculating stem
respiration in trees.
The rate of respiration of a tree stem has commonly been estimated from
measurements of CO2 efflux to the atmosphere. These estimates are based
on the assumption that all efflux of CO2 originates from respiration of local
tissues and that all CO2 produced by local tissues escapes to the atmosphere
through the bark. However, dissolved CO2 can be transported in the xylem
and the CO2 concentration in the stem can be up to three orders of magnitude greater than that of the atmosphere, suggesting that measurements
of CO2 efflux to the atmosphere do not account for all CO2 produced by
respiration. We developed a new mass balance approach for estimating the
respiration rate of tree stems that accounts for both external and internal
fluxes of CO2. We demonstrate this approach in three trees of different
species using measurements of CO2 efflux, sap flux, and internal CO2 concentration to calculate the total rate of CO2 production of a segment of
stem tissue in situ. When we applied the mass balance approach, we found
that CO2 produced by respiration of stem tissues moves in internal and
external flux pathways in different proportions at different times of day
and in different environmental conditions. Our calculations show that a
substantial portion of respired CO2 is not accounted for in measurements
of efflux to the atmosphere alone. In some cases, more than 75% of the
respired CO2 moved in the internal pathway, with less than 25% released
to the atmosphere. During daylight hours when sap is flowing, a large
proportion of respired CO2 is carried away in the xylem stream, while at
night more respiratory CO2 escapes to the atmosphere through bark. Measurements made using the mass balance approach can lead to an improved
understanding of woody tissue respiration rates.

THACKER, ROBERT W.,1,* PATRICK M. ERWIN,1 KEVIN P. BEVIS1

and VALERIE J. PAUL.2 1 University of Alabama at Birmingham, Birmingham, AL; 2 Smithsonian Marine Station, Fort Pierce, FL. Manipulations of sponge-cyanobacterial symbioses: Effects of shading on sponges with specialist and generalist symbionts.
The marine sponge Dysidea herbacea hosts large populations of the hostspecific, symbiotic, filamentous cyanobacterium Oscillatoria spongeliae.
Cyanobacteria may benefit their host sponges by fixing carbon or nitrogen
that is subsequently acquired by the host sponge. To determine whether
cyanobacteria significantly contribute to sponge metabolism through photosynthesis, we manipulated light availability in the field. Sponges were
held in plastic cages to remove the effects of fish predation. Light cages
were constructed with transparent tops, while shaded cages had opaque
tops. Sponges were distributed among the treatments in a randomized block
design, blocked by source sponge colony. After 16 days, we terminated the
experiment due to the dramatic mass loss of sponges in the shaded treatment. Sponges in the light cages lost 3.8% of their wet mass, an amount
that was not significantly different from zero loss. Sponges in the shaded
cages lost 28.5% of their wet mass, a significantly larger loss than the light
cages (ANOVA, P 5 0.002), despite a significant amount of variation
among sponge colonies (P 5 0.01). These results suggest that D. herbacea
obtains more nutrition from light-harvesting symbionts than by feeding
from the water column. Shading of sponges containing generalist symbionts
(e.g. Xestospongia spp. that host Synechococcus spp.) does not have a
measurable impact on sponge growth. The degree of physiological dependence on cyanobacterial photosynthesis may influence the degree of evolutionary specialization observed between sponges and their cyanobacterial
symbionts.
THOMAS, CAROLYN L.* and BOB R. POHLAD.* Ferrum College, Ferrum, Virginia, USA. The use of small watersheds as teaching tools.
Small watersheds make valuable tools for teaching in many different ecologically related courses. These watersheds can and should be local to your
campus and give valuable hands on experiences to your students. Some of
the upper level courses in which the Ferrum College, Ferrum Mountain
Creek Watershed, has been used are Ecology, Hydrology, Wildlife and Forest Management, Field Botany, Invertebrate Zoology and Environmental
Chemistry. The watershed has also been used for freshman and sophomore
level courses such as General Biology, Plant Science and Environmental
Issues. A methods manual has been developed for methods easily used in
small watersheds in the mountains with detailed procedures and descriptions of simple equipment that can be purchased or constructed for use by
students. This manual is online at a website developed for the Collaboration
through Appalachian Watershed Studies (CAWS) project. Methods will be
described including: 1) Leaf pack preparation and placement for studying
leaf decomposition in streams, 2) Aquatic fungi population analysis, 3)
Manual stream flow, 4) Stream chemical and physical parameters, and 5)
Meiofauna population analysis. These methods were developed by a group
of faculty at colleges in the Appalachian Mountains through two collaborative grant projects, Research Opportunities for Collaboration in the Appalachians (ROCA) and CAWS. Southern Appalachian watersheds at seven
different small colleges were instrumented, data collected, and methods
developed for the last two years. Weather and hydrologic data were collected and Geographic Information Systems (GIS) mapping was done on
these seven small watersheds with the help of undergraduate students. Data
has been shared through a project web site (www.ferrum.edu/caws) for
student study and learning purposes and training has occurred among the
seven colleges participating at frequent workshops attended by faculty and
students.
THOMAS, KATHRYN A.1,* and JOEL MICHAELSEN.2 1 US Geological
Survey, Southwest Biological Science Center, Flagstaff, Arizona; 2 Department of Geography, UC Santa Barbara, Santa Barbara, California. Response of plant species to climate change across a Southwest elevation
gradient.
The potential effects on plant species distribution in the arid Southwest of
predicted global climate change due to increase in atmospheric carbon dioxide were examined. Distribution data for dominant plant species in var-

Abstracts

333

ious vegetation types were collected and compiled across an elevation ecotone from the eastern Mojave Desert through the southwest Colorado Plateau. Three maps of precipitation averages during the last century and two
maps of mean maximum and minimum temperatures for the same time
period were developed. Potential future climate conditions were modeled
by developing maps of precipitation for the same monthly segments for
the three warmest years of the 1990s. These years were taken to represent
potential climatic conditions that might exist with 2x increase in CO2. The
current distribution of the plant species within the study area was spatially
modeled using decision tree analysis with the dependent variables of elevation, temperature, precipitation and geology, the current distribution surface. Plant habitat requirements were identified from the environmental
relationships expressed in the current distribution surface. Future plant habitat surfaces were modeled using elevation, geology and the warm year
temperature and precipitation maps. Potential future distribution surfaces
were developed by assigning species occurrence to the future plant habitat
surface using defined plant habitat requirements. The distance between current distribution and potential future distribution represents the migration
distance required for successful occupation of the new habitat locations.
Vulnerable landscapes in the elevation gradient are proposed based on identifying areas with high predicted loss of species due to loss of suitable
habitat for that species and low predicted colonization of species suitable
to the new habitat conditions at that location.
THOMAS, MICHAEL A.,1,* MARC PERRONE,2 BIXIA XIANG,2 DARREL MAZZARI,2 PEIGANG LI2 and MICHAEL J. CARVAN.3 1 Department of Biological Sciences, Idaho State University, Pocatello, ID, USA;
2
Marquette University, Milwaukee, WI, USA; 3 Great Lakes WATER Institute, Milwaukee, WI. Microarray experiments for non-model organisms: Estimates of error rates for cross-species hybridizations.
Microarrays are now routinely used in biomedical research as a tool for
gene discovery and expression profiling. This tool has yet to be extensively
used by researchers in evolutionary biology and ecology due to cost constraints and lack of species-specific arrays, despite its enormous potential.
While the cost is becoming increasingly attractive, the lack of well characterized genomes make the second problem difficult to overcome. Few
researchers are willing and capable of generating a cDNA library for their
organism of interest, and (when constructed) these libraries are often poorly
annotated, limiting their utility. One solution is the use of cross-species
hybridizations - using a microarray slide from a sequenced and annotated
model organism to explore gene expression in the organism of ecological
interest. For example, a microarray slide from Rattus norvegicus, the lab
rat, might be used to explore gene expression and behavior in Neotoma
floridana, the eastern wood rat, or a Danio rerio (zebrafish) microarray
slide might be used to examine gene expression and evolution among members of the Cichlidae of Lake Malawi. Understanding the expected error
rate of such experiments, and the affect of evolutionary divergence on this
error rate, is essential to an appropriate and meaningful interpretation.
Without such understanding, researchers will be easily misled, wasting
valuable resources. We conducted a computer simulation of cross-species
microarray hybridization that allowed us to predict error rates (hybridization of non-homologues) that can be expected for these experiments. The
results of this analysis allow us to make general recommendations for potential experiments, given a divergence time between the target species (of
ecological interest) and probe species (from which the microarray slide is
made).
THOMAS, SERGE,* EVELYN E. GAISER, MIROSLAV GANTAR,
RONALD D. JONES and LEONARD J. SCINTO. SERC/FIU, OE 148 UP,
33176, Miami, FL, USA. Dry benthic periphyton recovery and phosphorus buffering upon wetting in the Everglades: A microcosm survey.
Calcareous periphyton mats attached to limerock in the southern Everglades
(Florida, USA) were collected in June 2001 after 6 months of desiccation
and before the annual wet season inundation. Primary production, phosphorus kinetics and microbial community composition were followed for
20 days after flooding three microcosms (51x37x14 cm3) with 20 L of
artificial marsh water (AMW) in a greenhouse. The dry mat was dominated
volumetrically and numerically by cyanobacteria (90-99%) and this dominance remained unchanged after wetting, confirming that cyanobacteria

334

Abstracts

can both withstand 6 months of desiccation and recover faster than the
other taxa upon rewetting. Cyanobacteria recovery was rapid enough to
exhibit a positive net primary production just hours after wetting. The
limited development of the other taxa (green algae and diatoms) showed
the poor capabilities of these taxa to withstand 6-months of drying and to
recover fast upon rewetting. The persistence of low abundances of such
taxa during the dry season can be attributed to the moderate dry conditions
of the Floridian dry season and the moisture-retaining capability of the
mat. Light-corrected primary production was positively correlated with water total phosphorus concentration (TP, r250.6, P,0.001), which peaked 3
days after wetting (16 ug L-1) but leveled fairly rapidly after 5 days (6.6
ug L-1). This showed that TP limited the photosynthetic activity. Moreover,
the comparison between a simulated TP desorption to the water from the
C-111 mat (5 abiotic TP release) and the real TP concentration in the
water column exhibited that i) the mat microbial community recovers fast
enough to absorb TP initially released to the water column, thus preventing
high concentrations in water TP and ii) the mat does not absorb water TP
below concentrations of 6.6 ug L-1.
THOMPSON, ANDREW R.* University of California, Santa Barbara, Santa Barbara, CA. Influence of mutualism, competition, predation and
habitat on shrimp goby population dynamics.
Population dynamics of coral reef fish are influenced by a variety of biotic
and abiotic factors. To determine how mutualism, predation, competition
and habitat affect the population dynamics of the fierce shrimp goby (Ctenogobiops feroculus), I conducted field and laboratory experiments in Moorea, French Polynesia. Shrimp gobies interact mutualisticaly with snapping
shrimp (Alpheus sp.) as follows: shrimp construct burrows in which both
gobies and shrimp reside, and gobies warn shrimp of predators through
touch-based signals. Surveys of 686 3 x 3m quadrats throughout the north
shore of Moorea indicated that goby distributions were constrained by the
presence of shrimp, whereas shrimp were confined to habitats with a mixture of rubble and sand. Even within appropriate habitats, however, goby
densities were variable. Subsequent surveys demonstrated that recruitment
explained 88% of the variability in adult goby density among locations
with appropriate habitat. In locations with high recruitment, gobies competed intraspecifically for large shrimp, and large gobies were competitively
dominant. A strong, negative correlation between predator and large goby
densities indicated that predation also affected goby population structure.
In addition, a positive correlation between the occurrence of small gobies
residing with large shrimp and predators further suggested that predators
impacted goby populations. A field experiment demonstrated that predators
readily consumed large gobies, and that small individuals paired up with
large shrimp when a shrimp became available due to the death of a large
goby. These results indicate that shrimp availability, which is constrained
by appropriate habitat, affects goby presence/absence; variable recruitment
influences patterns of goby abundance; and predators influence goby sizestructure and turnover rates.
THOMSEN, MEREDITH A.* and BLAKE SUTTLE. University of California, Berkeley, Berkeley, CA. Importance of propagule supply and
ecological resistance in determining invasion success.
The observation that some plant communities contain more non-native species than others has important implications for community ecological theory and for management decision-making. The degree to which a site is
invaded is determined by two factors: the rate of exotic species arrival
(propagule supply) and the biotic and abiotic site characteristics that determine invader success (ecological resistance). Although a number of studies have experimentally manipulated elements of resistance and documented their effects on the success of invading species, relatively few have
manipulated propagule supply or tested for interactions between the two.
We evaluated the importance of propagule supply and resistance in determining invasion success of the exotic perennial grass Holcus lanatus in
California coastal prairie. For two years we decreased competition from
resident plants by clipping background vegetation and increased water
availability while adding Holcus seeds at four densities. Reduced competition positively affected Holcus seedling establishment in 2002 but not in
2001 while watering caused an increase in total Holcus seedling numbers
in both years. Nonetheless, even in watered plots, most seedlings died

before the end of the summer drought typical of this region, apparently of
water stress. The relationship between propagule supply rate and seedling
numbers was positive, with decreasing slope at higher values, suggesting
a threshold value of seed rain above which seedling numbers saturate.
There was no significant interaction between propagule supply rate and
manipulation of competition or abiotic stress (watering). It seems likely,
however, that the abiotic resistance of low water availability in this site is
too strong to be overcome by any level of propagule supply. In other
systems, propagule supply and resistance may be more likely to interact.
THORPE, ANDREA S.* and RAGAN M. CALLAWAY. University of
Montana, Missoula, Montana, USA. Interactions between the invasive
forb, Centaurea maculosa, and soil nitrogen cycling.
Positive feedback with soil microbes appears to play an important role in
the success of invasive plants. This feedback effect has been attributed to
invasive plants escaping species-specific soil pathogens and taking advantage of generalist mutualists such as AM fungi. Our research suggests that
feedback between the invasive forb, Centaurea maculosa Lam. (Asteraceae) and soil microbes may also involve the soil nitrogen cycle. Microbial
communities in soil collected from C. maculosa rhizospheres were significantly different than those collected from native grass rhizospheres and
there is evidence that such microbial effects may drive positive feedback
between C. maculosa and the soil community in Montana. Furthermore,
soil ammonium and nitrate are reduced in C. maculosa -invaded communities compared to adjacent, uninvaded communities. Nitrate was approximately 75% higher in native communities than invaded communities (P
, 0.0005). These patterns may be due simply to different uptake rates;
however, growing evidence suggests that an anti-microbial root exudate
produced by C. maculosa, (+)-catechin alters the activity of nitrifying bacteria and changes the nitrogen cycle in ways that benefit C. maculosa. Our
results indicate that the invasive success of C. maculosa may be due in
part to complex interactions with microbial components of the soil ecosystem.
THREAT, CALVIN J. and GRETCHEN B. NORTH.* Occidental College,
Los Angeles, CA, USA. Seedlings of desert agave grow better in sandy
than in rocky soil.
In the northwestern Sonoran Desert, Agave deserti occurs primarily in
rocky microhabitats and only infrequently in sandy soil. A previous study
indicated that the shoot growth and root production of mature agaves were
greater, however, in sandy soil, suggesting that seedling growth and requirements might account for greater plant frequency in rocky soil. In one
experiment, root and shoot growth were measured for seedlings in containers of field-collected sandy or rocky soil during 30 d of watering followed by 30 days without water. Seminal root length was nearly two times
greater in sandy than in rocky soil at the end of 60 days, and only seedlings
in sandy soil produced adventitious roots. Root dry weight was two times
greater for seedlings in sandy soil, although shoot dry weights were similar.
Shoot growth over the 60 days was higher for seedlings in sandy than in
rocky soil, although the pattern was reversed in the last week of the experiment. Therefore, we are currently investigating the effects of more prolonged soil drying. In a second experiment, the effect on seedling growth
of the shade provided by rocks was examined. Three sets of containers
filled with sandy soil were planted with seedlings 1) adjacent to rocks, 2)
adjacent to plastic screens of similar projected areas as rocks, or 3) unshaded. Containers were placed in full sun outdoors in late summer and
fall for 45 days without watering, with rocks and screens on the south side
of seedlings. Despite record-breaking heat in Los Angeles, seedling growth
was not improved by shade. We are now investigating whether rocks in
the soil improve seed germination and the anchorage of newly germinated
seeds.
THUM, RYAN A.* and RICHARD S. STEMBERGER. Dartmouth College, Hanover, NH, USA. Separating historical and contemporary factors that determine distribution and co-occurrence patterns in freshwater copepods.
Patterns of species occurrence are influenced by historical (e.g., phylogenetic and biogeographic) and proximal (e.g., species interactions, habitat

selection) factors. For example, two closely related species may not cooccur at local sites because of historical/contemporary dispersal barriers or
because they mutually exclude one another via reciprocal competitive interactions. Finally, the degree to which the two species are reproductively
isolated may also influence their patterns of co-occurrence via Allee effects
and/or incompatibilities of hybrid genotypes. Separating historical and contemporary factors is particularly difficult when direct estimation of dispersal capacity is impossible. If dispersal is not limiting to species9 distributions, mutually exclusive distribution patterns should result from fitness
differences at local sites. However, if dispersal is limiting, mutually exclusive distributions can be maintained in the absence of ecological and/or
reproductive differentiation. I addressed these issues to consider the potential mechanisms maintaining the parapatric distributions of sister species
of Skistodiaptomus (Copepoda: Calanoida). Mating experiments showed
these species to be almost completely reproductively isolated despite a very
high degree of morphological similarity. Discriminant function analysis had
great difficulty separating these species according to the range of habitat
variables among northeastern US lakes (e.g., water chemistry, physical
characteristics of lakes, and surrogate variables for potential competitors
and predators). Moreover, the functions discriminating lakes inhabited by
each species may be confounded by large-scale landscape patterns. These
results, in combination, suggest that allopatric speciation and range expansion of these species to distributional limits set by dispersal barriers, and
not competition or ecological differentiation, determine the distribution patterns of these closely related diaptomid copepods.
THYGERSON, TONYA,1,* BRUCE N. SMITH,1 LEE D. HANSEN,2
RICHARD S. CRIDDLE2 and V. W. MCCARLIE.2 1 BYU Dept Plant and
Animal Science, Provo, UT; 2 BYU Dept Chemistry and Biochemistry, Provo, UT. Temperature and temperature variability in determining gradients in plant species range.
There are many factors that determine where a plant (an ectotherm) will
grow. One of the primary factors is temperature. This paper develops the
hypothesis that temperature (T) and temperature variability (DT) are the
primary climatic variables determining global-scale gradients in species
range. The physiological factor defining adaptation of ectotherms to temperature is respiratory energy metabolism. Ectotherms adapt to the latitudinal/longitudinal gradients of T and DT which correspond to their gradients of adapted energy metabolism. Natural selection results in ectotherms
with metabolic properties corresponding with their environment. Measurements of the rates of respiration and energy production as a function of
temperature show that respiratory rates and efficiency of ectotherms are
closely adapted to their native environment. Principles developed in this
paper yield fundamental laws of ecology that allow the calculation of the
contributions of global temperature patterns to the formation of the global
gradient of species range. Relative values of range can therefore be calculated from data on abiotic variables. Predictions agree with known patterns of plant distribution.
TIAN, HANQIN,1,3,* JERRY M. MELILLO,2 JIYUAN LIU,3 DAVID
KICKLIGHTER,2 SHUFEN PAN,1,3 FANGFANG YU1,3 and MINGLIANG
LIU.1,3 1 The University of Kansas, Lawrence, KS 66045; 2 The Ecosystem
Center, Marine Biological Laboratory, Woods Hole, MA 02543; 3 Institute
of Geographic Science and Natural Resource Research, Chinese Academy
of Science, Beijing, 100101, China. The carbon cycle of terrestrial ecosystems in tropical and temperate Asia.
Newly developed data on three major determinants of the carbon storage
in terrestrial ecosystems are used with the process-based Terrestrial Ecosystem Model (TEM) to simulate the combined effect of climate variability,
increasing atmospheric CO2 concentration, and cropland establishment and
abandonment on the exchange of CO2 between the atmosphere and terrestrial ecosystems in tropical and temperate Asia. During 1860-1990, modeled results suggest that tropical and temperate Asia as a whole released
29.0 Pg C, which represents 50% of the global carbon release for this
period. For the recent decades, tropical Asia acted as net carbon source,
but temperate Asia acted as a net carbon sink. Substantial interannual and
decadal variations occur in the annual net carbon storage estimated by TEM
due to comparable variations in summer precipitation and its effect on net
primary production. At longer time scales, land-use change appears to be
the important control on carbon dynamics in this region.

Abstracts

335

TISSUE, DAVID,1,* MATTHEW TURNBULL,2 KEVIN GRIFFIN,3

MARGARET BARBOUR,4 JOHN HUNT,4 WILLIAM SCHUSTER,5 KIM
BROWN6 and DAVID WHITEHEAD.4 1 Texas Tech University, Lubbock,
TX; 2 University of Canterbury, Christchurch, New Zealand; 3 Columbia
University, Palisades, NY; 4 Landcare Research, Lincoln, New Zealand;
5
Black Rock Forest Consortium, Cornwall, NY; 6 Ohio University, Athens,
OH. Scaling photosynthesis from shoots to the canopy in a rimu dominated rainforest in New Zealand.
Our objective was to determine the potential for estimating CO2 exchange
in a complex, mixed species, natural rainforest in New Zealand using photosynthetic and internal CO2 (Ci) measurements taken at different scales.
Our study site is representative of an extensive multi-layered, mixed conifer-broadleaved forest ecosystem in South Westland, that is dominated
by rimu (Dacrydium cupressinum). The forest receives high rainfall (3.2
m), has highly leached and nutrient poor soils, and exhibits low productivity. Carbon exchange in this forest is sensitive to small shifts in climate,
including temperature, cloudiness and periods of seasonal water deficit. We
used direct measurements of leaf photosynthesis (ACicurves) to estimate
potential limiting factors to photosynthesis (e.g. Vcmax and Jmax). We used
the Leuning coupled photosynthesis-stomatal conductance model to estimate a shorter-term value of Ci at the leaf level, which was tested against
direct longer-term estimates of Ci based on delta 13C isotope discrimination
in leaves measured at different heights through the canopy. These results
were compared with longer-term whole ecosystem Ci values estimated using Keeling plots (i.e. measurements of delta 13C discrimination in air samples taken at different heights through the canopy). Decreasing values of
Ci with increasing height in the canopy indicate higher rates of photosynthesis (confirmed by gas exchange measurements) at higher light levels.
Although photosynthesis is limited by light availability, the primary limitation to photosynthesis in these forest trees appears to be low nutrient
availability that greatly reduced biochemical capacity (i.e. Vcmax and Jmax).
In general, there was good agreement between the three estimates of Ci
suggesting that scaling CO2 exchange from the shoot to the canopy level
is possible in this forest using a multi-layered model based on shoot-level
photosynthesis.

lacking rhizomes that reproduces apomictically; and var. curtipendula,

which reproduces sexually and from spreading rhizomes. At the regional
scale, var. curtipendula dominates the relatively mesic (eastern) portions
of the Great Plains, with var. caespitosa occupying the more xeric (western)
portions, extending into southwestern desert grasslands. We hypothesized
that if the differences in these varieties reflects natural selection for aridity
and/or disturbance, then within their zone of sympatry, the two growth
forms would be locally segregated in accordance with variables indicative
of landscape moisture/temperature, fire history or grazing gradients. To test
this hypothesis, we inventoried 321 - 0.5 x 1 m plots in pastures with
contrasting grazing/prescribed fire histories in an Edwards Plateau JuniperOak savanna site in central Texas. For each plot we recorded the % cover
of each variety and 11 explanatory variables including disturbance history
(fire and grazing), abiotic setting (topographic position, limestone type, soil
depth, % bare ground, rock size and rock cover) and proximity to the
nearest tree or shrub (source of radiant energy amelioration). Data were
analyzed using canonical correspondence analysis (CCA) with forward selection of explanatory variables. Significance of variables was tested using
Monte Carlo permutations. Local segregation of B. curtipendula varieties
was evident, in that only 1.9% of the plots sampled contained both varieties. The rhizomatous variety was associated with Buda limestone, small
rocks, no grazing and winter fire, whereas the caespitose growth form was
related to Edwards limestone sites with large rocks. However, environmental and disturbance variables we measured accounted for only 18.6 % of
the variation, suggesting factors other than those measured are affecting
local distributions of the two varieties.
TOWNSEND, ALAN R.,* CORY C. CLEVELAND and CARLETON R.
BERN. University of Colorado, Boulder, Boulder, CO. Biotic vs. geologic
controls on soil and foliar nutrient pools in a lowland neotropical rainforest.

Fire is an important disturbance in Neotropical rainforests. Here we report

a record of ancient fires along an elevational transect that extends through
old growth forest from the Continental Divide to the Caribbean lowlands.
Soil charcoal amount, depth distribution and age are measured from sixteen
1.5 m deep soil cores, in each 1 ha permanent plot at elevations of 50 m,
300 m, 500 m, 750 m, 1000 m, 1250 m, 1500 m, 1750 m, 2000 m, 2300
m and 2600 m along the Volcan Barva transect in Braulio Carrillo National
Park, Costa Rica. The transect crosses four life zones (lowland tropical,
premontane, lower montane and montane tropical forests) and extends from
50 m to 2906 m. Charcoal was found at every elevation ranging from as
much as 1027 g/m2 at 300 m to as little as 20 g/m2 at 1750 m. Charcoal
samples are radiocarbon dated and the dates range from 23,240 yr B.P. at
110-120 cm soil depth at 1750 m elevation to 140 yr B.P. at 30-40 cm soil
depth at 2600 m elevation. Many dates correspond with two intervals of
previously studied prehistoric burns of 2430 yr B.P. and 1110 yr B.P. at
the lower elevations of the transect. Interestingly, none of the high elevation
forest charcoal samples (.2300 m) are older than 170 yr indicating that
the forests near the Continental Divide may be relatively young, replacement stands that have come in since the last volcanic eruption. Soil charcoal
is a signature of historical land use and is also an indicator of the Holocene
fire history. These forests have regenerated multiple times as a consequence
of anthropogenic or naturally induced fires and disturbances.

Many of the worlds lowland tropical forests exist on old, highly weathered
soils that are typically poor in phosphorus (P) and the base cations calcium
(Ca), magnesium (Mg) and potassium (K). Prior evidence from other studies suggests that both soil and foliar pools of such rock-derived elements
should be substantially lower on these old soils than one might find in
much younger soils where substantial primary mineral sources of nutrients
remain. In addition, some evidence indicates that ecosystems on old soils
may rely almost entirely on atmospheric sources of P and base cations. We
established forest sites in southwest Costa Rica that share a similar climate
regime and several common tree species, but differ substantially in soil
type, ranging from forests on rich, Quaternary era alluvial Mollisols to
highly weathered Oxisols developed from basaltic material in excess of 50
million years of age. We then measured concentrations of multiple nutrients
in canopy leaves from several tree species across these sites, as well as
total and available soil nutrient pools in each site. We also used strontium
(Sr) 87/86 isotopic ratios to distinguish atmospheric versus local weathering
sources of Sr in soil and foliage. The results show that total and available
pools of soil P are substantially higher on the younger Mollisols than on
the older Oxisols, but that differences in foliar P, while present, are not
nearly as great as those in the soil pools. In addition, within a given soil
type, both between-species and seasonal differences in P and several other
elements often match or surpass the cross-site differences for any single
species. These results suggest that the diverse biota present in mainland
tropical forests may create small-scale variation in nutrient cycling patterns
that rival or exceed those seen across larger geologic gradients, with potential implications for both long-term ecosystem development and tropical
forest responses to environmental change. Finally, mass balance analyses
of the soil using immobile trace elements show nearly complete losses of
parent material base cations, yet the Sr isotopic data reveal a dominance
of parent material sources in soil exchangeable and foliar pools. Groundwater transport of local, basalt-derived Sr may account for this apparent
paradox.

TOMAS, ANDREA,1,* FRAN GELWICK1 and STEVE ARCHER.2 1 Texas

A & M University, College Station, TX; 2 University of Arizona, Tucson,
AZ. Local distributions of Bouteloua curtipendula varieties within their
zone of sympatry.

TOWNSEND, PHILIP A.,1,* ROGER W. BROWN1 and DEBRA A. WILLARD.2 1 University of Maryland Center for Environmental Science, Appalachian Laboratory, Frostburg, Maryland; 2 U.S. Geological Survey, Reston, Virginia. Long-term geomorphic and vegetation change on the Roanoke River floodplain, North Carolina.

Bouteloua curtipendula, an important component of native prairie ecosystems, consists of two varieties: var. caespitosa, a bunchgrass growth-form

We established a series of 50 transects across a 200-km extent of the lower

Roanoke River floodplain in North Carolina. Along these transects, we

TITIZ, BEYHAN* and ROBERT L. SANFORD, JR.* University of Denver, Denver, CO. Soil charcoal distributions along an elevational gradient in a Costa Rican rainforest.

336

Abstracts

collected sediment cores at regular intervals to conduct paleoecological

reconstructions of sedimentation history and vegetation change. Analyses
of ragweed pollen concentration from the cores indicate up to 5 m of
sediment deposition on the floodplain during the 300 years since European
settlement. Moreover, deep cores from the early Holocene suggest that total
amounts of sedimentation during recent centuries exceed total accumulations from the previous 10,000 years. We compared sediment characteristics and pollen assemblages from the pre-colonial landscape to surface pollen samples and a recent vegetation survey along the established transects.
We found that floodplain soils underwent a transition from a largely organic
substrate prior to European settlement to mineral soils during the period of
rapid sedimentation, and that the soils are showing signs of increasing
organic matter content in recent decades as the river has become sedimentstarved due to dam construction. Paleoecological evidence suggests the
development of less flood tolerant forest assemblages across large areas of
the floodplain that experienced deep sedimentation. Deepwater swamp habitats remain dominated by species of Taxodium and Nyssa, although selective logging of bald cypress in recent centuries has led to switch in dominance from Taxodium distichum to Nyssa aquatica. In addition, a rich fern
flora that was previously associated with Taxodium-Nyssa swamps appears
to have disappeared from the modern landscape, possibly due changes in
soil characteristics and flooding regimes. In a long-term context, the sedimentation that resulted from the clearing of uplands during the period of
European settlement appears to have led to substantial changes in floodplain forests.
TRAUT, BIBIT H. University of California, Davis, CA. The effects of
nitrogen addition on the structure and dynamics of salt marsh vegetation.
Nutrient limitation of component wetland species may differ from that of
the whole community and be critical in determining the effect of nutrient
addition on plant diversity. In salt marshes of Point Reyes, California,
where land-use practices (e.g. grazing) increase the input of nitrogen (either
directly or indirectly as runoff from within the salt marsh watershed), marsh
vegetation is dominated by Distichlis spicata and is less diverse than
marshes without excess nutrients. Nutrient addition experiments are helpful
in determining the response of natural vegetation to increased nitrogen
deposition. Using a field experiment, I investigated the role of soil fertility
on the high salt marsh plant community. I hypothesized that when soil
fertility is increased by nitrogen addition (1) plant productivity will increase, as represented by height, biomass, and cover, and (2) competitive
exclusion, presumably by Distichlis, will lead to a reduction in species
richness and evenness, especially where Distichlis is initially present in
greater amounts (from transplanting). After two growing seasons, biweekly
nitrogen addition to the high salt marsh lead to increased plant productivity.
However, diversity was not reduced, and space pre-emption (e.g. Distichlistransplants) did not confer a competitive advantage. Although the competitively superior species thrived (e.g. Salicornia virginica, Distichlis spicata,
Triglochin concinna), they did not displace subdominant species and decrease diversity. The high salt marsh system in this study does not support
the hypothesis that as productivity increases in response to increased nitrogen, competitive exclusion will occur and diversity will decrease. In the
inherently stressful environment of a salt marsh, a competitive superior
will have trade-offs with stress tolerance. The addition of nitrogen in this
study may affect not only the competitive ability of a plant but its salt
tolerance as well. High salt marsh plants may incorporate a suite of strategies to deal with stress and resource competition.
TRESEDER, KATHLEEN K.,1,* MARGARET S. TORN2 and CAROLINE
A. MASIELLO.3 1 University of Pennsylvania, Philadelphia, PA; 2 Lawrence Berkeley National Laboratory, Berkeley, CA; 3 University of California at Santa Barbara, Santa Barbara, CA. Do ectomycorrhizal fungi use
soil carbon? A field test with large-scale radiocarbon enrichment.
Ectomycorrhizal fungi acquire mineral nutrients by secreting exoenzymes
into the soil; these break down organic compounds for subsequent absorption by hyphae. However, the extent to which the fungi can augment carbon
acquisition through this process is unknown. We used a large-scale radiocarbon labeling experiment in Tennessee to directly test for this possibility.
In the summer of 1999, a pulse of 14CO2 was emitted to the atmosphere of

the Oak Ridge Reservation, and plants incorporated this label into their
tissues. In the fall of 2000, leaf litter was collected from enriched and
relatively unexposed areas, and a reciprocal transplant was established as
part of the Enriched Background Isotope Study (EBIS). Ectomycorrhizal
tips were collected from each area one year later. We found no evidence
of soil carbon uptake by the fungi. Only ectomycorrhizal fungi growing in
enriched areas were themselves enriched (D14C: 305 612). Where trees
had not been exposed to 14C, ectomycorrhizal fungi were not enriched even
when growing in plots with enriched litter (160 610, P , 0.001). Ectomycorrhizal fungi in this system had not switched between mutualistic
and saprotrophic functions at any point over the course of one year. These
results have important implications for models of nutrient cycling in ecosystems where ectomycorrhizal fungi dominate.
TREXLER, JOEL C.,1 JEROME J. LORENZ2 and WILLIAM F. LOFTUS.3
1
Florida International University, Miami, FL; 2 National Audubon Society,
Tavernier, FL; 3 U.S. Geological Survey, Homestead, FL. Monitoring fish
invasions and their impacts in southern Florida: Can we predict the
risks?
In a recent paper, we noted that the abundance of non-native fishes was
generally low in freshwater marshes of the Everglades at sites distant from
canals, but was relatively high in some habitats bordering that ecosystem.
Further, we reported a negative correlation over a 10-year period between
the numbers and biomass of native fish and the same parameter for the
non-native Mayan cichlid, Cichlasoma uropthalmus, in mangrove habitats.
Since that publication, we have observed an expansion of Mayan cichlids
into Everglades wetlands adjacent to the mangrove zone. These data permit
us to estimate a rate of spread of this species. At present, we can identify
no statistical evidence for effects from any non-native taxa on native fish
communities in freshwater wet-prairie habitats. However, in the mangrove
zone on the northern fringe of Florida Bay, the native species sheepshead
minnow (Cyprinodon variegatus) and marsh killifish (Fundulus confluentus) decline in abundance in periods when Mayan cichlids are common,
and increase following cold weather that decreases cichlid abundance. Two
other small, abundant native fishes, sailfin molly (Poecilia latipinna) and
rainwater killifish (Lucania parva), appear to be unaffected by the changing
abundance of Mayan cichlids. Our work indicates that Mayan cichlids have
altered the community structure and dynamics of native fishes in some
habitats in southern Florida, but not in others. We use our data to develop
a predictive model of the spread of Mayan cichlids into wetlands of the
Everglades, as well as their impact on freshwater fish communities there.
Further, we discuss the ability to generalize our observations to other expansions and evaluate if predictions can be made early enough to intervene
once a new fish species is detected.
TRIPATHI, GHANSHYAM,* SETHA RAM and BRIJ MOHAN SHARMA. Department of Zoology JNV University, Jodhpur, Rajasthan, India.
Impact of drought on faunal biodiversity and functional aspects of soil
in arid environment.
Impact of drought on faunal biodiversity and functional aspects of soil has
been studied in some selected silvipasture systems of arid zone of Jodhpur
in India. The silvipasture systems were Prosopis cineraria, Acacia nilotica,
Zizyphus nummularia, Capparis decidua and Acacia Senegal based having
Cenchrus ciliaris, Cenchrus biflorus, Eleusine compressa or Digitaria marginatus grasses in different combinations. The period from November 2000
to October 2001 was considered non drought year and November 2001 to
October 2002 as drought year depending on average rainfall. The drought
caused a significant reduction in density of major groups of soil fauna.
Populations of Acari, Myriapoda, Isoptera, Coleoptera, Collembola and
other arthropods showed about 50% reduction in response to approximately
80% decreased rainfall in the drought year. The drought effect on soil fauna
during rainy and winter seasons was more or less similar in all silvipasture
systems. Though there were also some season specific minor differences
in population reduction in different silvipasture systems, but the range of
drought dependant decrease in faunal population never exceeded 33 to
63.95%. Maximum drought effect on Acari was found in P. cineraria field.
Likewise, maximum drought effect on Myriapoda, Isoptera and Coleoptera
was in A. senegal system. However, maximum effect of drought on Coleoptera and other arthropods was recorded in C. decidua field. In contrast

Abstracts

337

to maximum drought effect, the least effect of drought on Acari was in A.

nilotica, Myriapoda in Z. nummularia and other arthropods in A. Senegal
based silvipasture field. The population of Coleoptera, Isoptera and Collembola exhibited minimum drought effect in traditional desert pedoecosystem of P. cineraria. This reflects a kind of adaptive association between
desert plant and soil fauna. Some soil nutrients significantly increased during drought year. However, there was a substantial reduction in soil respiration and dehyrogenase activity. The present observation suggests that
drought decreases faunal diversity and density but this reduction appeared
to be silvipasture system specific.
TRUDEAU, VANCE,* CHRIS MARTYNIUK, NATACHA GALLANT
and MAXINE CROTEAU. University of Ottawa, Ottawa, Ontario, Canada.
Expression profiling endocrine disruption in ecologically relevant and
non-modelvertebrate species.
The concept of endocrine disruption is one where chemicals and pollutants
mimic or upset the normal hormonal systems in the body. Such a disruption
can have deleterious effects on growth, reproduction and other physiological processes. These effects can lead to developmental abnormalities, reduced fertility and can threaten the health and ultimately the survival of
wildlife populations. One challenge in endocrine disruption research is that
often the effects cannot be predicted based on chemical structures or overt
toxicity. Moreover, the organisms affected by these compounds in the wild
are rarely the species used for laboratory-based studies, nor is there genome
sequencing projects for these environmentally relevant species. We have
used a molecular biological approach called differential display analysis
(DDA). This method is used because it does not require extensive gene
sequence knowledge. The effects of an estrogenic pollutant octylphenol
(OP) on brain function in two indigenous North American species, the
snapping turtle (Chelydra serpentina serpentina) and the Leopard frog
(Rana pipiens) tadpole was assessed using DDA. We found that OP affects
the expression of genes in neurotransmitter systems, and growth and cell
differentiation factors. In the case of tadpoles, OP can also disrupt the
timing of metamorphosis. We are also developing a DNA microarray or
DNA chip approach to assess the multiplicity of effects of endocrine disrupting chemicals. Specifically, we want to determine sex differences and
the effects of estrogenic pollutants on brain function in the common goldfish (Carassius auratus), a representative cyprinid used in endocrine research. This presentation will introduce the theory behind using these genomic approaches and discuss specific results obtained in three vertebrate
species. Funded by NSERC, CNTC, OGS and Univ. Ottawa (Canada).
TRUEMAN, REBECCA J.,1,* RAMESH MURTHY,2 DAN YAKIR,3
JOHN D. MARSHALL,4 JOSEPH A. BERRY5 and MIQUEL A. GONZALEZ-MELER.1 1 University of Illinois at Chicago, Chicago, IL, USA;
2
Columbia University Biosphere 2 Center, Oracle, AZ, USA; 3 Weizmann
Institute of Science, Rehovot, Israel, Israel; 4 University of Idaho, Moscow,
Idaho, USA; 5 Carnegie Institute of Washington, Washington, DC, USA. A
novel methodology for estimating mean residence times of ecosystem
carbon pools.
During the last century, anthropogenic fossil fuel emissions have been
causing an increase in atmospheric carbon dioxide concentration that is
altering the integrity and functioning of ecosystems worldwide. Since carbon exchange by vegetation far outweighs the anthropogenic carbon inputs
any carbon cycling changes will determine whether an ecosystem is a
source or a sink for atmospheric carbon. Mechanisms that control carbon
storage and allocation in ecosystems have been elusive to scientists. Ecosystem carbon dynamics can be determined using natural abundance of
stable isotopes. Stable isotopes provide the opportunity to look at ecosystem carbon processes by analyzing fractionation of the least abundant stable
isotope 13C. Plant isotopic composition is more depleted in 13C than the
atmosphere and the level of depletion is affected by climatic conditions.
The isotopic signature of plant tissues is reflected in the respired carbon
dioxide of plants, soils, and ecosystems. The release of carbon by respiration will be from a combination of ecosystem carbon pools of different
mean residence time (MRT). In natural ecosystems it is difficult to not only
measure carbon fluxes but also to estimate the MRT of carbon pools. In
order to determine the MRT of carbon at an ecosystem scale a quantitative
measurement technique has been needed. This methodology will be presented with data from Columbia University Biosphere 2.

338

Abstracts

TRUMBORE, SUSAN,1,* JULIA B. GAUDINSKI,2 ERIC A. DAVIDSON,3 LUZ MARIA CISNEROS DOZAL,1 PLINIO CAMARGO,4 EDWARD AG. SCHUUR5 and JEFFREY Q. CHAMBERS.1 1 University of
California-Irvine, Irvine, CA; 2 University of California-Santa Cruz, Santa
Cruz, CA; 3 The Woods Hole Research Center, Woods Hole, MA; 4 Centro
de Energia Nuclear na Agricultura, Piracicaba, Sao Paulo, Brazil; 5 University of Florida-Gainseville, Gainseville, FL. Sources of soil respiration
determined using radiocarbon.
Soil respiration combines CO2 derived from autotrophic (root metabolism)
and heterotrophic (organic matter decomposition) sources. A major challenge in ecological research is to determine how much of soil respiration
comes from each of these sources, and how each may respond to environmental changes. We have developed methods using the difference in radiocarbon among these substrates to quantify these sources, and in addition
to determine the relative contribution of different decomposing substrates
to overall heterotrophic C losses. This talk will present radiocarbon measurements of soil respiration and its components from tropical, temperate
and boreal ecosystems, as well as discuss how temperature and moisture
can influence the different components of soil respiration. Radiocarbon in
microbially-respired CO2 provides a measure of the mean residence time
of fast-cycling C pools in ecosystems, which vary from several years in
tropical systems to decades in boreal forests. In contrast, soil organic matter
is often comprised of more stable, slowly cycling components that contribute little to soil respiration. The pools contributing the most to heterotrophic
respiration include leaf and root litter; in particular soil respiration rates
can be sensitive to the moisture content of the leaf litter layer. More stable,
humified, organic matter pools contribute little to respired C on an annual
basis, but are important for C balance on longer (decades-centuries) time
scales.
TRZCINSKI, KURTIS M.,1,* SANDRA J. WALDE1 and PHILLIP D.
TAYLOR.2 1 Dalhousie University, Halifax, Nova Scotia, Canada; 2 Atlantic
Cooperative Wildlife Ecology Research Network, Acadia Universtiy, Wolfville, Nova Scotia, Canada. Stability of pitcher-plant microfaunal populations depends on food web structure.
Ecological theory suggests that top-down and bottom-up effects can be
either stabilizing or destabilizing, depending on community structure. In
this study we examined how food web structure affected the dynamics of
three microfaunal taxa living in pitcher plants. We manipulated predator
and resource levels and censused population in 60 communities that were
open to colonization by microfauna and arthropods. Predation tended to
decrease microfaunal abundances and to be destabilizing, leading to higher
temporal variability and lower persistence times. Bottom-up effects varied
with taxon, and depended on the manner in which resource availability was
increased. Rotifer populations tended to be destabilized by the addition of
resources under low predation and stabilized under high predation levels.
Bottom-up effects were stabilizing for microflagellate populations, but this
consumer responded, not directly to the addition of resources, but to the
presence of midges whose activity increased bacterial abundances. We conclude that top-down effects tend to be destabilizing, but that the way bottom-up factors influence population dynamics within pitcher plant food
webs depends on community structure, which in turn, depends mostly on
patterns of colonization.
TUCKFIELD, R. CARY TUCKFIELD.* Department of Environmental
Science and Technology, Savannah River Technology Center, Bldg. 77342A, Aiken, SC, USA. Statistical thinking, practical uncertainty, and
ecological evidence.
What an ecologist really wants is to know is how likely the hypothesis is
to be true given the data she collected. Unfortunately, the traditional education we receive in classical frequentist statistics does not solve this problem directly. Instead, it addresses a different problem, viz., how likely is
it to get these data, given the hypothesis. The issue to think about is whether the latter is tantamount to the former. The ecologist is free to interpret
the results of such statistical tests, but he is not free to state them. To state
the results is to relate the outcome in the parameter-space of the presumed
and underlying probability distribution function. To interpret them requires
a translation back into science-space. We crave answers to intriguing ques-

tions - research after all is fun - but the space-translated counterparts offered, in the long run, as nearly identical substitutes are seldom as intriguing as the original. This state of affairs and our ever increasing reliance on
p-values for getting knowledge derives from the straitness of a single proscriptive paradigm; not parsimony, objectivity. The insistence on bias minimalized hypothesis tests has placed mathematics on the pedestal it currently occupies in the western way of research and the acquisition of
knowledge about nature. All natural processes however, occur within the
context of a system and cannot be measured without uncertainty. This is
the root statistical thinking. The only certainty is in the disturbing fact that
nature and what we think we know about nature will never be one and the
same. The new methods of statistics will rely on Bayesian Thinking. They
are those that will address the original science-space hypotheses directly,
without translation. They will admit to bias and a subjective foundation
because measurement is not without bias. Finally, these methods will rely
on evidence not confidence.
TURCHIN, PETER.* University of Connecticut, Storrs. Nonlinear dynamics in a noisy world, or why deterministic skeletons should be relegated to the closet.
Numerical dynamics of real-world populations are a result of a complex
interplay between endogenous (density-dependent) feedbacks and exogenous influences. A naive approach to analyzing complex population dynamics is to attempt to decompose them into deterministic and stochastic
components, sometimes referred to as the "deterministic skeleton" and
"noise". For example, if we want to determine whether a population fluctuates chaotically or not, within this paradigm we need to first extract the
deterministic skeleton, and then if we find that it is chaotic, then we classify
the studied system as chaotic, and vice versa. Unfortunately, when nonlinear endogenous feedbacks are combined with stochastic exogenous drivers,
the resulting dynamics may be more than simply the sum of two parts. I
illustrate this idea with a theoretical example in which a deterministically
stable dynamical system becomes chaotic when white noise is added to it.
Several real-life ecological systems also appear to possess this characteristic. My conclusion is that we cannot characterize dynamical systems in
nature by attempting to subtract noise from them.
TURNER, GREG,1,* JEFF LICITRA,2 J D. LEWIS1 and WILLIAM
SCHUSTER.3 1 Calder Center, Fordham University, Armonk, NY; 2 Blind
Brook High School, Rye Brook, NY; 3 Black Rock Forest Consortium,
Cornwall, NY. Forest composition affects ectomycorrhizal fungal community composition on Quercus rubra seedlings.
Host composition is an important factor that may influence ectomycorrhizal
(ECM) fungal diversity. We examined ECM fungal diversity on northern
red oak (Quercus rubra) seedlings grown in situ and in soil cores from
three northeastern U.S. forests dominated by two distinct tree species:
northern red oak (Quercus rubra) and eastern hemlock (Tsuga canadensis).
We hypothesized that seedlings growing in oak-dominated stands would be
colonized by different ECM fungal morphotypes and exhibit greater morphotype richness than seedlings growing in hemlock-dominated stands,
which are declining due to invasion by the hemlock woolly adelgid. Bait
seedlings of northern red oak were planted and soil cores were taken from
each of three oak- and hemlock-dominated stands located in a single watershed. Bait seedlings were grown for 20 weeks in the field. Oak seedlings
planted in the cores were grown for 24 weeks in a climate-controlled greenhouse. For both bait and greenhouse seedlings, ECM fungal composition
varied between oak- and hemlock-dominated stands. Moreover, ECM colonization and richness were significantly higher on bait and greenhouse
seedlings grown in oak- compared to hemlock-dominated stands. Reduced
ECM richness and colonization in hemlock-dominated stands was associated with significantly reduced bait seedling growth. These results suggest
that spatial patterns in ECM fungal community composition may reflect
forest composition or health. Further, ECM fungal community structure
may regulate oak recruitment in hemlock stands as hemlocks decline in
response to the hemlock woolly adelgid.

TWEDT, DANIEL J. and SCOTT G. SOMERSHOE.* USGS Patuxent

Wildlife Research Center, 2524 South Frontage Road, Vicksburg, MS,
USA. Breeding birds on reforested bottomlands in forested and agricultural landscapes.
In the Lower Mississippi Valley, restoration of bottomland forest adjacent
to mature forest has been advocated for conservation of forest birds. This
placement increases contiguous forest area and forest core habitats and
thereby benefits forest interior birds. However, density and demographic
response of birds to this landscape placement is largely unknown. We assessed avian densities and nesting success on 36 reforested bottomland sites
(2 - 16 years post-planting) within two different landscape contexts: (1)
sites that abutted large tracts of mature bottomland forest or (2) sites adjacent to agricultural fields that were distant from large tracts of mature
forest. Density was assessed from transect counts (2000) or spot-mapping
(2001, 2002). Grassland birds (Red-winged Blackbird, Dickcissel, Eastern
Meadowlark, Northern Mockingbird, and Mourning Dove) characterized
sites that abutted agricultural fields. White-eyed Vireo and Indigo Bunting,
birds typical of shrub-scrub habitats, characterized sites adjacent to mature
forests. Brown-headed cowbirds were more abundant near mature forests.
Generalized nest survival of all songbirds (17%) was similar on sites adjacent to agricultural fields and on sites abutting forests. Of 2142 nests that
had eggs or chicks, 611 fledged young. Estimated nesting successes (Mayfield) were: Yellow-breasted Chat (37%), Orchard Oriole (36%), Indigo
Bunting (29%), Northern Cardinal (27%), Yellow-billed Cuckoo (26%),
Dickcissel (18%), Mourning Doves (17%), Northern Mockingbird (14%)
Red-winged Blackbird (14%), and Common Grackle (6%). Predation was
the primary cause of nest failure. Overall parasitism rate was low (,4%)
but those species that characterized reforestation abutting mature forest
suffered higher levels of parasitism (White-eyed Vireo [36%]; Indigo Bunting [32%]). Vegetation height varied with the age of the site but mean tree
heights did not differ between landscape contexts. Tree density was greater
on reforested sites in forested landscapes than it was in agricultural landscapes. Bottomland restoration that abutted existing forest accelerated colonization by forest birds but nesting success was similar on reforested
bottomland sites regardless of their landscape context.
TYLER, ANNA C.,* EDWIN D. GROSHOLZ, JANIE C. CIVILLE and
JOHN G. LAMBRINOS. University of California - Davis, Davis, CA,
USA. Changes in carbon and nitrogen cycling following Spartina alterniflora invasion of Pacific estuaries.
The introduction of Spartina alterniflora into Willapa Bay, WA (WB) and
San Francisco Bay, CA (SFB) has impacted ecosystem primary production
and the cycling of carbon (C) and nitrogen (N). S. alterniflora was introduced into WB in the early 1900s, and is currently invading mudflats dominated by microalgae and the non-native intertidal seagrass, Zostera japonica, which has been in Willapa Bay since at least the 1950s. In 1975, S.
alterniflora was introduced to SFB; a subsequent hybridization with the
native S. foliosa has produced an aggressive hybrid. This hybrid is now
expanding into both unvegetated mudflats and native marsh vegetation.
Changes in C and N cycling are most dramatic where Spartina has replaced
mudflats. Sediment organic matter in unvegetated mudflats is historically
low, but labile (low C:N) in comparison to the Spartina-dominated areas
(high C:N). In WB, belowground C and N (living tissues only) are 10-25
times higher in S. alterniflora-dominated areas; collectively, S. alterniflora
and Z. japonica have increased the C and N storage in the intertidal sediments of the estuary (23,000 ha of mudflat + marsh) by approximately
7- and 2-fold, respectively. In SFB, the impact of Spartina on C and N
storage depends on the initial conditions: little change when compared to
Salicornia sp.; a 2-4 fold increase compared to S. foliosa; and a 50-300
fold increase when compared to bare mudflats. Senescence of aboveground
tissue results in the eventual release of 3E+05 kg N and 1E+07 kg C
into WB and 2E+04 kg N and 7E+05 kg C into SFB annually. The sheer
quantity of this input has profound implications for ecosystem function,
including system metabolism and food web structure.
TYRE, ANDREW J.,1,2,* BRIGITTE TENHUMBERG1,2 and C. MICHAEL
BULL.3 1 University of Queensland, St. Lucia, QLD, Australia; 2 University
of Nebraska-Lincoln, Lincoln, NE; 3 Flinders University of South Australia,
Adelaide, SA, Australia. Could density dependent engorgement success
regulate populations of ticks on Sleepy Lizards?
Density dependence in one or more demographic processes is often detected statistically as a relationship between a demographic process and

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339

density at small experimental scales. It is not always clear whether or not

such a relationship contributes significantly to population regulation at larger scales, especially if the life history is complex and subject to substantial
spatial and temporal heterogeneity. Recent reanalysis of population and
experimental data on Aponoma hydrosauri, an ectoparasitic tick found on
Sleepy Lizards (Tiliqua rugosa) in Australia, suggests that larvae of this
tick have reduced engorgement success as numbers of infesting larvae increase. We examined the potential of this process to regulate tick populations at larger spatial and temporal scales in the presence of strong spatial
and temporal heterogeneity with an individual based model of tick population dynamics. The results clearly indicate that the observed density dependent relationship is sufficient to regulate the population over long time
periods. The model also highlights the importance of understanding the
nocturnal refuge use behavior of the host lizards, as this can significantly
influence survival rates of life history stages that are waiting for hosts.
UDOVIC, DANIEL.* University of Oregon, Eugene, OR, USA. Phenological analysis of flowering and fruiting in Hesperoyucca whipplei.
Detailed tracking of the timing and spatial patterning of flowering and
fruiting in populations of Hesperoyucca whipplei provides a framework for
analyzing the effects of major selective agents on the evolution of the
plants reproductive strategy and the maintenance of highly specialized interactions with its mutualist pollinators (Tegeticula maculata) and with florivorous sap beetles (Anthoneus agavensis). In this presentation, I will (1)
describe a method for visualizing the fates of individual flowers as a function of their position on inflorescences, (2) describe how such positional
information can be used to develop models reconstructing a populations
flowering phenology, and (3) apply this approach to test the hypothesis that
the plants flowering phenology can act as a buffer against within-season
variability in pollinator abundance and overexploitation by sap beetles.
UNDERWOOD, NORA.* Florida State University, Tallahassee, FL. Population dynamics in a heterogeneous environment: Herbivore population response to variance in plant quality.
The quality of individual plants and mean quality of plant populations can
influence herbivore populations, but little is known about the effects of
variance in plant quality. Using field experiments with wild strawberry
(Fragaria chiloensis) and strawberry aphid (Chaetosiphon fragaefollii) I
examined how variance in plant quality affected aphid population dynamics. I identified plant genotypes with differing effects on aphid population
dynamics, and constructed plant populations that were monocultures of
individual genotypes or mixtures of the genotypes used in the monocultures. I followed aphid population dynamics on each of these plant population types in the field and fit population dynamic models to the resulting
time series. These data allow me to ask how dynamics on individual plant
genotypes in the monocultures relate to dynamics in herbivore populations
on mixtures of plants. Results suggest that the variance in quality among
plant genotypes affects aphid population dynamics. Estimated aphid population growth rates and carrying capacities on mixtures of plant genotypes
are often not the average of those parameters on monocultures of the genotypes included in the mixture. Depending on the particular genotypes
included in the mixtures, aphid population sizes can be either higher or
lower than the expected population size based on the average of the monocultures. Aphid movement among plants within a population appears to be
an important mediator of the effects of variance in plant quality on aphid
populations.
UNNASCH, THOMAS R.,1,* EDDIE W. CUPP,2 GEOFF HILL,2
CHARLES KATHOLI,1 HASSAN K. HASSAN,1 CHARLES S. APPERSON3 and HARRY M. SAVAGE.4 1 University of Alabama at Birmingham,
Birmingham, AL, USA; 2 Auburn University, Auburn, AL, USA; 3 North
Carolina State, Raleigh, NC, USA; 4 Centers for Disease Control and Prevention, Fort Collins, CO, USA. Host preference and arboviral encephalitis transmission in the Eastern USA.
Arthropod-borne encephalomyelitis viruses, including West Nile (WNV)
and Eastern Equine encephalomyelitis (EEE) virus represent a significant
and growing public health threats in the United States. These viruses are
primarily infections of the endemic avifauna and are maintained in an en-

340

Abstracts

zootic cycle by mosquitoes that feed primarily on birds. The viruses escape
this enzootic cycle to infect mammals through the activity of bridge vectors
with catholic feeding patterns. It is likely that an important variable in viral
amplification and escape from the enzootic cycle is the degree of contact
of the avian host and its mosquito vector. To investigate this, bloodfed
mosquitoes from confirmed EEE vectors were collected from a site in the
Tuskagee National Forest during an EEE outbreak in 2001. The bloodmeals
were identified to the species level and the proportion of bloodmeal derived
from each species compared to the overall composition of the avifauna at
the study site. EEE vector mosquitoes were found to demonstrate a marked
preference for certain bird species, and that this preference was not determined by the overall density or biomass of the avian species present at the
site. In particular, a single species, the Brown-headed Cowbird, was significantly over-represented in the bloodmeals, with over 40% of the bloodmeals having been derived from this species. Taken together, the data suggested that the vector mosquitoes were targeting young of the year and
species that inhabited the fringes of the study site. A similar host preference
pattern was demonstrated in vectors for WNV collected in three states
during the WNV outbreak of 2001. Taken together, these data suggest that
forest fragmentation and increases in the populations of the Brown-headed
Cowbird may result in an increase the potential for the development of
outbreaks of encephalomyelitis viruses, and in particular EEE.
VAITKUS, MILDA R.,* GEOFFREY M. HENEBRY, BRIAN C. PUTZ
and JAMES W. MERCHANT. Center for Advanced Land Management
Information Technologies (CALMIT), School of Natural Resources, University of Nebraska, Lincoln, NE, USA. Evaluating the use of statistical
decision trees for modeling avian habitats and regional range distributions in the Great Plains.
Attempts to regionalize species models by mosaicking range distributions
produced by neighboring state Gap Analysis Projects have proved problematic. Variations in habitat modeling have resulted in significant differences in predicted species distributions within and across state lines. The
use of national geospatial data to map surrogates of habitat enables a regional scope. Yet, there is a decided knowledge gap between the scales of
biogeography and those of wildlife management. Can the flexibility of statistical decision trees help fill this gap? We generated regional distributions
of 20 selected breeding birds in a six-state region (IA, KS, MN, ND, NE,
SD) using four recursive partitioning algorithms (CART, QUEST, CRUISE,
GUIDE). BBS route level summaries over two time periods (last 10 and
30 years) were used for the occurrence data (presence/absence and abundance). Environmental variables included land cover, daily climatic means
and variances, soil texture, and terrain. Multiple statistical decision trees
were generated for each target species to evaluate the relative strengths and
weaknesses of the different algorithms. Principal considerations were speed
of tree identification, interpretability of the cross-validated tree, and plausibility of the predicted range distribution resulting from tree inversion. As
expected, classification trees (from CART, QUEST, CRUISE) yielded predicted range distributions different from regression trees (from CART and
GUIDE). Unbiased variable selection in QUEST, CRUISE, and GUIDE
appeared to facilitate the identification of parsimonious, robust models and
plausible range distributions.
VAN DE BOGERT, MATTHEW C.,1,* JONATHAN J. COLE,1 MICHAEL
L. PACE1 and JAMES R. HODGSON.2 1 Institute of Ecosystem Studies,
Millbrook, NY; 2 St. Norbert College, DePere, WI. Whole-system metabolism in small lake ecosystems: Challenging continuous free-water oxygen methods with mechanistic models.
With the advent of reliable, continuous, free-water measurements of dissolved oxygen, researchers can directly measure net ecosystem production
which is the difference between gross primary production (total photosynthesis; GPP) and ecosystem respiration (R). Since R can be estimated from
dissolved oxygen and gas exchange at night, these continuous oxygen records can be used to estimate both GPP and R at relatively short time
scales (minutes). Physiologically based models provide independent estimates of GPP from photosynthetic parameters (Pbmax, alpha), phytoplankton
and/or periphyton biomass, and photosynthetically-available radiation. How
well do physiologically-based models perform in reproducing continuous
measurements of dissolved oxygen in the mixed-layers of small lakes? We

generated physiologically-based models for 4 lake basins for which we had

continuous oxygen data from sondes which recorded every 5 minutes for
about 100 d during the ice-free season. The lake basins differed in DOC
and food-web structure and one of the basins (Peter L. in 2002) was also
eutrophied by daily additions of N and P. The lake basins also received
daily additions of inorganic 13C which allowed us to further constrain carbon cycling parameters. Measured values of GPP varied from 31 to 104
mmol C m-2 d-1 among the basins and R from 32 to 79 mmol C m-2 d-1.
GPP/R ratios were less than 1 in all but the fertilized lake and NEP ranged
from -8 to +24 mmol C m-2 d-1. Physiological models using only pelagic
compartments failed to reproduce the diel pattern and magnitudes in measured dissolved oxygen. The addition of benthic GPP and R significantly
improved the fit. Inferred values of benthic GPP and R were large enough
to affect the spatial patterns and magnitude in dissolved oxygen. The diel
amplitude in oxygen increases dramatically from offshore to onshore, consistent with the models.
VAN GESTEL, NATASJA C., APRIL D. NESBIT, LUKE HALEY, ELLEN B. PEFFLEY and DAVID T. TISSUE. Texas Tech University, Lubbock, Texas. Effect of photoperiod on photosynthetic down-regulation
and carbon gain in Allium fistulosum.
We investigated the effect of increased source activity (24-h continuous
light) on the growth, photosynthesis and biochemistry of onion (Allium
fistulosum) by growing plants for 56 days in a 12-h (control) or 24-h photoperiod. Plants grown in 24-h light exhibited decreased leaf net photosynthesis (A) and photosynthetic capacity (Amax) compared with plants in 12h light, indicating photosynthetic down-regulation. The reduction in A and
Amax for leaves of 24-h plants was due to reductions in leaf N and increased
soluble sugar. The reduction in leaf N suggested that less N was available
to produce Rubisco. We suggest that the accumulation of sugars is due to
the inhibition of respiration in 24-h plants, thereby allowing carbohydrates
to accumulate in the leaf and initiate a feedback signal to reduce photosynthesis. These results demonstrate that carbohydrate utilization in respiration affects photosynthesis the following day. On a specific leaf area
basis, 24-h plants generally gained more carbon than 12-h plants because
they had a positive A over the entire 24-h period. However, 24-h plants
produced much less total leaf area, because 24-h plants may have allocated
their carbon to other functions (e.g. root exudates, volatile carbon emissions, growth of other plant organs, and other physiological processes).
Therefore, on a whole plant basis, 24-h plants gained less carbon than 12h plants. In conclusion, increased source activity (i.e. continuous light) was
detrimental to plant growth due to reductions in photosynthetic capacity
and changes in carbon allocation.
VAN MANTGEM, PHILLIP J.,1,* NATHAN L. STEPHENSON,1 MARYBETH KEIFER2 and JON E. KEELEY.1 1 U.S.G.S., Western Ecological
Research Center, Three Rivers, CA; 2 National Park Service, Three Rivers,
CA. White pine blister rust, fire and long-term demographic trends of
sugar pine in the Sierra Nevada.
An exotic pathogen, white pine blister rust, has caused declines in fiveneedled pines throughout western North America. While the range and
severity of white pine blister rust infection are relatively well known, longterm demographic data that document the effects of the disease are rare.
Simultaneously, fire exclusion may reduce recruitment opportunities for
these shade-intolerant pines. Fire exclusion has also allows high fuels accumulations, making it unclear how affected populations will respond to
the reintroduction of fire. We present population trends from 1,854 unburned and 321 burned sugar pine (Pinus lambertiana) individuals over 515 years at several burned and unburned sites at Sequoia and Yosemite
National Parks. Populations are declining at most unburned sites (l range:
0.82-1.04), and bootstrapped confidence intervals of l did not provide unequivocal evidence for population growth or stability at any site. Retrospective analysis suggests that declines are primarily a result of poor recruitment, although it is not clear whether small tree mortalities are most
closely tied to biotic attacks or resource competition. Data from 16 prescribed fires showed that fire caused sharp, but transitory, increases in tree
mortality, which does not appear to affect long-term population viability.
Our models suggest sugar pine will not become extinct at any of our sites
within 50 years. Nevertheless, some populations show rapid rates of decline

for a long-lived species, and additional stressors, such as increased pathogen virulence, or climatic change, could accelerate the rate of decline.
VAN NIEL, KIMBERLY P.1,2,* and MIKE P. AUSTIN.3 1 The Australian
National University, Canberra, ACT, Australia; 2 The University of Western
Australia, Perth, WA, Australia; 3 CSIRO Sustainable Ecosystems, Canberra, ACT, Australia. The effects of error in environmental variables on
predictive vegetation modelling.
The modelling of species responses to the environment and the prediction
of species geographical distributions have important implications for vegetation theory and conservation evaluation. However, error and uncertainty
can confuse these procedures and their results. The effect of digital elevation model (DEM) error on environmental variables, and subsequently
on predictive vegetation models, is often acknowledged as a concern in
modelling, but has not been explored. Based on an error analysis of a DEM,
multiple error realisations of the DEM are created and then used to develop
both direct and indirect environment variables for input to predictive vegetation models. The effects of this error and the resultant uncertainty of
results are explored in the context of the typical steps in the modelling
procedure for prediction of forest species presence/absence on the south
coast of New South Wales, Australia. Results indicate that all of these steps
and results, including the statistical significance of environmental variables,
shapes of species response curves in generalised additive models (GAMs),
stepwise model selection, coefficients and standard errors for generalised
linear models (GLMs), prediction accuracy (Cohens kappa and overall accuracy), and spatial extent of predictions, are greatly affected by this type
of error. Unconsidered error in the DEM can affect the reliability of interpretations of model results and level of accuracy in predictions, as well as
the spatial extent of the predictions.
VAN ZANDT, PETER A. University of Toronto, Department of Botany,
25 Willcocks St., Toronto, ON, Canada. Community-level impacts of induced plant responses to herbivory.
Numerous forces affect herbivorous insect communities, but the role that
induced plant responses play in altering community structure has been little
studied. To better understand the potential for early season herbivory to
affect subsequent herbivore species richness, I conducted field experiments
over two years involving the community of milkweed herbivores. Plants
were either undamaged (controls) or damaged by one of three herbivores:
monarchs (Danaus plexippus), milkweed leaf beetles (Labidomera clivicollis), or weevils (Rhyssomatus lineaticollis). Following treatment, naturally colonizing herbivores were counted on all plants for two months to
assess host plant usage and herbivore species richness. In 2001, monarch
damaged plants had 26% greater species richness, while weevil damaged
plants had 17% lower richness than undamaged plants. In 2002, herbivory
by monarchs reduced subsequent insect species richness on host plants by
33%, but weevil damaged plants did not differ from controls. In 2001,
monarch treated plants exhibited induced susceptibility, with over 6 times
more damage compared to controls, while in 2002, weevil damaged plants
received nearly 4 times more damage. In general, colonizing herbivores
were less likely to be found on weevil treated plants in 2001, whereas in
2002 they were attracted to weevil treated plants. The net effects of initial
weevil treatment were unaffected by subsequent plant damage, while the
effects of monarchs dissipated in strength as natural herbivore damage accumulated. This suggests that induced responses to initial weevil feeding
persisted through the season while monarchs were more likely to affect
colonization via alterations in the trajectory of community assemblage.
Therefore, induced plant responses to herbivory can substantially alter herbivore community structure, but the nature of the effects are likely to depend on the identity of the initial herbivore.
VANCE, RICHARD R.1,* and ANDREW L. NEVAI.2 1 University of California at Los Angeles, Los Angeles, California, USA; 2 University of California at Los Angeles, Los Angeles, California, USA. Competition for
light between plant species with different foliage profiles: Formulation.
Mathematical study of how differences in plant architecture might influence
interspecies competition for light requires population growth equations that
include the benefits and costs of plant architectural features and yet remain

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341

simple enough to analyze. We will describe the formulation of such a

model, beginning with the single-species case. This formulation incorporates explicit descriptions of three essential plant features: the vertical distribution of leaves and the vertical gradient in light intensity it creates, the
photosynthetic response of leaves to light intensity throughout this gradient,
and the metabolic cost of growing and maintaining stems whose lengths
determine the vertical leaf distribution. These descriptions give rise to a
population growth model that consists solely of a scalar ordinary differential equation of the Kolmogorov form, dx/dt 5 xf(x). The measure x of
population abundance is the leaf area index, i.e., the total leaf area deployed
by the plant population above each unit area of ground surface. Relatively
direct scrutiny of the specific growth rate function f reveals the qualitative
manner in which each feature of plant structure and function influences
population growth, persistence, and equilibrium abundance. Straightforward extension of this formulation to two competing species sets the stage
for examining how architectural differences might facilitate plant species
competitive coexistence.
VANCE-CHALCRAFT, HEATHER D.1,2,* and DANIEL A. SOLUK.1,2
1
University of Illinois at Urbana-Champaign, Urbana, IL; 2 Illinois Natural
History Survey, Champaign, IL. Prey density can influence the effects
of multiple predator species on the preys predation risk.
Changes in prey density can influence the consumption rate of a single
predator but it is not clear how prey density influences our ability to predict
the combined consumption rates of multiple predator species. These combined consumption rates influence the distribution and abundance of prey.
To determine whether interactions among predator species affect the preys
risk of predation at varying prey densities, we conducted experiments in
artificial stream tanks that crossed 6 treatments consisting of 0, 1, or 2
predator species with three treatments varying in prey density. Our predators were hellgrammites, greenside darters, and creek chubs and our prey
were mayfly larvae. We found that each predator species, when alone,
consumed an approximately equal proportion of prey at all prey densities.
The consumption rates of the multiple predator combinations increasingly
deviated from our models predictions as prey density increased. Thus,
making generalizations about the preys risk of predation in the presence
of multiple predator species may be difficult based on single prey density
experiments. Empirically derived functional response curves of multiple
predator treatments may be needed in order for prey density to be incorporated into models that predict the combined impact of multiple predators
on prey populations.
VANDERMAST, DAVID B.,* JASON D. FRIDLEY, DANE KUPPINGER
and ROBERT K. PEET. University of North Carolina - Chapel Hill, Chapel
Hill, North Carolina. Assessment of habitat specialization of southeastern trees using large-extent co-occurrence data.
Much ecological theory is based on the characterization of ecological habits
of species as generalist or specialist, but standard measures for placing
species along a generalist-specialist gradient do not exist. We have developed a novel approach to quantifying degree of habitat specialization by
using species co-occurrence data. Generalists should co-occur with many
species across their range, while specialists should co-occur with relatively
few species, given equal plot occurrences. We quantify this concept using
a beta diversity statistic whereby a constant number of plots containing a
given species are randomly selected, and mean plot richness is subtracted
from the total number of species in the sample to obtain a species-centered
beta diversity statistic. This calculation is repeated via permutation to obtain a mean beta value that places a species along a generalist-specialist
gradient. Species that tend to occur in single habitats with the same species
are specialists and have low beta values; a higher beta value indicates high
species turnover among plots containing the focal species and thus that the
species is a generalist. We introduce our approach with the large woody
flora (. 10 cm dbh) of the Southeast using 2500 vegetation plots from the
Carolina Vegetation Survey, and rank the generalist-specialist tendencies
of 112 common species. Common understory forest species (American
holly, ironwood) have among the highest beta values, and species associated with few habitat types (longleaf pine, bald cypress) are among the
lowest. On average, generalist species are deciduous, shade-tolerant, and
have short lifespans and bird-dispersed seed. Generalism is also correlated

342

Abstracts

with the range of species along certain environmental gradients, including

annual temperature, potential evapotranspiration, and soil pH and organic
matter. We conclude that applying our co-occurrence-based approach to
large-extent plot databases provides a robust means to rank species on a
generalist-specialist gradient.
VANNI, MICHAEL J.,1,* WILLIAM H. RENWICK,1 MARIA J. GONZALEZ1 and SCOTT HALE.2 1 Miami University, vannimj@muohio, Oxford, OH; 2 Inland Fisheries Research Unit, Ohio Division of Wildlife, Hebron, OH. Response of an agriculturally impacted reservoir ecosystem
to improved watershed management.
Agricultural practices are changing in landscapes throughout the world.
However, implications of these changes for downstream lakes are not well
documented, particularly for reservoirs, which receive large inputs of sediments and nutrients from watersheds. We assessed effects of improved
agricultural practices on a eutrophic reservoir (Acton Lake, Ohio) that resides in an agricultural watershed. During this 9-year period (1994-2002),
the percentage of agricultural land implementing conservation tillage increased from about 15% to about 70%. Associated with changes in tillage
practices, concentrations and export rates of suspended sediments and phosphorus (P) in streams draining into Acton Lake declined markedly (5-15%
per year). These trends are consistent with known effects of conservation
tillage, which tends to reduce soil erosion and P export. Despite large
decreases in watershed inputs of P (the limiting nutrient), phytoplankton
biomass in Acton Lake increased over this period. This increase was most
likely caused by reduced light limitation, due to decreased concentrations
of suspended inorganic sediment in the lakes water column. From 1994 to
2002, concentrations of inorganic suspended sediment in the lake, and attenuation of light by this sediment, have declined by about 50%. In the
early years of this study, attenuation of light by sediment exceeded attenuation of light by phytoplankton self-shading, but currently phytoplankton
attenuate more light than sediment. In addition, photosynthesis-irradiance
curves and experiments show that Acton Lake phytoplankton are frequently
light limited. Phytoplankton can maintain high biomass in the face of decreased watershed P inputs because of P translocation by sediment-feeding
fish (gizzard shad). These fish consume P from sediment detritus and excrete dissolved P into the water column. We expect that in future years,
phytoplankton biomass will eventually decrease as watershed P inputs continue to decline, and gizzard shad biomass decreases in response to decreased sediment inputs.
VARNER, J. M.,1,* FRANCIS E. PUTZ,1 DORIA GORDON,1,2 J. K.
HIERS,3 ROBERT J. MITCHELL4 and ROGER OTTMAR.5 1 University
of Florida, Gainesville, FL, USA; 2 The Nature Conservancy, Gainesville,
FL, USA; 3 Eglin Air Force Base, Niceville, FL, USA; 4 Joseph W. Jones
Ecological Research Center, Newton, GA, USA; 5 USDA Forest Service
Pacific NW Research Station - FERA, Seattle, WA, USA. Restoring fire
to long-unburned old-growth southeastern pine savannas: new approaches for novel forests.
Traditional models of southeastern coastal plain pine savannas are based
on pristine ecosystems now rare in the contemporary landscape. Instead of
open pine savannas with high groundcover richness frequented by lowseverity surface fires, greater than half of all remnants are degraded and
fire-excluded. To fill the gap between these understandings, we reviewed
the literature and investigated natural experiments of fire exclusion in
southeastern longleaf pine (Pinus palustris) savannas. Our primary interests
were the influence fire exclusion has had on vegetation structure and composition, its role in the advent of forest floor accumulations, and outcomes
of fire re-introduction. Fire excluded pinelands have shifted structurally
from nearly mono-specific open savanna-woodlands to closed canopy
mixed pine-hardwood forests. Both overstory and understory species composition has changed; canopies are more diverse, groundcovers more depauperate. Without frequent surface fires, forest floor accumulations have
developed deep (5-50 cm), densely rooted decomposition horizons (Oe and
Oa) absent in frequently-burned stands. In fire re-introductions, resulting
overstory pine mortality has been substantial (from 10 to 100% of larger
individuals), a major deterrent to further restoration. We review four primary hypotheses related to pine death resulting from fire re-introduction:
(1) smoldering Oe and Oa horizons damage vascular tissues, (2) smoldering

Oe and Oa horizons damage organic and surficial mineral soil roots, (3)
associated flaming combustion damages canopy meristems, and (4) firecaused damage and stresses lead to whole-tree carbohydrate deficits. Restoration of long-unburned pineland ecosystems will require more understanding of vegetation dynamics, pine physiological response, and smoldering fire ecology; all research areas that have received little attention
considering the extent and importance of this novel forest.
VELLEND, MARK,* JONATHAN A. MYERS, SANA GARDESCU and
PETER L. MARKS. Cornell University, mv59@cornell.edu, Ithaca, NY,
USA. Seed dispersal by white-tailed deer: Implications for long-distance dispersal and plant migrations.
For many plant species in eastern North America, observed seed dispersal
distances (ranging up to only a few tens of meters) fail to explain inferred
rates of post-agricultural colonization and post-glacial migration. This discrepancy points to a substantial gap in our knowledge concerning the mechanisms by which seeds are dispersed long distances. Here we report seed
dispersal of .70 plant species via ingestion and defecation by white-tailed
deer in central New York, USA. This is the first comprehensive study of
seed dispersal by white-tailed deer, despite a vast body of research on other
aspects of white-tailed deer ecology. The plant species dispersed by deer
include herbs, shrubs, and trees from a variety of habitats, and the full
range of presumed dispersal modes in eastern North America (ants, wind,
vertebrates, ballistic, and unassisted). We found a mean of .30 germinable
seeds per pellet group, and seeds were dispersed during all months of the
year. The majority of seeds found possessed no obvious adaptations for
dispersal. Notable taxa included Trillium grandiflorum, an otherwise antdispersed forest herb, and four of the top 20 invasive plants of greatest
concern in New York State: Lonicera spp., Rosa multiflora, Rhamnus cathartica, and Lythrum salicaria. A deer-generated seed shadow modeled
using data from the literature on movement patterns and gut retention times
revealed that most deer-dispersed seeds should travel several hundreds of
meters, and occasionally .3 km. We conclude that white-tailed deer represent a significant and previously unappreciated vector of seed dispersal
across the landscape, likely contributing an important long-distance component to the seed shadows of hundreds of plant species, and providing a
mechanism to help explain rapid rates of plant migration.
VERBURG, PAUL,* JAY ARNONE and DAVID SCHORRAN. Desert
Research Institute, Reno, NV. NSF-IRCEB interannual climate variability and ecosystem processes in tallgrass praire: Soil respiration.
Soil respiration represents one of the main C fluxes from terrestrial ecosystems to the atmosphere. Yet it is unclear how interannual climate variability will affect soil respiration and how long effects of an anomalously
warm year on soil respiration will persist. In order to address this problem,
we are measuring soil respiration in twelve intact tallgrass prairie monoliths
(divided over four environmentally controlled growth chambers) using dynamic and static chambers. Measurements started in the spring of 2002
when temperatures in all growth chambers were similar. First results indicate that, despite variations in soil respiration rates between monoliths,
average soil respiration rates were similar in each growth chamber. These
results show that no pre-treatment differences existed between growth
chambers despite variability in plant community structure. In addition, measurements of dynamic chambers agreed well with static chamber measurements. Soil respiration rates showed clear seasonal patterns and strong increases in soil respiration were observed after watering events. In February
2003, temperature in two growth chambers was increased by four degrees.
Results of the first six months of this warming treatment on soil respiration
will be discussed.
VERMILLION, BRIAN, INDU BALA, WILLIAM RETZLAFF and
RICHARD BRUGAM.* Southern Illinois University Edwardsville, Edwardsville, IL, USA. Industrial development on the American Bottomsa test of stable isotope analysis in paleolimnology.
Many investigators have shown that lakes in industrial areas contain sediment records of past pollution. Techniques of stable isotope analysis have
only recently been applied to the interpretation of these records. We compared the stable isotope record with pollen, diatom, and chemical analysis

of sediment. We studied Horseshoe Lake on the American Bottoms, a

floodplain of the Mississippi River east of St. Louis, MO. Horseshoe Lake
is an oxbow lake located in Granite City, IL, a town with two Superfund
sites and an integrated steel mill. Fifteen sediment cores and 84 surface
sediment samples have been collected from Horseshoe Lake. Five of these
cores were dated using pollen analysis and 210Pb. Diatom analysis shows
that in pre-settlement times the lake was clear and shallow with lots of
macrophytes. The modern lake is very eutrophic. In the late 1930s and
40s, Pb, Zn, and Cd sediment concentrations increased. The isotopic composition of the Pb indicates that it originates from industrial contamination.
During the 1920s, Si, Al, and Ti (elements indicative of erosion) decrease
as Ca increases in concentration. An increase in d15N above 10 o/oo during
the 1920s also indicates contamination of the lake by sewage or animal
waste. We hypothesize that the increase in Ca results from increased deposition of CaCO3 resulting from the eutrophication of Horseshoe Lake by
sewage inputs.
VIDRA, REBECCA L., GEORGE R. HESS and THEODORE H. SHEAR.
North Carolina State University, Raleigh, NC. Can we design urban forest corridors to resist invasion by exotic plant species?
Management and restoration of native ecological communities would be
greatly enhanced with knowledge of characteristics that make them resistant to exotic species invasion. This is particularly important in urban areas,
where remnant habitats provide important ecological functions but are often
degraded by invasive species. While many studies have addressed the influences of disturbance, resource availability, and species richness on invasibility, certain structural features of habitat patches are also thought to
control patterns of exotic species invasion. For example, larger patches of
intact habitat are thought to be less susceptible to invasion than small
isolated fragments with high edge to area ratios. The character of the surrounding landscape may also affect invasion dynamics. We studied the
influence of landscape structure and context on the invasion of forested
riparian corridors. Specifically, we examined the relationship between the
distribution of exotic plants in forested greenways and the nature of the
surrounding landscape, including width and surrounding development density. We used a modified belt transect approach to record presence, frequency, and percent cover for 16 invasive exotic plant species in each
corridor. The frequency of occurrence and mean percent cover of four common species, Microstegium vimineum, Ligustrum sinense, and Lonicera
japonica, were significantly negatively correlated with corridor width. Several species were positively correlated with development density, but an
interaction between width and context warrants further exploration. Here
we present these results, with special attention to the distribution of M.
vimineum, a pervasive exotic annual grass in these riparian corridors. We
also examine the within-site variability in exotic species presence and abundance to account for local-scale characteristics that may promote invasion.
The results of this study can guide forest corridor protection and restoration
in urbanizing areas, where there are many sources of invasive propagules
and few opportunities for preservation.
VIERLING, KERRI T.* and DALE J. GENTRY. South Dakota School of
Mines and Technology, Rapid City, SD. The influence of burn age on
Lewis9s Woodpecker (Melanerpes lewis) reproductive success and productivity.
Lewis9s Woodpeckers (Melanerpes lewis) have been characterized as burn
specialists due to their preference of nesting within burned pine forests.
Saab and Vierling (2001) suggest that recently crown-burned pine forests
might function as source habitat for this species; however, the importance
of a burned forest to Lewis9s Woodpecker reproduction might change
throughout time with increased understory development and changes in
snag availability. While Lewis9s Woodpeckers are noted to occur both in
young and old burns, the relative importance of old burns (i.e. .10 years
post fire) to Lewis9s Woodpecker populations is not known. The purpose
of this study was to compare reproductive success and productivity of
Lewis9s Woodpeckers breeding in an old crown-burned pine forest with
published accounts of breeding activities in young crown-burned pine forests. In 1991, the Shirttail fire burned approximately 1700 ha in Wind Cave
National Park and the Black Hills National Forest, South Dakota. We surveyed two 300ha plots in the Shirttail burn during the summer of 2002

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343

and located Lewis9s Woodpecker nests in these areas. We located 16 nests

and monitored them every 3-4 days until they failed or fledged. Productivity in the Shirttail burn was higher than in the young burn (2.3 fledglings
vs. 1.8 fledglings), and reproductive success was similar (76% in the old
burn vs. 78% in the young burn). Predation was a major cause of nest
failure in both young and old burns. The results of this study suggest that
older crown-fire burns may provide important breeding habitat for Lewis9s
Woodpeckers.
VIERLING, LEE A.,1,* JEFF FRYKHOLM2 and GEORGE GLASSON.3
1
South Dakota School of Mines and Technology, Rapid City, SD,
Lee.Vierling@sdsmt.edu; 2 University of Colorado, Boulder, CO; 3 Virginia
Tech, Blacksburg, VA. The Earth Systems Connections elementary curriculum: Contextual learning opportunities for building ecological
knowledge.
As humans continue to subject the Earth9s dynamic and interdependent
systems to greater levels of stress and demand, it is paramount that educators help young children appreciate the significance and interrelated nature of ecological systems. The Earth Systems Connections (ESC) curriculum project aims to provide integrated learning opportunities for primary
children across a diverse range of cultural backgrounds that will help them:
1) identify and recognize the interconnected nature of the Earth9s systems;
2) appreciate technological tools (e.g. remote sensing imagery and analysis
techniques) that scientists use to conduct earth system science; 3) recognize
the extent to which mathematics, science, and technology are not only
interrelated, but also can help us understand natural phenomena; and 4)
cultivate a spirit of curiosity and confidence in themselves as emerging
thinkers and scientists who will be responsible for understanding and caring
for the Earth in the decades to come. The ESC curriculum is being designed
around seven themes, each representing a different perspective of earth
system science that encompasses several disciplines yet is appropriate in
scope for elementary aged children. The content of each lesson is presented
in contexts that require students to gather and assimilate ecological information not only through science and mathematics, but social studies, art,
and the language arts as well. The initial phase of the ESC curriculum will
provide teachers with a core of 50 thematic, integrated, yet stand-alone
lessons (containing over 100 activities) that are connected across K-5 grade
levels. While lessons are designed with pedagogical and content innovation
in mind, each reflects specific national content and process standards for
elementary level mathematics and science education.
VILE, DENIS,1,2,* BILL SHIPLEY1,* and ERIC GARNIER.2,* 1 Universite
de Sherbrooke, Sherbrooke, Quebec, Canada; 2 Centre dEcologie Fonctionnelle et Evolutive, CNRS, Montpellier, France. Specific leaf area and
dry matter content estimate leaf thickness in laminar leaves.
Leaf thickness is a functional trait related to leaf and plant functioning, and
which has an impact on the ecological distribution of species. Thickness
is not straightforward to measure however, due both to the diversity of
structure among leaves, and to the heterogeneity of the trait within the leaf
itself. In laminar leaves, it can be demonstrated that average thickness is
the inverse of density 3 SLA 3 LDMC; (SLA, specific leaf area 5 leaf
surface area to dry mass ratio; LDMC, leaf dry matter content 5 leaf dry
mass to fresh mass ratio). Assessing that density is constant across species,
we tested whether leaf thickness can be estimated from SLA and LDMC
which are easier to measure, and included in large scale screening programmes of comparative ecology. This was done at the interspecific level
for a total of 1047 data points, using both original and published data sets
from widely-differing geographical areas. We fit the following general linear covariance model: log(measured thickness) ; dataset + log(1/(SLA 3
LDMC) + interaction. As predicted by the approximation, T 5 (SLA 3
LDMC)-1, there was no significant difference in the intercepts across studies, the overall intercept was not significantly different from zero (indicating a mean leaf density of 1), and the slope associated with log(SLA 3
LDMC) did not differ significantly from -1. We conclude that a simple
estimation of average thickness could be accurately given by the inverse
of SLA 3 LDMC, also called the Hughes constant (K 5 projected surface
/ fresh mass). Although these findings suggest that the average leaf density
is close to unity, that of individual species varies around this value, which
produced some scatter around the relationship. This scatter can therefore
be interpreted in relation to the tissue composition of leaves.

344

Abstracts

VILE, MELANIE A.,1,* PAMELA A. MATSON2 and LARS O. HEDIN.1

1
Department of Ecology and Evolutionary Biology, Princeton University,
Princeton, NJ, USA; 2 Department of Geological and Environmental Sciences, Stanford University, Stanford, CA, USA. Thermodynamic controls
on soil trace gas (CH4, CO2, N2O) fluxes from the Maui rainfall gradient.
Predictions of ecosystem-scale emissions of trace gases are mired in large
temporal and spatial variability. Much of this variability is likely caused
by thermodynamic factors operating at the microsite scale. We wanted to
determine if a thermodynamic perspective can enhance our mechanistic
understanding and predictive capability of trace gas emissions at sites on
the Maui rainfall gradient, where all state factors are constant but rainfall
varies between 2200 mm, and 4050 mm. In laboratory incubations, we
manipulated water and carbon (C) in soil cores from both sites to test the
importance of these two thermodynamic factors in constraining trace gas
fluxes. Both water, and water plus C additions increased CH4 production
significantly at the drier site, and caused a shift from net CH4 consumption
to net CH4 production. At the wet site, both addition treatments caused
significant increases in CH4 production, with net production of CH4 measured in all cores, including controls. Nitrous oxide production showed the
same responses as CH4 to water and water plus C additions with a much
larger increase in production measured at the drier site. Soils from the drier
site always showed net CH4 production while water and water plus C treatments provoked a shift from net NO2 consumption to production at the wet
site. Production of CO2 decreased significantly in both water and water
plus C-amended cores relative to controls at both sites. Production of NO2,
CH4 and CO2in water and water plus C treatments were statistically indistinguishable at both sites. These results suggest that water has a greater
thermodynamic control than labile C over trace gas emissions from soils
along the Maui rainfall gradient.
VINTON, MARYANN* and LEANNE VIGUE.* Department of Biology,
Creighton University, Omaha, NE, 68178. The role of plant-soil feedbacks in the persistence of reed canary grass.
Reed canary grass (Phalaris arundinacea) is a persistent and invasive species of tallgrass prairie lowlands. We performed field experiments to evaluate the role of soil nitrogen and litter decomposition in the growth and
persistence of this species relative to native tallgrass species. Soil manipulations involved nitrogen fertilization with ammonium nitrate and lowered
soil nitrogen treatments with the addition of high carbon materials, sucrose
and sawdust. Litter bags were constructed using recently produced litter of
big bluestem and reed canary. Reed canary grass showed a stronger positive response to added nitrogen than did native species. The growth of both
reed canary grass and native species seemed to be affected by a preemption
of soil nitrogen by weedy dicots that eventually dominated many plots.
Soil carbon additions affected plant-available nitrogen in complex ways
and may not have the desired effect of lowering nitrogen mineralization.
Reed canary grass litter likely persists longer than that of other species and
this mechanism may be key to its ability to dominate sites.
VITOUSEK, PETER M.,1,* OLIVER CHADWICK,2 THEGN LADOFOGED,3 MICHAEL GRAVES,4 SARA HOTCHKISS,5 PATRICK KIRCH,6
PAMELA MATSON1 and SHRIPAD TULJAPURKAR.1 1 Stanford University, Stanford, CA, USA; 2 University of California - Santa Barbara,
Santa Barbara, CA, USA; 3 University of Washington, Seattle, WA, USA;
4
University of Hawaii - Manoa, Honolulu, HI, USA; 5 University of Wisconsin - Madison, Madison, WI, USA; 6 University of California - Berkeley, Berkeley, CA, USA. Climate, nutrient availability, and the distribution of pre-contact dryland agriculture in Hawaii.
Hawaiians developed and maintained intensive dryland agricultural systems
for hundreds of years prior to European contact. One such system occupied
55 kilometers squared on the leeward flank of Kohala Volcano, the oldest
volcano on the island of Hawaii. This leeward slope supports one of the
steepest rainfall gradients on Earth, from ,200 to .3000 mm/yr in less
than 15 km, and remains of the Kohala field system fill the area receiving
between 750 and 1500 mm/yr. As part of an effort to understand relationships among soil fertility, agricultural intensification, human effects on ecosystems, and the development of social and cultural complexity in Hawai-

ian society, we determined soil characteristics along several transects across

the field system. The soils of areas above (wetter than) the field system are
infertile, with low base saturation and available P averaging 3.9 ug/g. Within the field system, soil P concentrations are much greater, averaging 124
ug/g, while soils in dry sites below the field system have high base saturation and intermediate available P (26 ug/g). Intensive agriculture here
appears to have been poised between conditions that are too wet (and so
infertile) and too dry (and so water-limited).
VOGELSANG, KEITH M.* and JAMES D. BEVER. Indiana University,
Bloomington, IN. The interaction of native plants and mycorrhizal fungi on plant growth and soil stabilization processes.
Native plant communities are increasingly threatened by introduced species, some of which exhibit invasive qualities. Our work with California
grassland species suggests that ecological differences exist among herbaceous native and exotic plants, and that these differences ultimately lead
to changes in ecosystem function. Specifically, exotic species appear less
dependent on arbuscular mycorrhizal fungi than native species. We tested
the community and ecosystem consequences of these differences using manipulated mesocosms. These mesocosms were inoculated with soil communities derived from native-dominated sites and then cultured with assemblages of native or exotic host plants. The divergence of the mycorrhizal communities under their respective host communities caused patterns
of differential growth in two related indicator species: Gnaphalium californicum, a California native forb; and Carduus pycnocephalus, an invasive
exotic thistle. The native indicator grew best with soil microbes cultured
with native plant communities, while the exotic indicator grew relatively
better with soil microbes cultured with exotic plant communities. Conversely, the exotic indicator was strongly inhibited by soil microbes from
native plant communities. These same mycorrhizal communities also influenced the stability of soil macroaggregates, which form due to the interaction of plant roots, mycorrhizal fungi, and other soil microbes. Stable
soil aggregates improve water infiltration, are less erodible, and are associated with multiple ecosystem processes. We show that soil aggregate
stability is higher in soil cultured with native plant species than in soil
cultured with exotic plant species. Using soil microbes from unmanaged
native and exotic plant communities, we also find that soil aggregate stability increases more in response to inoculants derived from native communities. Our results suggest an important link exists between patterns of
species introductions and below-ground ecosystem processes.
VOLIN, JOHN C.,* MICHAEL J. LOTT, JORDAN D. MUSS, DIANNE
OWEN and JOY E. STEWART. Florida Atlantic University, Davie, FL.
The life history of the invasive fern Lygodium microphyllum: from reproductive strategy to landscape dispersal.
We studied the life history of Lygodium microphyllum, a highly invasive
non-indigenous fern species with a kudzu-like growth form that is currently
spreading rapidly throughout southern Florida. Examination of its reproductive biology showed that L. microphyllum is capable of using all three
mating systems: intragametophytic selfing, intergametophytic selfing, and
outcrossing. This complex reproductive biology is highly unusual for the
vast majority of fern species studied to date. The ability of L. microphyllum
to utilize all three mating systems lends support to our hypothesis that its
reproductive strategy partially explains its invasive spread across the landscape. In addition, we found that its invasiveness is enhanced by the ability
of a female gametophyte to determine the sex of nearby gametophytes,
making them male and thus ensuring cross-fertilization. The distribution of
L. microphyllum was also examined at the community level in forested
wetland sites within the Big Cypress Swamp. The observed patterns indicated that L. microphyllum is predominantly found in moderately hydric
conditions; relatively low elevation, deep substrate and high substrate water
content were significantly related to its presence. We also developed a
cellular automaton dispersal model for L. microphyllum. The model was
calibrated using actual data from 1978 to 1993, then validated independently using actual data from 1993 through 1999. A projection to 2009
used actual 1999 flight survey data as a starting point and showed an
alarming increase in L. microphyllum establishment across South Florida,
in particular in the cypress-dominated wetlands of the Big Cypress Swamp.

VON HOLLE, BETSY,* DAVID R. FOSTER and GLENN MOTZKIN.

Harvard Forest, Harvard University, vonholle@fas.harvard.edu, Petersham,
MA. Historic and current influences on habitat invasibility in a mosaic
landscape: Cape Cod National Seashore.
Effects of non-native species are a hazard to global biodiversity, second
only to habitat destruction. For informed management decisions, we must
determine factors that contribute to ecological resistance to biological invasion. Habitat invasibility to plant invaders was investigated in the highly
resistant Cape Cod ecosystem through a spatially-explicit study of historical
disturbances and current environmental and biotic properties of 20 x 20m
field plots, randomly located across the landscape. These 352 plots were
within natural areas, half within Cape Cod National Seashore, MA. This
region experienced intensive land use during European settlement of the
area, followed by large-scale reforestation in the last century and fire suppression in the last 60 years. Historic disturbances, current vegetation,
edaphic properties and other environmental conditions were determined for
each plot using historic and field methods. These factors were assessed for
their influence on exotic species richness and cover with multiple linear
regressions and canonical correspondence analyses. The most influential
factors for non-indigenous plant cover and richness were current soil nutrient conditions. Plots that had been plowed or burned in the past did not
have significantly greater richness of non-indigenous species. While land
use history in Cape Cod National Seashore is a strong determinant of native
community assemblages, this is not the case for non-native species in natural areas. The resistance of these natural areas may be due to the harsh
conditions provided by the depauperate soils in this area.
VONESH, JAMES R.,* CRAIG W. OSENBERG and BENJAMIN BOLKER. Department of Zoology, University of Florida, Gainesville, Florida.
Multi-predator effects across life-history stages: Non-additivity of eggand larval-stage predation in an African treefrog.
Recent studies of multiple predator effects on shared prey indicate that
aggregate predator effects on prey are frequently less than expected from
their independent effectsa result referred to as risk reduction. However,
studies to date have focused on interactions of predators of a single lifestage, where risk reduction results from interactions among predators such
as intraguild predation. For prey that undergo ontogenetic habitat shifts,
multiple predators may feed on different life stages that occupy different
habitats and may not interact directly. Thus, the primary mechanism that
gives rise to risk reduction in previous studies is absent. However, predator
effects across stages, habitats, or sizes may still be non-additive due to
indirect effects mediated through prey traits (e.g., size, morphology, behavior) and density. We examined the effects of sequential stage-specific
predators of the arboreal eggs and aquatic larvae of the African reed frog,
Hyperolius spinigularis. To quantify the overall effect of egg and larval
predators and isolate the mechanisms leading to emergent effects, we conducted an experiment in which we mimicked the effects of the egg predator
on initial tadpole density and initial larval size/age in the presence and
absence of aquatic larval predators. Egg and larval predator effects were
non-additive: more Hyperolius survived than predicted from their independent effects. Egg-stage predator effects on both larval density and size/age
contributed similarly to reduce the effectiveness of larval-stage predators
to a level indistinguishable from background mortality. Because many animals from diverse taxa exhibit complex life histories, non-additive multipredator effects across stages and habitats are likely common in nature. In
addition, it appears that risk reduction may arise through a variety of mechanisms and thus may be a common feature of many within- and acrossstage multi-predator systems.
VOSE, JAMES M.* Land and Water Interactions in Southeastern
Landscapes.
Water is a key integrator of ecosystem processes in the southern U.S. High
rainfall, coupled with frequent high intensity storm events, results in ecologically significant interactions among terrestrial and aquatic ecosystems.
Water quality within the 1,530,000 km of rivers in the south reflects the
cumulative impacts of historical and contemporary natural and anthropogenic disturbances. Recent assessments of surface water quality and biotic
integrity south-wide suggests that water quality has been improving over

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345

the past several years; however, a variety of contemporary land disturbances are impacting the health and integrity of southeastern watersheds.
The magnitude, duration, and ecological consequences of these impacts
varies considerably across the south, beginning in the steep/high gradient
streams in the mountains, passing through the low gradient streams in the
piedmont, and ending in the wetland dominated coastal plain region. The
importance of land-water interactions is reflected by the significance of
non-point source (e.g., roads, construction sites, agricultural runoff) contributions to degraded water quality and overall biotic integrity. Rapid human population growth in the southeastern region is accelerating land disturbing activities and has the potential to significantly reduce water quality
in the future. In addition, most of the southern U.S. depends upon surface
water as the primary source of drinking water. Conflicts among demands
for clean and plentiful drinking water, land development pressures, and
maintenance of watershed health and integrity has major sociological implications for the southern U.S.
VOSS, JOSHUA D.* and LAURIE L. RICHARDSON. Florida International University, Department of Biological Sciences, Miami, FL, USA.
Environmental factors and coral community composition associated
with coral diseases in the northern Florida keys and Bahamas Exuma
Chain.
The increase in the number of coral diseases and the expansion of their
ranges over the past 25 years pose serious threats to the health of coral
reef ecosystems worldwide. Thus far, little is known about coral epidemiology or the conditions that promote coral infection. While some studies
provide data on the distribution of these diseases, few studies have addressed the factors that may drive the distributions. Anthropogenic nutrient
loading, increased sedimentation, and global climate change (including increased temperature) have been suggested in the literature as potential causes of the increase in coral disease incidence. This research specifically
addressed 11 environmental factors (total nitrogen, nitrite, nitrate, ammonia, soluble phosphorus, total phosphorus, turbidity, salinity, depth, temperature, and sedimentation) and 4 coral community composition factors
(species richness, species diversity, colony size, and density [colonies/ m2])
associated with coral disease incidence and prevalence on reefs of the
northern Florida Keys Reef Tract and reefs near Lee Stocking Island in the
Bahamas Exuma Chain. Intra- and inter-regional comparisons were made
between transects containing infected individuals and transects with no
signs of disease. The Florida Keys exhibited greater disease prevalence
coupled with higher levels of total nitrogen, nitrate, nitrite, and salinity.
Lower disease prevalence, higher phosphorous levels, and greater species
richness, diversity, and density were observed in the Exumas. Intra-regionally, temperature, sedimentation, density, and colony size differed between
diseased and healthy sites. This is the first study to provide direct regional
comparisons of coral disease along with the environmental factors and
community composition associated with infections.
WAGNER, C. MICHAEL* and GARY D. GROSSMAN. Warnell School
of Forest Resources, University of Georgia, Athens, GA, USA. Conditionspecific asymmetric competition between threatened native and aggressive non-native minnows may mitigate an ongoing invasion.
Approximately 13 years ago the yellowfin shiner (Notropis lutipinnis) was
introduced into the headwaters of the Little Tennessee River of western
North Carolina, USA where it exhibits microhabitat preferences that overlap significantly with those of a threatened native minnow, the smokey
dace (Clinostomus funduloides ssp.). Previous field observations have established that each of these drift-feeding minnows routinely forage in both
single- and mixed-species groups, and employ aggression to maintain the
forward positions in profitable habitat patches. Laboratory experiments
have further established that the introduced shiner is: 1) approximately
twice as aggressive as the native dace in intraspecific foraging groups; and,
2) a less efficient forager at higher stream velocities than the native dace.
We conducted a response-surface competition experiment to examine the
effects of water velocity (10 or 20 cm s-1) and group size (4 or 8 individuals) on intra- and interspecific aggression and foraging performance in
these species (six treatments: 4 dace, 4 shiner, 8 dace, 8 shiner, 2 dace +
2 shiner, 4 dace + 4 shiner). Each group was tested at both velocities on
subsequent days. When competing with native dace, the more aggressive

346

Abstracts

shiner occupied significantly higher ranked positions at both velocities and

group sizes. However, shiner only successfully monopolized the available
prey at low velocities. At high velocity clear competitive release occurred
for the native dace as: 1) in small groups dace acquired the largest proportion of the available prey (i.e., competitive reversal); and, 2) in large
groups dace received an equal proportion of the available prey. Such asymmetric condition-specific competition implies maintenance of habitat diversity (high and low velocity patches) may provide competitive refugia
for the smokey dace.
WAHL, EUGENE R.* and CASPAR AMMANN. National Center for Atmospheric Research, Boulder, CO, USA. Using AOGCM output to assess
the performance of teleconnection patterns derived from climate proxies.
Long continuous runs of coupled atmosphere/ocean general circulation
models (AOGCMs) are now being developed using the full suite of forcings that are particularly relevant over the last millenium: greenhouse gases,
tropospheric sulfates, variations in solar output, and volcanic aerosols. Output from these models is reported as monthly averages, allowing a number
of new avenues to compare the modeled values with high-resolution paleoclimatic reconstructions and to use the physically consistent model
world to examine issues in paleoclimatic analyses. In pursuit of the second
goal, output from a variety of long transient model runs is being studied
to analyze the fidelity of proxy-based measures of the teleconnection patterns of important global climate modes. Initial work has focused on
ENSO/drought teleconnections in six runs from two models at the U.S.
National Center for Atmospheric Research (NCAR). In these runs, the teleconnection patterns associated with "real" (within-model) Nino3 SST variations are compared to those produced by a within-model "proxy", December-March precipitation in Ecuador/NW Peru. Both "real" and "proxy"
ENSO measures are stratified into high/low 25 percentile composites, representing warm/cool events respectively. Results indicate that the models
reproduce known ENSO/moisture teleconnection patterns during the period
of instrumental data (c.1870-present) quite well, indicating they provide a
reasonable basis for judging the performance of the simulated proxy patterns. The teleconnection patterns produced by the simulated proxy show
generally good fidelity to the "real" ones at the global scale; they are generally somewhat weaker in intensity and suggest regions where proxy performance is less good in either one or both of the ENSO phases, e.g.
western North America. Analysis using running-average low pass filters
indicates that the simulated proxy patterns may be more sensitive to temporal smoothing than the "real" patterns, amplifying the need to establish
maximal dating accuracy when developing and using proxy-derived climate
reconstructions.
WALBRIDGE, MARK R.,1,* BARBARA CADE-MENUN2 and WILLIAM J. DRESS.1 1 West Virginia University, Morgantown, WV; 2 Stanford
University, Stanford, CA. Landscape analysis of soil characteristics of
12 forested floodplains in the southeastern US.
We compared soil characteristics of forested floodplains selected as part of
a broader study to analyze phosphorus transformations during flooding
events. Soils were collected from 12 floodplains bordering two types of
riverine systems (alluvial vs. blackwater), with underlying parent materials
of two ages (Young - Pliocene/Miocene - 2-24 millions years old vs. Old
- Cretaceous - 66-144 million years old) (n 5 3 per river type/parent material combination). Surface soils were collected form all 12 sites during
July and August 2002, and analyzed for pH, texture, total C, N, and P, and
base cations. Soils were also extracted with EDTA/NaOH, and then analyzed by 31-P NMR. Soil pH averaged 4.7 across all sites. Total C, N, and
P concentrations were all significantly greater in soils collected from blackwater vs. alluvial floodplains. Soil N:P ratios ranged from ,1 to .13, and
were greater in blackwater systems on old parent materials. A significantly
greater proportion of P was present as organic P in soils from blackwater
vs. alluvial floodplains (59.2 vs. 40.9 %, respectively). Phosphate monoesters were the predominant from of organic P in all soils, comprising 6589% of total organic P, followed by phosphate diesters (11-31 % of total
organic P). Small percentages of pyrophosphates (1-4 % of total P) were
found in all soils. Small percentages of phosphonates were found primarily
in soils of blackwater floodplains. Results emphasize the importance of

considering landscape features (e.g., river type; parent material) when

studying P transformations in field experiments.
WALDROP, THOMAS A.1,* and JAMES MCIVER.2 1 USDA Forest Service, Southern Research Station, Clemson, South Carolina; 2 USDA Forest
Service, Pacific Northwest Research Station, Le Grande, Oregon. The national fire and fire surrogate study: an interdisciplinary comparison of
methods to reduce fuels and fire risk.
Forest ecosystems throughout the United States are denser today and have
heavier fuel loads than historic ecosystems that were maintained by frequent fires. The result has been an increase of catastrophic wildfires and a
loss of ecosystem integrity. The need to reduce live and dead fuels is widely
accepted and silvicultural treatments, such as prescribed fire and thinning,
are readily available to accomplish this objective. These treatments can be
used to mimic the effects of historic fire regimes on forest structure but
virtually no comparative data exist on how they mimic the ecological functions of fire. The National Fire and Fire Surrogate Study uses an interdisciplinary approach to compare ecological and economic impacts of fuelreduction treatments in ecosystems throughout the United States. Study
sites were selected at 13 locations in ecosystems as diverse as western
Ponderosa pine, southern longleaf and slash pines, southeastern pine-hardwood mixtures, and northeastern oak-hickory forests. Strict treatment and
sampling protocols were followed at all sites. Treatments include prescribed
burning alone, mechanical fuel reduction alone, burning and mechanical
treatment together, and an untreated control. Each was replicated at least
three times at all 13 locations. Core variables were measured from disciplines including wildlife (mammals, birds, and herpetofauna), soils (fertility, nutrient cycling, microfauna, and decomposition), entomology, pathology, vegetation, fuels, fire behavior, and economics. Treatments and variables of local interest were added at some locations. Early results suggest
that each treatment creates a unique ecosystem structure which has varying
impacts on most ecosystem functions. A compilation of results from all 13
sites will provide a better understanding of how choices for fuel management impact multiple components of ecosystems.
WALKER, AMBER M.* and CHARLES H. NILON. University of Missouri-Columbia, Columbia, MO. Land use, land cover and socioeconomic variables as predictors of bird species composition in St. Louis, MO.
Using a patch dynamics to approach to understand patterns of bird species
composition and abundance in cities requires information on patch habitat
features and on the characteristics of the matrix that surrounds the patch.
Azerrad and Nilon (2001) studied bird species composition and abundance
in 55 greenspace patches in inner city St. Louis, Missouri, finding that four
patch habitat variables: percent canopy closure, canopy height, shrub stem
density, and percent forb cover were the best predictors of bird species
composition. Our objectives in this study are to describe the matrix surrounding these 55 habitat patches at three different scales (200m, 500m,
and 1000m from patch center) and identify matrix variables that are predictors of differences among patches in bird species composition. Bird
species composition and abundance for each patch were determined by
Azerrad (2001) in 1998 and 1999. We are using land cover, land use,
landscape pattern and socioeconomic data to characterize the matrix surrounding each patch.
WALKER, MARILYN, * AMY CARROLL, CARL H. WAHREN and
ANDREW BORNER.1,3 1 Boreal Ecology Cooperative Research Unit, University of Alaska Fairbanks, Fairbanks, AK; 2 USDA Forest Service PNW
Station, Fairbanks, AK; 3 Dept. of Biology and Wildflie, University of Alaska Fairbanks, Fairbanks, AK. Phenological responses to changing climate in arctic plant communities.
1,2,

1,3

Phenology is a critical process in arctic plants adapted to short growing

seasons. Most arctic species are highly sensitive to timing of snowmelt in
particular, and changes in timing of events in plant species may be of
extreme importance to certain animals. We have been collecting data on
the phenology and growth of arctic plant species under conditions of experimental manipulated winter snow cover and summer temperature for 8
continuous years. We use path analysis to examine a system of closelyrelated variables and to look at the sensitivity of species to this environmental change.

WALLACE, LINDA L.,1,* REBECCA A. SHERRY,1 YIQI LUO,1 NANCY

ZEHRBACH1 and SHIQIANG WAN.2 1 University of Oklahoma, Norman,
OK; 2 Oak Ridge National Laboratory, Oak Ridge, TN. Phenological response of tallgrass prairie to experimental warming and clipping.
We have warmed 6 plots of tallgrass prairie 2o C above ambient since 1999
in a fully crossed design with both annual clipping (hay mowing mimic)
and warming. Although we have not yet seen substantial community structure changes, the phenology of plant species has been altered, particularly
in the fall. This site has heavy clay soils and a depauperate winter annual
flora. Hence, few early spring differences in phenology have been noted
between warmed and unwarmed plots, despite soil temperatures above 10o
C occurring from 6 to 22 days earlier in the warmed plots. Perhaps the
species present in these plots are more sensitive to daylength than soil
temperature. However, in the fall we have found that green tissues are
present for longer periods on Schizachyrium scoparium and Sorghastrum
nutans, both C4 species. Ambrosia psilostachyia, a C3 forb, also maintains
green tissues and flowering structures longer in the warmed plots for time
periods up to 14 days. Soil temperatures above 10o C have been maintained
in the warmed plots from 1 to 11 days longer than in the unheated plots.
In a site with a more diverse winter annual flora, we would expect larger
phenological differences to occur in the early spring, as well. We have not
yet seen phenological changes that would suggest uncoupling of plant/
pollinator systems.
WALLENSTEIN, MATTHEW D.,1,2,* WILLIAM H. SCHLESINGER,2
RHEE S.K.3 and JIZHONG ZHOU.3 1 Duke University, University Program
in Ecology, Durham, NC; 2 Duke University, Nicholas School of the Environment, Durham, NC; 3 Oak Ridge National Laboratory, Environmental
Sciences Division, Oak Ridge, TN. Nitrogen fertilization alters fungal
and bacterial biomass and microbial community structure in three
long-term experiments.
We collected soil samples throughout the 2002 growing season in three
long-term nitrogen fertilization experiments in New England, USA (Harvard Forest, Bear Brook Watershed, Mt. Ascutney). Fungal and bacterial
biomass were assessed using the chloroform fumigation-extraction technique and by substrate induced respiration coupled with selective inhibitors.
At two of the sites, we found a large and significant decrease in total
biomass with increasing nitrogen inputs. The ratio of fungal:bacterial biomass also decreased with increasing N. Across all 3 sites, there was a
negative correlation between microbial biomass and annual net N mineralization, and a positive correlation with pH. We also used functional gene
microarrays to assess long-term changes in the functional diversity and
gene abundance in DNA extracted from these soils. These microarrays
provide abundance data for over 5000 genes important in nitrogen, carbon,
phosphorus, and sulfur cycling. These analyses revealed consistent shifts
in the diversity and abundance of several nitrogen cycling genes with fertilization treatments. These results suggest that long-term changes in nitrogen inputs to forest soils result in fundamental changes in microbial community structure that are likely to alter nitrogen cycling.
WALLER, DEBORAH A.* and JOHN MARCH. Old Dominion University, Norfolk, Virginia, USA. Ant species diversity in a mainland coastal
ecosystem and nearby barrier island.
This study examined ant species diversity related to habitat in the Virginia
Coast Reserve. Ants were collected in different habitats along the coast
near Nassawadox, Virginia, and on nearby Parramore Island. Samples were
taken from tidal forest, marshland and agricultural areas on the mainland
and from forest and beach on the island. Ant baits and pitfall traps were
placed along transects in each habitat; baits were examined for ants after
an hour and pitfall trap contents were collected the following day. Litter
samples were taken near pitfall traps and returned to the laboratory for ant
extraction over a week using Berlese funnels. Searches of vegetation and
fallen logs were also conducted in each habitat. Ant species were most
diverse in the mainland tidal forest and least diverse on the island beach.
Some ant species were restricted to either the mainland or the island, while
other species were widespread.

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347

WALSH, PETER D.,1,* PAUWEL WACHTER,2 ELLA AKOU MARC,2

BAS HUIJBREGTS,2 DANIEL IDIATA MAMBOUNGA,3 SALLY A.
LAHM,4 SOSTHE`NE NDONG OBIANG3 and LEE J.T. WHITE.5 1 Department of Ecology and Evolutionary Biology, Guyot Hall, Princeton, NJ,
USA; 2 WWF Central Africa Regional Program Office, Libreville, Estuaire,
Gabon; 3 Ministe`re de lEconomie Forestie`re, des Eaux, de la Peche charge
de lEnvironnement et de la Protection de la Nature, Direction de la Faune
et de la Chasse, Libreville, Estuaire, Gabon; 4 Institut de Recherche en
Ecologie Tropicale, Libreville, Estuaire, Gabon; 5 Wildlife Conservation
Society, Bronx, NY, USA. Group structure, population density and the
emergence of Ebola in African Apes.
The great majority of the worlds gorillas (Gorilla gorilla) and common
chimpanzees (Pan troglodytes) live in Western Equatorial Africa (WEA).
Over the last decade, an epidemic of what appears to be Ebola hemorrhagic
fever has killed a substantial proportion of the gorillas and chimpanzees in
WEA. The epidemic continues to burn and the rest of the regional population is at serious risk. Unfortunately, there is very little published information on Ebola transmission dynamics with which to plan management
responses. Here we use data on gorilla derived from observations at clearings and extensive nest surveys to test alternative models of Ebola transmission. Our analyses suggest a high rate of virus transmission between
gorillas within a group. Results from a spatially explicit, individually based
simulation model suggest density dependent transmission dynamics in
which Ebola cannot persist below some threshold gorilla population density. Observations on the location of Ebola outbreaks are consistent with
this hypothesis as are survey data. Gorilla group size may also have an
effect on transmission rate that is independent of local population density.
We discuss maximum likelihood estimation of simulation model parameters
with both observations from habituated gorillas and nest survey data and
describe how the model might be used to evaluate and optimize Ebola
management strategies such as vaccination, barrier cutting, and translocation.
WALTER, ACHIM,1,* MAJA M. CHRIST,1,2 GREG BARRON-GAFFORD1 and RAMESH MURTHY.1 1 Biosphere 2 Center, Columbia University, Oracle, AZ, USA; 2 Forschungszentrum Julich, ICG III (Phytosphare), Julich, Germany. Diurnal leaf growth variations in Populus deltoides grown at different external CO2 -concentrations.
Cottonwood trees (Populus deltoides Bartr.) were grown in three different
CO2-concentrations (40, 80 and 120 Pa) within the environmentally controlled Intensive Forestry Mesocosm at the Biosphere 2 Center, Oracle, AZ.
Diurnal leaf growth characteristics of trees within three mesocosms were
monitored both with a conventional and a digital image sequence processing method for several months. The novel imaging method was performed
for the first time under relatively realistic environmental conditions, as
opposed to a growth chamber. The results show that diurnal courses develop gradually throughout the vegetation period and that their amplitude
depends on ambient CO2-concentration. Decreased amplitudes were found
in elevated CO2. Phasing of the diurnal courses of different leaves was
identical, irrespective of treatment and positions along the architecture of
the plant. The distribution of growth within the plant, however, was affected
by the CO2 treatment. Results are discussed on the background of environmental constraints, the photosynthetic performance and the overall carbon
balance of the different treatments.
WAN, SHIQIANG,1,* KURT PREGITZER,2 ELIZABETH ONEILL1 and
RICHARD NORBY.1 1 Environmental Sciences Division, Oak Ridge National Laboratory, Bldg.1059, P. O. Box 2008 (Fedex address: Bethel Valley
Road), Oak Ridge, TN; 2 School of Forestry and Wood Products, Michigan
Technological University, Houghton, MI. Responses of roots of two deciduous tree species, Acer rubrum (red maple) and A. saccharum (sugar
maple), to elevated CO2 and temperature.
As a central component of ecosystem carbon and nutrient cycling, root
production and turnover may be substantially affected by global change
and act as a key link between plant responses and long-term changes in
soil organic matter and ecosystem carbon balance. A four-year temperature
and CO2 interaction experiment was conducted in open-top chambers at
the Oak Ridge National Environmental Research Park in Tennessee, USA.

348

Abstracts

Both elevated CO2 and elevated temperature significantly enhanced total

root production, root mortality, and net root production. At the end of the
experiment, elevated CO2 also increased fine-root biomass. However, higher temperature reduced fine-root biomass for all size classes, which could
be attributable to the warming-induced heat stress and reduction in aboveground growth in two hot summers. Specific root length significantly decreased with elevated CO2 and the interaction of temperature and CO2, but
temperature alone did not affect it. Elevated temperature and the interaction
of temperature and CO2 had significant positive effects on N concentration
of roots smaller than 0.5 mm whereas elevated CO2 had a negative effect.
Root C concentration increased with root size class, but root N concentration decreased. Both elevated CO2 and temperature reduced the ratio of
root biomass to aboveground biomass, suggesting that there might be a
relative decrease in belowground C allocation in these deciduous trees in
a warmer, CO2 enriched world. The interactive effects of elevated temperature, CO2, and interannual variability of climate in regulating root production, turnover, and biomass may all be important in understanding forest
C cycling.
WANG, LIXIN,1 PAUL P. MOU1 and JONES H. ROBERT.2 1 University
of North Carolina at Greensboro, l_wang5@uncg.edu, Greensboro, North
Carolina, U.S.; 2 Virginia Polytechnic Institute and State University,
Blacksburg, Virginia, U.S. Physiological and morphological plasticity of
plants nutrient foraging.
Plant root nutrient foraging ability in heterogeneous soil is enhanced by
morphological, physiological, and demographical plasticity. We conducted
a greenhouse experiment to examine root physiological and morphological
plasticity of three plant species (Erechtites hieracifolia (L.) Raf, Pinus taeda L. and Liquidambar styraciflua L). Seeds were sown in 19-liter pots
that were randomly assigned into two treatments: even fertilization or quarter fertilization. Each pot received 5 grams of slow-release fertilizer (N-PK) after the seedlings were established (one plant per pot). In the even
fertilization treatment (H), fertilizer was evenly applied to the surface of
each pot, while in the quarter fertilization treatment (Q), all fertilizer was
applied to one randomly selected quarter. As the plants reached adequate
size (roots touching the pot edge), we injected 20 ml 5 atom % 15 N ammonium-nitrate solution (0.1 M or 0.29 M with red food dye) into a soil
column of each pot. This procedure was performed to fertilized or unfertilized quarters in the Q treatment, or one randomly selected quarter in H
treatment. This injection was repeated 24-hrs later at the same location.
Plants were harvested 24-hrs after the second injection. The 15 N fertilized
plug was collected first, then, aboveground plant parts and roots from each
quarter were harvested. Fine roots from each plug were sorted, oven dried,
weighed and prepared for 15 N analyses. The preliminary results demonstrated that root physiological plasticity for nutrient uptake differed significantly (p50.05). Nutrient uptake rate by roots grown in enriched soil was
the highest, followed by the roots grown in evenly fertilized pots. The root
nutrient uptake rate in unfertilized quarters was the lowest. Root morphological plasticity of these species was significantly different (p,0.001), and
the differences among species were significant. This finding agreed with
our previous study results.
WANG, XIANLI* and BO SONG. Clemson University, Georgetown, SC,
US. Forest dynamics visualization using models.
Visualization of forest dynamics using simulation models can aid in comparing various management alternatives. In this study, the PnET-II model
was used in determining ecosystem biomass. Combining the stand information with the modeling output, the dynamics of the forest were visualized. Using the PnET-II model, we calculated aboveground biomass and
stand age, which were then used to estimate tree height and density for 3D visualization. This study showed that 3-D visualization can realistically
generate virtual forest whereby the researchers and the public can use different modeling assumptions to look at the various management alternatives.
WANG, XIANZHONG1 and KEVIN L. GRIFFIN.2 1 Indiana UniversityPurdue University Indianapolis, Indianapolis, Indiana; 2 Lamont-Doherty
Earth Observatory, Columbia University, Palisades, New York. Carbon
assimilation in wild type and starchless mutant of Nicotiana as affected
by CO2 concentration.
Research based on the use of genetically manipulated plants can improve
our understanding of the regulation of carbon metabolism. We examined

the relationship between carbon assimilation physiology and carbohydrate

contents in leaves of wild type and a starchless mutant (NS 458) of Nicotiana sylvestris (Speg et Comes) grown at ambient (365 mmol mol-1) or
elevated (730 mmol mol-1) CO2 concentration. Elevated CO2 increased net
photosynthesis (A) in wild type plants by 27%, but had no effect on A in
starchless mutant plants. Leaf dark respiration (Rd) was not significantly
affected by CO2 level in wild type or mutant plants. There was a positive
correlation between A and Rd in both wild type and mutant plants, regardless of CO2 concentrations. However, the coefficient of determination
(r2) was greater for elevated than for ambient CO2 grown plants. Elevated
CO2 increased contents of soluble sugars and total non-structural carbon
(TNC) of both wild type and mutants, but the magnitude of increase was
much greater for mutants than for wild type plants. Total soluble sugar
content (glucose, fructose and sucrose) and TNC more than doubled at
elevated CO2 for mutant plants, while their increases in response to CO2
enrichment was 87% and 90%, respectively, for wild types. Although leaf
Rd of the starchless mutant had a statistically positive correlation with
glucose, total soluble sugar and TNC under elevated CO2, the percentage
of variation in leaf Rd attributable to carbohydrate content was low. Leaf
Rd of wild type plants was uncorrelated with carbohydrate contents at
ambient or elevated CO2. Our study demonstrated that leaf Rd tends to
increase with leaf A, especially at a higher CO2 concentration, and that leaf
carbohydrate content is not the most significant determinant of leaf Rd in
N. sylvestris.
WANG, YUNG-HO (OPHELIA)* and CAROL K. AUGSPURGER. University of Illinois at Urbana-Champaign, Urbana, IL 61801, USA. Understory palms and cyclanths impose strong recruitment limitation on
seedlings in a Neotropical rain forest.
Recruitment limitation, which occurs when species either fail to reach or
establish in sites suitable for regeneration, has been proposed as an important factor that controls population dynamics and species coexistence in
tropical moist forests. Palms in some Neotropical forests dominate understory strata, and may cause seedling recruitment limitation because their
umbrella-like architecture may prevent seed rain and cast deep shade. This
study surveyed seedlings under dwarf palms/cyclanths in four habitats at
La Selva Biological Station, Costa Rica. It compared seedling recruitment
in 2001 and 2002 at control sites vs. sites under dwarf palms of two heights
and two densities. Overall, dwarf palms/cyclanths had great density (2540
individuals/ha) and covered 20% of the understory in the primary forest.
Overall, seedling growth, survivorship, density, and species richness were
23%, 37%, 34%, and 26%, respectively, lower at under-palm sites than at
control sites. Each of these variables was greatest at control sites and lowest
at high-density palm sites. Seedlings at different stages and of different
taxonomic groups responded to the suppression differently. The main restriction by dwarf palms occurred on post-cotyledonous and woody seedlings. Percent canopy openness, leaf area index (LAI), and photon flux
density (PFD) measured and estimated with hemispheric photos each indicated that light availability was greatest at control sites and lowest at
high-density palm sites. However, overall seedling responses under a given
palm were only weakly correlated with its light environment. The study
revealed significant recruitment limitation under individual dwarf palms,
with greatest negative effects where a palm was surrounded by nearby
palms. This restriction by individual dwarf palms accumulates to a significant level at the landscape scale as well, based on the great dwarf palm
density and coverage in the four habitats.
WARD, BESS B.1,* and GEORGE A. JACKSON.2 1 Princeton University,
Princeton, NJ; 2 Texas A&M University, College Station, TX. Microbial
diversity in functional guilds: Does diversity matter in ecosystem function?
Complex biogeochemical cycles, such as the microbially mediated nitrogen
cycle, are deceptively simple when viewed in terms of the net chemical
transformations they include. For example, the oxidation of ammonium to
nitrite or the denitrification of nitrate to N2, can be determined from the
net nitrogen fluxes measured using geochemical methods. Every biologically mediated process in a particular environment can be ascribed to the
activity of an enzyme, encoded by a functional gene (e.g., ammonia monooxygenase, amoA, or nitrite reductase, nirS). Sequence analysis of such

genes from the environment reveals a vast diversity within functional

guilds; many different variants or alleles of the same functional genes are
seen to be associated with each biogeochemical transformation. We are
investigating the extent to which this genetic diversity is important in determining or regulating the overall rates of biogeochemical processes. In
order to compare diversity/function relationships among sites and times, it
is necessary to assess the degree to which the total diversity of the environment has been sampled. The results of our comprehensive diversity
analysis of genes involved in nitrification and denitrification have been used
to assess the shape of the species abundance curve for functional genes.
While the range of sequence diversity varies significantly between genes
(e.g., amoA vs. nirS), their abundances appear to be lognormally distributed
and should allow us to assess the main features of the community composition in relation to environmental variables and function. The initial
development of DNA microarrays used to evaluate the microbial assemblage along a gradient of measured biogeochemical transformation rates
within Chesapeake Bay will be presented.
WARDROP, DENICE H.,* ROBERT P. BROOKS, WAYNE L. MYERS,
G. P. PATIL and CHARLES TAILLIE. Penn State University, University
Park, PA, USA. Modeling ecological condition in upstream watersheds
and downstream estuaries in the Atlantic Slope region.
The Atlantic Slope Consortium (ASC) is a project designed to develop and
test a set of indicators in coastal systems that are ecologically appropriate,
economically reasonable, and relevant to society. It extends the area relevant to coastal condition into freshwater aquatic ecosystems in the headwater regions of the Appalachian Mountains, and the project area covers
the Delaware, Chesapeake, and Albemarle-Pamlico drainage basins. The
project utilizes ecological and socioeconomic information compiled at the
scale of estuarine segments and small watersheds. The research mandate
of the ASC project is the following: Using a universe of watersheds and
estuarine segments, covering a range of social choices, we ask two questions: * How "good" can the environment be, given those social choices?
* What is the intellectual model of condition within those choices, i.e.,
what are the causes of condition and what are the steps for improvement?
As a basis for compiling ecological indicators, a watershed/estuarine segment classification system was developed representing the range of social
choices. These watershed/estuarine segment classes were the basis for an
analysis of the explanatory power of various indicators through the following steps: 1. Identification of critical areas of low biological integrity on
the watersheds across the project area. This was accomplished by use of
the scan statistic originally developed for geographical surveillance of disease, and adapted to regional environmental and ecological settings. 2.
Reassessment of critical areas taking into account potential explanatory
factors. Three categories of factors (or indicators) were evaluated: descriptive (e.g., land use and landscape), structural (e.g., drainage pattern, geomorphology), and network-related. 3. Modeling of those critical areas
which were not satisfactorily accounted for in the preceding step by general
landscape and drainage variables This exercise provides an initial screening
of ecological indicators with regard to their explanatory power at various
spatial scales, utilizing steps 1 and 2 above. The utility of various indicators
for each of the watershed/estuarine segment categories is reported.
WARREN, PAIGE S.,1,* ANN P. KINZIG,2 CHARLES H. NILON,3 MORGAN GROVE,4 MARY COX5 and CHRIS MARTIN.2 1 Virginia Tech,
Blacksburg, VA; 2 Arizona State University, Tempe, AZ; 3 University of
Missouri at Columbia, Columbia, MO; 4 USDA Forest Service, Burlington,
VT; 5 Parks and People Foundation, Baltimore, MD. Human socioeconomic factors and avian diversity: A cross-site comparison.
A variety of measures have been advanced as predictors of biodiversity in
urban areas, such as human population density, building density, and canopy cover. These measures, either singly or in combination, describe only
a portion of the habitat structure that is important for wildlife. For example,
neighborhoods with the same housing density can be landscaped with different kinds of plants. In our studies of small, residential parks in Phoenix,
Arizona, we found that the socioeconomic status (SES) of the neighborhoods around the parks was one of the best predictors of bird community
structure within them. Human behaviors, values, and resource consumption
levels, which may vary by SES, can influence factors such as the habitat

Abstracts

349

and food availability for other organisms. An advantage of using SES factors over direct measures of the ecological factors influencing biotic communities is that information on SES is widely available. We tested whether
human SES showed a similar correlation with avian species richness in
parks in two cities: Phoenix and Baltimore, Maryland. We used PRIZM
market cluster data to classify the SES of the neighborhoods surrounding
the parks. Since PRIZM use the census block group as its geographical
unit, we treated the set of block groups immediately surrounding each park
as its neighborhood, selecting only parks found in relatively homogeneous
neighborhoods with respect to PRIZM market clusters. We found that bird
species richness is indeed correlated with SES in both cities, but SES explains much higher proportions of the variance in Phoenix. Birds in Baltimore appear to be less sensitive to differences associated with human
SES and more sensitive to overall human density than birds in Phoenix. In
addition, bird diversity in Baltimore parks also appears to be more strongly
area-dependent than in Phoenix. We propose several possible reasons for
these differences between the two cities.
WARREN, ROBERT J.,1,* IRENE ROSSELL2 and KEVIN MOORHEAD.2
1
Western Carolina University, Cullowhee, NC; 2 University of North Carolina at Asheville, Asheville, NC. Colonization and Establishment of
Red Maple (Acer Rubrum) in a Southern Appalachian Wetland.
Following a disturbance, there may be a recruitment window when conditions are optimal for the colonization and establishment of woody species.
Research in terrestrial habitats has indicated that such a window for Acer
rubrum establishment lasts approximately 1-14 years following a disturbance. The objective of this study was to investigate A. rubrum colonization
and establishment in a southern Appalachian wetland by examining (1) the
recruitment window for seedling colonization and (2) size and age dynamics in established stands. The study site was the Tulula Creek wetland
complex, which includes both forested (dominated by A. rubrum) and early
successional fen and floodplain habitats. The DBH and core age of understory and overstory trees were measured in 108 plots in three areas of the
Tulula Creek wetland complex in 1994 and 2001 as part of a larger ecological study. The three areas were a cleared fen (open fen), forested fen
(closed fen) and a cleared floodplain (open floodplain). A fourth site, a
forested floodplain (closed floodplain), was sampled in 2001. Heights of
A. rubrum seedlings were measured in 379 quadrats in the open floodplain
in 1996 and 2001. Diameter and height distributions were compared with
established distribution models in order to determine A. rubrum stand dynamics at the four sites. Results showed that recruitment continued (at a
diminishing rate) at sites that were last cleared 7 years (open floodplain),
14 years (open fen) and approximately 30 years (closed fen) earlier. Seedling recruitment ceased after approximately 45 years of tree growth (closed
floodplain). These results suggest that A. rubrum readily colonizes and
becomes established in wetland habitats, and that the opportunity for its
establishment may last longer in wetlands than in terrestrial systems.
WATANABE, MORIYOSHI,* MASAHIKO SEKINE, MASAO UKITA
and TSUYOSHI IMAI. Yamaguchi University, Ube, Yamaguchi, JAPAN.
Monitoring the ecosystem condition of the shallow sea by using the
sound of snapping shrimp.
Although detecting the impacts of pollution on sea ecosystem is very important, measuring the condition of living organisms in the sea requires
some skill and patience. Snapping shrimp (Genus Alpheus), which are bottom-living animals, are distributed all over the world coastal area and yet
have not suffered from fishing pressure. The rapid closure of the chela
emits a powerful pulse sound accompanied by water jet. An idea came out:
doesnt this sound represent the number of snapping shrimps (and hopefully
the ecosystem condition, too) on the bottom of the sea? The objective of
this research is to establish a monitoring method of the ecosystem condition
in the shallow sea by monitoring the snapping shrimps sound. The pulse
count (the number of snap per minute) measuring method is quite simple.
The pulse count can be achieved merely by recording their sounds for a
few minutes without special biological knowledge by using a hydrophone
and a PC with the software we developed. From field surveys conducted
since 1996, the following findings have been obtained; (1) A few minutes
measurement of pulse count is sufficient to acquire the pulse count value
that represents the study period. (2) The pulse count changes according to

350

Abstracts

water temperature under the normal condition. The shrimp population can
be estimated from the pulse count and the water temperature. (3) The pulse
count seems to decrease in the place where the water quality is not very
good, like the head of Tokyo Bay or the Bosporus. (4) The pulse count
sharply falls by the occurrence of hypoxia and red tide. This method has
shown premise to be a new, simple and effective technique of monitoring
the ecosystem health in coastal areas.
WATKINS, JAMES E.,1,* CATHERINE CARDELUS1 and ROBBIN
MORAN.2 1 University of Florida, Department of Botany, Gainesville, FL,
USA; 2 New York Botanical Garden, Bronx, NY, USA. Diversity and
abundance of epiphytic (canopy and understory) and terrestrial ferns
along an elevational gradient: Implications for dispersal and migration.
Along a continuous wet forest elevational gradient in Costa Rica, 303000m, we examined the distribution and diversity of pteridophytes, both
epiphytic and terrestrial, at each of six sites: 2600m, 2000m, 1600m,
1000m, 500m and 30m. To determine terrestrial and understory (low trunk)
epiphytic species diversity and abundance, 4: 100m x 4m transects were
laid out in randomly chosen areas at each site. Transects were subdivided
into 5m x 4m quadrats in which both diversity and abundance were recorded. To determine diversity and incidence of canopy species, we
climbed ten individual canopy trees per site except at 2600m where 5
individuals were studied. Two: 4m by 0.5m canopy transects, subdivided
into 0.5m x 0.5m quadrats, were laid out and all ferns found in these
quadrats were sampled and their presence recorded. Thus far, a total of 276
species of pteridophytes in 61 genera were recorded along the entire gradient: 114 canopy epiphytes, 122 understory epiphytes and 79 terrestrial
species. Pteridophyte diversity shows a clear mid-altitudinal bulge at
1000m for rarefied data. Predictably, overlap of species among elevations
decreases with increasing distance between sites. Overlap among habitats
is higher between the canopy and understory epiphyte communities (18%)
while overlap of both epiphytic groups with terrestrial species is 0 and 1
species respectively. The differences in overlap suggest extreme recruitment limitation on local and regional scales in a dispersal unlimited system.
WATTS, SEAN* and O. J. REICHMAN. University of California, Santa
Barbara, Santa Barbara, CA, USA. The evolutionary response of plants
to root herbivory.
Most work conducted on plant-herbivore interactions has focused on
aboveground herbivory. Plants, however, suffer greater negative consequences from belowground herbivory. We investigated the capacity for
plants to evolve in response to root herbivory by comparing populations
of two grassland forbs (Eschscholzia californica and Hemizonia fasciculate) on the California Channel Islands, which have no record of gophers,
and mainland areas with high densities of gophers. We used this system to
test the hypothesis that, due to the absence of root herbivory, island plant
species have a reduced capacity to both deter and tolerate root loss. Island
and mainland plants were collected to determine if island plant forms contain less defensive compounds than mainland conspecifics. In addition, we
censused island and mainland populations to determine if island plant forms
exhibit lower fecundity and greater mortality in response to simulated belowground herbivory. Results from the 2002 field season indicate that the
roots of mainland Eschscholzia californica are defended by a greater
amount and greater variety of alkaloid compounds. Censuses conducted
after simulated root herbivory showed greater mortality among island Hemizonia fasciculat and a delayed recovery in island Eschscholzia californica
which translated to lower total annual reproduction relative to mainland
populations. In addition to characterizing the evolutionary responses of
island and mainland plants to root herbivory, this set of experiments should
contribute to a comprehensive, understanding of the ecological tradeoffs
between growth, defense and reproduction in plants.
WAYSON, CRAIG,* JC RANDOLPH and HANS PETER SCHMID. Indiana University, Bloomington, Indiana. Comparison of methods to estimate annual soil respiration in a mid-latitude deciduous forest.
The single largest component in the calculation of net CO2 flux from midlatitude deciduous forests is the flux from forest floor and soil respiration;
however, accurate measurements of this flux are difficult. Since May, 1998

at the AmeriFlux site in Morgan-Monroe State Forest (MMSF), located in

south-central Indiana, we have made periodic (usually monthly) measurements of soil respiration using portable instruments in plots with differing
topography. These data allow spatial comparison but are limited temporally.
However, since September, 2001 soil respiration has been measured continuously using an automated chamber system. The data are limited to a
single location near the flux tower, but the CO2 flux, soil moisture, soil
temperature, air temperature, and precipitation data are continuous. Estimates of seasonal and annual soil respiration rates for 2002 made by direct
measurement using the chamber system are compared with estimates generated by eddy covariance flux measures and estimates made using a simple
model and spatially specific data from the portable system.
WEAR, DAVID N.* USDA Forest Service, Southern Research Station,
Research Triangle Park, NC, USA. The changing face of Southern forest
lands: findings from the Southern Resource Assessment.
The forests of the South are diverse and dynamic. They have been heavily
utilized since European settlement and their current conditions reflect the
legacies of a long land use history. A century of especially intensive agricultural exploitation gave way to a period of forest recovery and growth
at the beginning of the twentieth century. In the last quarter of the century,
however, timber harvesting and in-land development for urban uses has
increased substantially. As a result, questions have emerged regarding the
health, productivity, and ultimately the sustainability of the Souths forests
and the benefits they provide. The Southern Forest Resource Assessment
(SFRA) was initiated in the spring of 1999 to address broad questions
concerning the status, trends and potential future of southern forests. The
SFRA represents an unusual situation where government evaluates the status and future of a largely private sector: 89 percent of the Souths forests
are held by private owners. With this in mind, the SFRA was chartered as
an information gathering exercise. The overall objective of the Assessment
was to develop a thorough and objective description of forest conditions
and trends in the South, compiled in a way that would help the public
understand a complex and dynamic resource situation. In this presentation,
we discuss the major findings of SFRA, focusing especially on the several
factors that are reshaping southern forests and examine the implications for
the sustainability of southern forests and the conduct of research in the
South.
WEARING, HELEN J.,1,* PEJMAN ROHANI,1 TOM C. CAMERON2 and
STEVEN M. SAIT.2 1 University of Georgia, Athens, Georgia; 2 University
of Leeds, Leeds, U.K. Dynamical effects of demographic stochasticity
in stage-structured population models.
Stage-structured population models have often ignored the role of demographic stochasticity when investigating both single and multi-species dynamics. In this talk, we describe a framework that can be used to simulate
stochastic birth, death and maturation processes. This approach also permits
a systematic analysis of the dynamical significance of the variance in the
duration of each stage, assuming a gamma delay distribution. Applying this
to the well-studied Plodia and Plodia-Venturia systems, we demonstrate
that certain dynamical patterns predicted by deterministic delay-differential
equation models are sensitive both to demographic stochasticity and to the
assumed variance of the distributed delays. We discuss how our findings
may explain why specific phenomena (e.g. half-generation cycles) predicted by the deterministic models, despite occurring for a wide range of parameter values, are seldom observed in real systems.
WEATHERLY, HEATHER E.,1,* BO SU,2 LINDA WALLACE2 and JOHN
A. ARNONE.1 1 Desert Research Institute-Reno, Reno, NV, USA; 2 University of Oklahoma, Stillwater, Oklahoma, USA. NSF IRCEB interannual climate variability and ecosystem processes in tallgrass prairie:
Controls on litter decomposition.
The 48C increase in average air temperature that is expected to occur over
the next decade is likely to result in an increase in the decomposition of
plant litter and potentially enhance the rate of nutrient return to the soil.
Evidence for the positive relationship between temperature and decomposition is abundant. This may be especially significant in a tallgrass prairie
ecosystem where moisture is not typically a limiting factor for microbial-

mediated decomposition processes, in contrast to more arid systems. In the

first year of a four-year study to determine the effects of a 48C increase in
air temperature on ecosystem processes, we decomposed representative
mixtures of tallgrass prairie litter in litterbags, as well as standard cotton
and popsicle sticks, for 8 months in laboratory mesocosms (EcoCells) in
order to determine what may control decomposition under current ambient
temperature in the tallgrass prairie ecosystem. One half of the litterbags
were exposed under the intact canopy, while the remaining half were exposed under a mowed layer. We also decomposed the same litter in field
plots in Oklahoma to eliminate possible artifacts of the controlled laboratory conditions. Linear regressions of percent decomposition with aboveground net primary production (both during the year the litter was produced
and during the period of litterbag exposure) showed no relationship. Furthermore, no relationships existed between percent decomposition and species diversity or between percent decomposition and average soil moisture
in the top 15 cm of soil (TDR) over the duration of litterbag exposure.
However, litterbags that were exposed beneath the intact canopy of each
monolith (plot) decomposed more rapidly (35%) than those exposed on the
mowed side (26%) of each monolith (P50.005). Both temperature and
relative humidity (averaged over the exposure period) were higher on the
soil surface of the mowed canopy than on the soil surface beneath the
intact canopy (P,0.000 for both temperature and relative humidity), suggesting that abiotic factors may be less important in controlling decomposition than microbial processes in this system. The possibility that microbial activity is localized in areas that are more nutrient rich (i.e. not
mowed) may explain the faster decomposition observed in the intact canopy.
WEBSTER, THEODORE M.* USDA-ARS, twebster@tifton.usda.gov, Tifton, GA. Polyethylene mulches suppress purple nutsedge (Cyperus rotundus) and yellow nutsedge (Cyperus esculentus) growth.
Purple and yellow nutsedge are the most troublesome weeds of commercial
vegetable crops in the southern region. Methyl bromide has been the pest
management foundation for nearly 50 years. However, methyl bromide is
a suspected ozone-depleting substance and its use will be halted in 2005.
Nutsedges will be among the most difficult of these pests to control. Polyethylene mulches suppress many weeds, however pointed tips of emerging
nutsedge shoots readily pierce polyethylene mulch. The objective of this
study was to evaluate purple and yellow nutsedge growth in systems with
different polyethylene mulches. A single purple or yellow nutsedge tuber
was planted in 58 cm diameter pots filled with field soil. Pots were covered
with 32 mm black polyethylene, 32 mm clear polyethylene, or no-mulch
(six experimental units per replication). Treatments were arranged as a
RCBD with three replications, repeated over time. Relative to no-mulch,
yellow nutsedge shoots that pierced the black and clear polyethylene mulch
were reduced 95% and 99%, respectively. Greater than 91% of yellow
nutsedge shoots that emerged in mulch systems did not pierce the mulches
(were trapped underneath). Purple nutsedge shoots emerged through clear
mulch were reduced by 79% relative to no-mulch. However, black polyethylene reduced purple nutsedge shoots that emerged through the mulch
by only 29%, compared to no-mulch. Most (58% and 80%) emerged purple
nutsedge shoots remained below black and clear polyethylene mulches,
respectively. Successful nutsedge management must target tubers, the primary means of reproduction. In 16-weeks, yellow nutsedge in no-mulch
produced 486 tubers, while tuber production was reduced 51% by both
mulches. Purple nutsedge in no-mulch produced 223 tubers, while black
and clear polyethylene mulches reduced tuber production 34 and 82%,
respectively. Where feasible, polyethylene mulch can be an important component for nutsedge management in a methyl bromide alternative system.
WEICHT, THOMAS R.* and DARYL L. MOORHEAD. University of Toledo, Toledo, Ohio. Impacts of temperature and moisture regimes on
Scottnema lindsayae population dynamics, an Antarctic soil nematode.
An earlier model of Scottenema lindsayae population growth was modified
to include observed sex ratios and response to gravimetric soil moisture
(data provided by the McMurdo Long-Term Ecological Research program).
The sex ratio of field population approximated 63% 6 25 females, which
altered the slope of the relationship between the coefficient of population
growth (l) and cumulative annual degree-days (DD): l50.6 + 0.002DD,

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351

n 5 17, r2 5 0.97. The relationship between soil water content and the
fraction of the resident nematode population that was inactive (anhydrobiosis) was approximated with a sigmoid curve: AN 5 0.652 / (0.652 + M2)
where An is the inactive portion of the population and M is the percent
gravimetric water (n 5 74 r2 5 0.82). This model then was used to explore
the combined impacts of temperature and moisture on population dynamics,
given the observed sex ratio. Estimates of l were found to be sensitive to
both temperature and gravimetric moisture: l5 0.68 + 0.0019DD + 0.025M
- 8.18310-7DD2 - 7.5310-4M2, n 5 213, R2 5 0.68. In particular, the model
was most sensitive to conditions found in the McMurdo Dry Valley soils
(M , 10% and DD , 300).
WEIHER, EVAN.* University of Wisconsin - Eau Claire, Eau Claire, WI.
Functional traits, diversity, and other components of community assembly: tools, linkages, and beyond.
Assembly rules (the constraints that limit which subsets of a species pool
can exist as communities) remains an important yet daunting field. Critical
questions persist. Are plant communities functionally underdispersed (selected through environmental filtering) or overdispersed (show limiting
similarity)? If the former holds then plant functional groups and plant strategy theory are important avenues forward. Does functional diversity covary
with taxonomic diversity? How do diversity and functional plant traits vary
and covary along fundamental environmental gradients (i.e. with stress and
disturbance)? How does the intensity of assembly rules vary from place to
place? Here, I will discuss the strengths and weaknesses of three tools (null
models, structural equation modeling, and constraint envelopes) for testing
these hypotheses and for gaining understanding in several ecosystems (prairies, oak savannas, coastal sand dunes, wetlands). Successful use of these
methods obviates a balance between creative and critical thinking - as well
as courage (or is it hubris?). Gaining understanding also requires explicitly
linking our study of community assembly phenomena with other levels of
ecological organization and conceptual areas (e.g. evolution, landscapes,
ecosystems). This may also seem daunting, but it may be relatively simple
to layer these other factors onto our investigations. Examples of how this
can be achieved will also be discussed.
WEIN, GARY R.* WSRC-Soil and Groundwater Closure Program,
gary.wein@srs.gov, Aiken, SC, USA. Monitored natural remediation at
U.S. Department of Energys Savannah River Site, South Carolina.
Monitored Natural Attenuation (MNA) is emerging as a cost-effective minimally invasive way to remediate diffuse groundwater contaminant plumes
or surface contamination. Research at the U.S. Department of Energys
Savannah River Site (SRS) in South Carolina through its Monitored Natural Remediation (MNR) Program seeks not only to understand and document natural attenuation (NA), but to enhance natural attenuation through
engineered approaches such as phytoremediation, biostimulation, and
bioaugmentation. The MNR Program at the SRS seeks to: 1) determine if
NA occurs at contaminated sites; 2) Identify the mechanisms of NA; 3)
determine the rates at which NA occurs; and 4) identify the appropriate
MNA/MNR techniques for application at the SRS. In addition to these
objectives, efforts are underway to develop cost-effective monitoring techniques and technologies. The SRSs MNR program is an integrative multidisciplinary approach to develop the tools necessary to use MNA as a
remediation tool. Scientists and engineers from the Westinghouse Savannah
River Companys Soil and Groundwater Closure Program, Savannah River
Technology Center, University of Georgia, U.S. Forest Service, Clarke Atlanta University, University of South Carolina, and others are engaged in
a diversity of MNR efforts. This poster will present ongoing efforts and
results from the SRS MNR Program including: documenting NA of trichloroethylene (TCE) by microbial degradation and native plants, spray
irrigation of native vegetation and hybrid poplars with TCE;, microbial
degradation of TCE by biostimulation and bioaugmentation, microbial degradation of pesticides by windrow composting, phytoremediation of tritium
by spray irrigation, natural attenuation capacity of soils contaminated with
metals, and sequestration of cesium-137 by illitic clays.

352

Abstracts

WEINTRAUB, MICHAEL N.* and JOSHUA P. SCHIMEL. University of

California, Santa Barbara, Department of Ecology, Evolution, and Marine
Biology, Santa Barbara, CA. Plant uptake of amino acids in the Arctic
tundra of Alaska.
Amino acid uptake by plants was first reported in the Arctic tundra, and is
thought to be important there because both plants and soil microbes are
strongly N limited, and estimates of plant nitrogen uptake exceed the traditional measures of nitrogen availability. Because estimates for plant amino acid uptake range widely and have not accounted for microbial competition, it is still hard to draw solid conclusions about the importance of
amino acids to tundra plants. To assess the competitive partitioning of
amino acids between tundra plants and soil microbes, we added either
labeled ammonium or amino acids labeled with both carbon and nitrogen
isotopes to intact cores of the tundra sedges Eriophorum vaginatum and
Eriophorum angustifolium several times in the summers of 2000 and 2001.
Recoveries of the amino acids from the plants were not significantly different when calculated independently using either the carbon or nitrogen
isotopes, indicating that amino acids were taken up by the plants as intact
molecules. Up to 3% of the added amino acids and 5% of the added ammonium were recovered in Eriophorum vaginatum; and up to 1%, but
typically less than 0.5% of the amino acids and up to 1.5% of the ammonium were recovered in Eriophorum angustifolium. We also found that
Eriophorum vaginatum took up adenine in similar proportions to amino
acids. On all but one occasion both plant species took up more of at least
one of the added amino acids than they did ammonium, indicating that
these tundra species often compete well for amino acid N relative to ammonium. To determine the role of uptake kinetics in the competitive partitioning of amino acids and ammonium, we separately quantified the uptake kinetics of both Eriophorum vaginatum excised roots and soil at the
same times as the intact core experiments in the summer of 2000. We
hypothesized that plants would compete best for the compounds that had
the lowest uptake rates in soil. Arginine and adenine did fit this pattern,
and were the two compounds beside ammonium that were best taken up
by Eriophorum vaginatum. Ammonium did not fit this pattern, however,
and was the compound that had the highest uptake rates in both plants and
soil.
WEIR, ENRIQUE H.,1,* KARINE C. GIL-WEIR2,* and CLARK L. CASLER. 1 Texas A&M University, Dept. Wildlife and Fisheries Science, 210
Nagle Hall, 2258 TAMU, College Station, Texas, USA; 2 Texas A&M University, Dept. Wildlife and Fisheries Science, 210 Nagle Hall, 2258 TAMU,
College Station, Texas, USA. Bird richness and abundance at The Olivitos Wildlife Refuge and Fishing Reserve, Zulia State, Venezuela.
We studied bird species composition and abundance monthly from January
1999 to June 2000, in The Olivitos Wildlife Refuge and Fishering Reserve,
Zulia State, Venezuela. This is a Ramsar wetland, national protected area
and the only location in the country where the flamingos nest. The bird
community was evaluated in six areas associated to the Olivitos Mangrove
Ecosystem: one area located in the swamp, two estuarine areas and three
areas located inside the mangrove forest. We used a boat with a 40 Hpmotor to cover each habitat, during one hour, with a slow constant speed.
The birds were observed with the help of binoculars (50x20 and 40x8).
We observed differences in the abundance, richness and composition between habitats and the seasonality. We observed changes in the species
richness, abundance and composition in the Olivitos Mangrove between
our research and the reports of Casler and Lira (1982) and Casler, Weir
and Aniyar (1996).
WEISER, MICHAEL D.,1,* BRIAN J. ENQUIST,1,2 BRAD BOYLE1,2 and
SRINIVAS REDDY.1 1 University of Arizona, Tucson, AZ, USA; 2 Center
for Applied Biodiversity Science- Conservation International, Washington,
DC. Rapaports rule, latitudinal patterns, and the frequency distributions of range sizes of New World woody plant species.
Rapaports rule posits that species from higher latitudes have ranges with
greater latitudinal extent. This pattern has been claimed for North American
trees but has not been analyzed for temperate and tropical trees across
North, Central, and South America. Recent analyses indicate that tropical
trees may have larger ranges that originally thought, casting doubt on the

importance of Rapaport patterns driving tropical alpha diversity. Here we

use a global compendium of forest plot and herbarium data to examine the
range sizes of New World woody plants. Latitudinal range extents are determined by using specimen-based range maps collected from a network
of herbarium datasets. We specifically focus on the how the frequency
distributions of range sizes and latitudinal extent of range vary with latitude
from Canada to Chile.
WEISHAMPEL, JOHN F.1,* and GEOFFREY G. PARKER.2 1 Department
of Biology, University of Central Florida, Orlando, FL, USA; 2 Smithsonian
Environmental Research Center, Edgewater, MD, USA. Characterizing
successional volumetric patterns of canopy architecture.
Structurally, a forest canopy is analogous to a fractal spongea volume
of spaces and phytomass between two (as found with a planar object) and
three (as found with a solid object) dimensions. The spatial organization
of canopy components (i.e., leaves, twigs, branches, etc.) directly influences
the biotic habitat (e.g., light, temperature, humidity) and hence, atmospheric
exchange, and tree growth and physiology. To measure canopy architectural
dynamics associated with forest succession, data were collected using a
handheld laser instrument to measure zenith-projected surfaces in four (30
x 30 x 40 m) forest stands. These stands represent different successional
stages of the "tulip poplar" association, a mixed deciduous forest type
found in eastern Maryland, USA. Its climax stage typically possesses an
overstory dominated by Liriodendron tulipifera. This chronosequence, representing young, intermediate, mature, and old-growth stands, has been
characterized vertically as having: a compressed monomodal canopy, an
expanded monomodal canopy, a bimodal leaf distribution, and an uneven
leaf area distribution, respectively. Using three-dimensional textural analysis techniques coupled with neutral modeling, we explored the lacunarity
and multifractal patterns of the morphological elements and spaces that
comprise these volumes as determined by the laser measuring system. This
neutral modeling approach is a derivative of percolation theory, which is
commonly used by landscape ecologists for depicting patterns and processes between one and two dimensions. Here, it is applied to provide a
basic interpretation and discrimination of these complex, hierarchical structures.
WELCH, NICOLE T.,1,* JONATHON M. BELMONT2 and J.C. RANDOLPH.2 1 Middle Tennessee State University, Murfreesboro, TN; 2 Indiana University, Bloomington, IN. Carbon and macronutrient content of
herb layer vegetation in an Indiana temperate deciduous forest.
While herb layer tissue nutrient content (mg/g) often decreases with CO2
enrichment, the increase in plant biomass under these conditions results in
an overall increase in nutrient content of the herb layer (g nutrient/m2).
Baseline herb layer nutrient and biomass data will be crucial for detecting
if similar changes occur in the field with global climate change. This study
assessed biomass and carbon and macronutrient concentrations of herb layer vegetation (forest vegetation ,1 m in height) at the AmeriFlux Tower
Site in the Morgan-Monroe State Forest, Indiana. Two hundred and fifteen
1-m2 plots were sampled for herb layer species composition and cover.
Forty of these plots were harvested for above ground biomass determination and tissue nutrient analysis. Lindera benzoin, Boehmeria cylindrica,
Viola sp., Parthenocissus quinquefolia, and Ploystichum acrostichoides
were the most important herb layer species. Mean herb layer biomass was
38.50+8.08 g/m2. Forbs accounted for approximately 14.56 g/m2 of the herb
layer vegetation followed by grasses (5.73 g/m2) and woody shrubs and
seedlings (21.06 g/m2). Forbs, grasses, and woody shrubs and seedlings
averaged 6.04, 2.32, and 9.54 g C/m2, respectively. In general, macronutrient concentrations for these herb layer components were N . K . Ca
. Mg . P. Mean macronutrient concentrations were highest in forbs (0.35
g N/m2, 0.45 g K/m2, 0.15 g Ca/m2, 0.06 g Mg/m2, 0.26 g P/m2) followed
decreasingly by woody shrubs and seedlings, and grasses. Predicted increases in CO2 with global climate change could alter these nutrient concentrations and, in turn, alter forest nutrient cycles and herbivore food
quality.

WELLBORN, GARY A.* and RICKEY D. COTHRAN. University of

Oklahoma, Norman, OK. Ecology and mechanisms of coexistence in
three sympatric cryptic species in a freshwater amphipod species complex.
Among the most conceptually challenging findings arising from the growing application of molecular phylogenetic analyses to natural populations
is the frequent discovery of unexpected species diversity within seemingly
phenotypically homogeneous taxa. Although traditional coexistence concepts contend that coexistence of congeneric species is sustained by traitmediated niche differentiation, the close phenotypic similarity of sympatric
cryptic species implies limited scope for niche separation, thus taxing our
understanding of species coexistence. We examined phenotypic diversification, microhabitat distribution, and vulnerability to predation in three
undescribed cryptic species of Hyalella amphipods that commonly coexist
in the littoral zone of lakes. Because we know of no distinguishing morphological characters for the species, all species identifications were established using a PCR screening method. We found that, although there is
extensive overlap in body size, the three species differ in maturation size
and mean adult size. Quantitative differences in some other life history
traits were also observed. A study of the relative frequency of the species
across a depth gradient in the littoral zone revealed strong complementary
structure in species distributions, with the smallest species most prevalent
in the deepest region, and the largest species most prevalent in the shallowest microhabitats. Finally, a study of fish feeding patterns in the field
demonstrated that relative predation risk differed among the species and
was positively related to the mean body size of species. Because resource
consumption rate and competitive ability increase with body size, our results suggest that these sympatric cryptic Hyalella species experience a
tradeoff between resource exploitation and predator avoidance, a condition
that may foster coexistence. We interpret our study in the context of food
web models addressing coexistence of consumer species that share a common resource and predator. In particular we explore the congruence of our
results with somewhat counterintuitive model predictions that close phenotypic similarity among species may promote coexistence in heterogeneous environments.
WELLMAN, MARJORIE C.,1,* SUSAN B. NORTON,1 RICHARD A.
PARK2 and JEFFREY B. FRITHSEN.1 1 US Environmental Protection
Agency, Washington, DC, USA; 2 Eco Modeling, Diamondhead, MS, USA.
Stressor identification with the ecosystem model AQUATOX, Release
2.
Increasingly, the regulatory, remediation and restoration actions taken to
manage impaired environments are based on measurement and analysis of
biotic communities. When an aquatic community has been identified as
impaired, the cause of the impairment must be determined so that appropriate actions can be taken. The US Environmental Protection Agencys
Stressor Identification (SI) Guidance (EPA 2000) describes a methodology
for identifying the causes of observed impairments in aquatic systems.
Stressor identification requires extensive knowledge on a variety of stressors as well as depth of knowledge on the mechanism, symptoms, and stressor-response relationships for specific stressors. AQUATOX is a processbased, time-varying, fate and effects simulation model that integrates aquatic ecology, chemical dynamics, bioaccumulation, and ecotoxicology.
AQUATOX can be used to predict not only the environmental fate of
nutrients, pesticides, and other chemicals in aquatic ecosystems, but also
their direct and indirect effects on organisms. Time-varying process rates
and limitations are provided for detailed analyses. Therefore, the AQUATOX model is a valuable tool for stressor identification, specifically to
evaluate the relative importance of multiple stressors, provide ecologically
plausible causal scenarios, and eliminate unlikely causes. Examples will
include an application to Coralville Reservoir, Iowa, where model results
indicated that pesticide runoff, not nutrient or organic enrichment, was the
primary reason that largemouth bass were not present. Model results were
supported by the recovery of bass in Coralville Reservoir after the banning
of dieldrin.
WELLNITZ, TODD and LEROY POFF. Biology Department, Colorado
State University, Fort Collins, Colorado. Does heterogeneity in near-bed
current mediate competition between stream herbivores?
Herbivore guilds in streams can be speciose despite the fact grazers are
often food-limited. This suggests that variation in environmental conditions

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353

may influence the outcome of species interactions and thus potentially mediate herbivore co-existence. To examine the role of heterogeneity in nearbed current in this regard, we conducted a 3 week competition experiment
in streamside channels along the upper Colorado River using 3 current
velocities (5, 15 and 30 cm/s), and 5 densities (0, 0.5x, 1x, 2x and 3x
ambient densities) for each of two grazers, the mayfly Drunella, and the
caddisfly Glossosoma. Grazer weight gain and periphytic responses to herbivores were both influenced by current velocity. For example, Glossosoma
grew faster and were more effective at removing periphyton as current
velocity increased, suggesting higher feeding efficiency under high flows.
At the slowest velocity, periphytic abundance actually increased with Glossosoma density. Caddisfly weight gain was inversely related to larval density, implying intraspecific competition. Interestingly, Glossosoma weight
actually increased with increasing Drunella density, suggesting facilitation
by this mayfly on Glossosoma feeding. Evidence from this study and others
indicate that heterogeneity in near-bed current can influence species interactions among stream herbivores, and thereby potentially mediate patterns
of species co-occurrence.
WELLS, CHRISTINA E.* and DESMOND LAYNE. Department of Horticulture, Clemson University, Clemson, SC, 29634. Fine root dynamics
of peach (Prunus persica) under two irrigation regimes.
The use of minirhizotrons (root observation tubes) has generated information on fine root dynamics in many plant species and ecosystems. However, the extent to which specific environmental variables affect rates of
root production, aging and mortality is still poorly understood. We installed
72 minirhizotron tubes beneath 4-yr-old trees in a peach production systems
trial to determine how root morphology, architecture and demography differed in trees with and without supplemental irrigation. Red Globe peach
trees were grown on one of two rootstocks (Lovell or Guardiany) and
received one of two irrigation regimes (rainfall only or 1 inch of supplemental irrigation per week). Beginning in May 2002, videotaped images
from each minirhizotron tube were collected at two week intervals; notes
on tree phenology were also recorded on these dates. Videotapes were
digitized in the lab, and information on root length, diameter and appearance was extracted from the images. Soil temperature, soil volumetric water
content and pan evaporation were also measured in the orchard throughout
the growing season. Initial data indicate that non-irrigated trees produced
thinner fine roots (P , 0.05) that were more highly branched (P , 0.05)
than those of irrigated trees. Non-irrigated trees also allocated more of their
fine root length to deeper soil layers (P , 0.08). There were significant
differences in root length production between rootstocks, with the nematode-tolerant rootstock, Guardiany, producing more length per tube under
both irrigation regimes (P , 0.01). Analysis of fine root demography will
begin in 2003, one year after the disturbance of minirhizotron installation.
Previous research in peach has shown that thinner roots and roots of lower
branching order have shorter life spans than thicker, higher-order roots.
Whether the thinner, highly-branched root system of non-irrigated trees
experiences higher rates of root turnover is a question we will address in
the coming year.
WELTZIN, JAKE F.* University of Tennessee, Knoxville, TN. Changing
climates, invasive species, and encroachment of woody plants in arid
and semi-arid ecosystems.
Coincident with a world-wide increase in the abundance of woody plants
within former grasslands, grasslands throughout the western United States
have experienced a substantial increase in the abundance of woody plants
(e.g., Prosopis, Juniperus)within the last century. These changes in grassland and savanna structure have been accompanied by changes in other
biotic and abiotic factors, including grass invasions, livestock grazing, fire
suppression, landscape fragmentation, and climate change. Combined, these
environmental driving variables may alter disturbance regimes, soil resources (e.g., water, nutrients), biotic interactions, and seed dispersal, with
likely consequences for woody plant demography, savanna structure, and
biological diversity. Predicting the response of semi-arid systems to climate
change or biological invasions will require an increased understanding of
the processes that control population dynamics, under a variety of plant
invasion and climate change scenarios, with the recognition that other factors (e.g., soil texture, grazing) may also be important at a variety of dif-

354

Abstracts

ferent scales. Moreover, interactions between driving factors may control

the response of a system to a given perturbation. I briefly review our current
understanding of the importance of changes in precipitation regimes and
plant invasions with respect to woody plant recruitment, discuss the importance of spatial and temporal scales when considering these factors, and
provide a case study that illustrates the multifaceted role that climate
change and invasions may play in structuring grassland and savanna ecosystems.
WENDEL, HEATHER E.* and DARREN SANDQUIST. California State
University, Fullerton, CA. The effects of invasive exotic annuals on the
water relations and productivity of native endangered Eriastrum densifolium ssp. sanctorum (Polemoniaceae) shrubs.
Flood control programs along the Santa Ana River prevent the natural
removal of perennial shrubs and leaf litter which modify alluvial scrub
habitat and reset community succession. The lack of periodic flooding and
deposition of wind-blown soil particulates from adjacent mining industries
create suitable soils for invasive exotic grasses and forbs. Eriastrum densifolium ssp. sanctorum is a California endangered subshrub restricted to
alluvial communities immediately following sand deposition by floods. I
hypothesize that competition with exotic annuals will decrease the amount
of soil water available to the deep taproot of E. d. sanctorum and ultimately
inhibit growth and productivity. I established three removal treatments (removal of other native shrubs only, removal of shrubs and annuals, no removal) within solitary E. d. sanctorum plots (N59) and measured predawn
water potential, transpiration, stomatal conductance and soil water content
values. To assess productivity differences, I determined the leaf carbon and
nitrogen isotopic ratios and followed growth and seed production for one
season. Although trends suggest the release from competition increases
water use (greater water potential and transpiration values for plots with
shrubs and annuals removed), shoot growth, and results in higher seed
counts, there were no significant differences found between treatments (pvalue.0.05). Nutrient-level differences as determined by percent nitrogen
in the leaves (p-value 5 0.002), imply annuals may be altering the soil
chemistry. While there is no evidence to support an effect of annuals on
the water relations of E. d. sanctorum, compensation through increased leaf
production or nutrient cycling may yield different results. Further experimentation is needed to test these hypotheses in years of non-drought and
with larger sample sizes.
WENDER, NAOMI J. and KATHLEEN DONOHUE. Department of Organismic and Evolutionary Biology, Harvard University, 22 Divinity Ave,
Cambridge, MA, 02138. Genetic constraints on the evolution of dispersal are mediated by phenotypic plasticity.
We investigated the quantitative genetics of seed dispersal in Arabidopsis
thlaiana and found a novel genetic constraint on the evolution of dispersal
that is mediated by phenotypic plasticity. We grew several ecotypes of A.
thlaiana in high and low density, measured maternal plant characters, and
then characterized seed dispersion patterns in a wind tunnel. We identified
maternal plant traits that influenced seed dispersion patterns and found
significant genetic variation for these maternal traits. The genetic variation
for maternal traits that influence seed dispersion patterns, moreover, did
not differ with density. However, these plant traits determined dispersal
more strongly when plants were grown at high density than when they
were grown at low density. In addition, we found significant genetic variation for both dispersal distance and for seed density after dispersal. Genetic variation for seed density after dispersal, however, was only detectable
at high density. Therefore, as increased dispersal ability evolves and plants
experience lower densities as a result, the expression of genetic variation
for dispersal is expected to decline. Thus, a limit to the evolution of dispersal would result. In this study, we therefore detected a novel environment-dependent genetic constraint that is mediated by the evolution of the
trait itself.
WENGER, SETH J.,1 BYRON J. FREEMAN,1,2 MARY C. FREEMAN,3
ALLISON H. ROY1 and WILLIAM E. ENSIGN.4 1 University of Georgia,
Athens, GA; 2 Georgia Museum of Natural History, Athens, GA; 3 United
States Geological Survey, Athens, GA; 4 Kennesaw State University, Kennesaw, GA. Assessing effects of development on imperiled fishes for the
Etowah Regional Habitat Conservation Plan.
Nineteen counties and cities have begun development of a regional Habitat
Conservation Plan (HCP) for imperiled aquatic species in the Etowah River

watershed, a highly diverse river system just north of Atlanta, GA, USA.
Rapid urbanization in the region threatens the survival of nine fish species,
some of which still await formal description. The purpose of the HCP is
to minimize impacts of development on these species through growth management policies and regulations. However, we presently lack sufficient
scientific information to determine the most effective tools for achieving
this goal. To address this problem, we have initiated a program of study
designed to answer some of our most pressing research questions: (1) What
are the most significant mechanisms by which altered hydrology impacts
aquatic species? (2) Is channel erosion from altered stream flows a significant source of sedimentation in developing watersheds? (3) Do riparian
buffers play a significant role in protecting aquatic habitat in urban areas,
or are they bypassed by storm drainage networks? (4) Do some culvert
designs present greater genetic barriers than others? (5) Is development in
specific subwatersheds impacting mainstem species? (6) Does clustered development provide better protection of aquatic habitat than sprawling development, on the landscape scale? We will use the results of these studies
both to direct and to justify the policies and regulations recommended for
the HCP.
WENNERBERG, SARAH B.,* KATRINA R. KLUGH, CLINT J.
SPRINGER and ERIC J. PYLE.* West Virginia University, Morgantown,
WV. Ecological education enrichment in grades 5-10 of West Virginia
schools by NSF GK-12 teaching fellows.
Teams of interdisciplinary graduate fellows in science, mathematics, and
engineering make up the TIGERS project at West Virginia University
(WVU). The TIGERS project is part of the National Science Foundations
GK-12 program. Middle and high school (5-10) math and science teachers
work cooperatively with teams of graduate fellows to co-plan and co-teach
science and math topics throughout the school year. During this co-planning
and co-teaching, graduate fellows provide real world scientific expertise
while the teachers help guide the fellows in preparing a series of hands-on
inquiry based lessons to present to the students. The TIGERS Humans and
the Environment team is comprised of graduate students from the departments of biology and physics at WVU. The developed lessons cover many
ecological themes such as nutrient dynamics of plants in disturbed ecosystems, plant identification, and forest succession. Experimental design and
data collection in field and laboratory settings along with computerized
data analysis are emphasized in most lessons. Student mastery of the intended objectives is determined using multiple assessment strategies. The
results of student assessments show that West Virginia middle and high
school students display an increase in knowledge of human and environmental interactions as well as a greater understanding of humans role as
stewards of Earths ecosystems.
WENTWORTH, THOMAS R.,1,* PETER S. WHITE,2 BROOKE E.
WHEELER,2 KRISTIN TAVERNA,2 DANE KUPPINGER,2 LEE ANNE
JACOBS,2 JASON D. FRIDLEY2 and ROBERT K. PEET.2 1 North Carolina State University, Raleigh, NC; 2 University of North Carolina, Chapel
Hill, NC. Nested versus non-nested characterization of vegetation composition and species richness at multiple spatial scales.
The importance of characterizing vegetation composition and species richness at multiple spatial scales is increasingly recognized by ecologists, but
there is no consensus as to whether variation with scale is best characterized
by subplots arranged in a design that is nested or non-nested. Many crucial
aspects of biodiversity management and research may depend on this design choice. The fundamental difference between these designs is in how
they are influenced by spatial autocorrelation structure and in how well
they allow characterization of change in vegetation structure with change
in grain of observation. We compared nested and non-nested designs for
characterizing species richness (as species-area and species-accumulation
curves) of vascular plants using data collected by the Carolina Vegetation
Survey in 0.04 ha plots. If the goal is to provide a strategy to inventory
all species in a large plot, neither design presents a clear advantage. Although either design can be used to examine species-area relationships, the
nested design constrains the result to the desired monotonic response for
each replicate. The non-nested design is superior when the goal is to find
as many species as possible within a specific subsample of a larger target
area. The nested design offers a means to interpret underlying spatial pat-

terns of richness that the non-nested design does not. Only the nested design is appropriate if the goal is to assess changes in species composition
with changing grain size of observation, because the non-nested design
confounds the influence of grain with that of extent. We also explored the
complex question of which design allows more effective extrapolation of
richness to larger spatial extent. We conclude that the nested design is
equivalent or superior to the non-nested design for most applications and
should be the standard method for multi-scale inventories.
WEST, JASON B.,1,* DAVID WEDIN,2 SARAH E. HOBBIE1 and PETER
B. REICH.1 1 University of Minnesota, St. Paul, MN; 2 University of Nebraska, Lincoln, NE. The dominant role of plant species in controlling
the response of N mineralization to altered plant diversity, atmospheric
CO2, and N deposition.
Declining biodiversity and increasing nitrogen inputs and atmospheric CO2
are important global changes that strongly affect terrestrial ecosystems. We
present results from a long-term experiment where we manipulated plant
species biodiversity, atmospheric CO2, and N deposition (BioCON) in Nlimited grassland plots in central Minnesota and examined the consequences for net N mineralization rates (Nmin). As N limitation is a common
feature of terrestrial ecosystems, the response of N cycling to global change
factors may shape overall ecosystem response to global change. Consistent
with our predictions, diversity strongly decreased Nmin. This effect was
evident in the first year of plant growth. The difference among diversity
treatments dampened over time, suggesting that greater species diversity
may accelerate the return of plant-soil interactions characteristic of prairie
ecosystems. The results from year 5 revealed an interaction between elevated CO2 and N. Elevated CO2 decreased Nmin at ambient N, whereas it
stimulated Nmin at elevated N. The observed changes in Nmin in mixedspecies plots are likely the result of shifts in dominance in response to the
treatments. Consistent with previous research, we observed large species
effects on Nmin. Most of the effects in ambient plots corresponded to
known plant trait differences between and within functional groups (i.e. C3
& C4 grasses, C3 forbs, legumes). However, both the magnitude and direction of changes in Nmin in response to CO2 and N differed widely
among species. For example, elevated CO2 dramatically stimulated Nmin
in Koeleria cristata plots, whereas there was no effect of CO2 for Agropyron repens, both C3 grasses. The overall time course likely represents
initial immobilization of N, followed by N cycle changes from plant effects.
Understanding the mechanisms of these responses improves our ability to
generalize about the effects of global change. Gross N mineralization measured in year five suggests that plant effects on Nmin are caused by changes
in N immobilization. Our results demonstrate important effects of global
changes on N cycling, and the primary role of plant composition in determining the direction and magnitude of these responses.
WESTBURY, HUGH M.* USACE ERDC CERL, Fort Benning, GA, USA.
Conducting ecological studies on military reservations.
The Department of Defense (DoD) manages large tracts of land in support
of training and mission readiness. These reservations, particularly in the
eastern U.S., are some of the least developed lands and offer substantial
opportunities for ecological studies. In addition to numerous regulatory
requirements that mandate good stewardship, the military must also maintain their training areas in good condition to insure the long-term sustainability of their mission. To meet these goals, DoD installations require
detailed understanding of their ecosystems. DoD installations provide the
opportunity and funding for research in a variety of ecological conditions,
as well as presenting unique challenges to traditional research practices.
The Strategic Environmental Research and Development Program
(SERDP) Ecosystems Management Project (SEMP) is an ecological research and development project that is improving the knowledge, tools and
techniques that are needed to support management of Department of Defense lands. SEMP provides a framework for SERDP-funded, large, longterm ecological research and monitoring projects at Fort Benning, Georgia.
An obvious initial concern was for the safety of ecological field crews on
this active military installation where the conduct of realistic training requires field exercises using live ammunition, pyrotechnics and heavy vehicles. To insure crew safety and minimize the impact of research on the
training mission, SEMP funded a full-time Host Site Coordinator at Fort

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Benning. By the end of 2002, SEMP had conducted more than 1800 field
trips without a safety incident and without interfering with training. The
Host Site Coordinator was also tasked with documenting problems encountered and solutions achieved at Fort Benning in order to smoothly
migrate SEMP ecological research to other installations. Examples of these
"lessons learned" are scheduling procedures, safety briefings, team responsibility, awareness of military culture, and research equipment survivability.
WESTOVER, KRISTI M.* and JAMES D. BEVER. Soil pathogens and
maintenance of plant community diversity.
The coexistence of competing species may be mediated through interactions with other trophic levels. For plants, many of these trophic interactions occur below ground. The direct effects of soil microorganisms on
plant growth are known to be profound. Yet, with the difficulty of monitoring soil organisms, their importance to plant community processes is
easily overlooked. In this talk, we test the role of soil community dynamics
in the maintenance of plant species diversity in a North Carolina grassland.
We find that the presence of a particular plant species changes the composition of its soil community in a manner that decreases the growth rate
of that plant species relative to that of a second plant species. This negative
feedback results in part from the accumulation of host-specific soil pathogens in the genus Pythium. But these negative feedbacks can also result
from shifts in the species composition of rhizosphere bacteria and arbuscular mycorrhizal fungi. The negative feedbacks through host-specific
changes in the composition of the soil community can directly maintain
plant species diversity. We test this possibility with for two common plant
species: Anthoxanthum and Panicum. We find that Anthoxanthum is predicted to exclude Panicum in the absence of soil community dynamics. In
the presence of soil community dynamics, however, negative feedbacks on
Anthoxanthum prevent it from dominating and both species coexist. In this
system, the soil community dynamics appear to be essential for their coexistence.
WETZEL, PAUL R.,1,* WILEY M. KITCHENS2 and MARSHA DUSEK.2
1
Smith College, Northampton, Massachusetts, USA; 2 University of Florida, Gainesville, Florida, USA. Using classification tree analysis and
structural equation models to investigate changes in tidal marsh communities.
The tidal delta of the Savannah River (Georgia, USA) supports a dynamic
marsh plant community that responds to environmental gradients that include river flow, sediment distribution, salinity regimes, and seasons of the
year. Recent harbor development, the installation (1977) and removal of a
tidal gate (1993) and the dredging of the main river channel (1993-1994)
resulted in differential compression/expansion of these gradients. This
study examined the responses of the marsh communities to natural and
anthropogenic variations in these gradients. Vegetation, sediment, and interstitial salinities were sampled at 4-7 sites along the tidal gradient. Sites
were sampled 11 times (7 different years) over a 17 year period. This long
term information provided a unique opportunity to investigate tidal marsh
community in two ways: 1. To determine which factors best differentiate
plant functional groups and to use that information to predict future community level changes, and 2. Use structural equation modeling to confirm
conceptual models of relationships between plant species, environmental
parameters, and time for tidal marsh ecosystems. Multivariate classification
tree analysis was used to differentiate eight species functional groups. Spartina alternaflora and Spartina cynosuroides/Scirpus validus functional
groups were found at interstitial soil salinities . 5.25 ppt. The S. alterniflora group disappeared after 1993 and was replaced by the S. cynosuroides/S. validus group. In low salinity environments, the Eleocharis montevidensis group grew in soils with a bulk density . 0.2 g/cm3. Distance
from distributary channels built for past rice cultivation differentiated the
S. validus and Zizaniopsis milacea groups. A conceptual model of tidal
marsh vegetation was developed from published literature and information
collected over the sample period. Structural equation modeling was used
to confirm the strength of direct and indirect pathways between variables
in the conceptual model. It was also used to compare conceptual models
through time, specifically before and after major known natural and human
disturbances to the tidal marsh communities.

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WHITBECK, JULIE L.* University of New Orleans, Julie_Whitbeck@

alum.swarthmore.edu, New Orleans, LA, USA. Setting bounds on belowground NPP: Evaluating three approaches in a bottomland hardwood
forest.
Because existing methods for assessing belowground net primary production (BNPP) have important limitations, my objective is to set upper and
lower bounds on the fine root component of the carbon (C) budget by using
three distinct approaches in parallel: the compartment flow method employing sequential coring with decomposition corrections, minirhizotron
based root demography, and the Cbalance (or total belowground C allocation) method calculated from soil respiration and fine litterfall. In addition to bracketing BNPP, I evaluate points of agreement and difference
among these widely used methods. This study is set in an old growth
bottomland hardwood forest at the southernmost end of the Lower Mississippi Alluvial Valley. My research team sampled at three points along a
hydrologic gradient in this forest, ranging from a rarely flooded natural
levee "ridge" site to a regularly inundated (59 month hydroperiod)
"swamp" site. Tree species richness and total basal area are similar at the
three focal sites, and soils along the entire gradient are fine textured Sharkey clay. My multi-year dataset spans extremely dry through normally wet
years. In contrast to many upland ecosystems, fine root growth in periodically flooded bottomland forests is at least as sensitive to changes in soil
water content as to variation in temperature. My data indicate that fine root
production and turnover in this ecosystem are limited by both low and high
soil water content and are correlated with ambient temperature. While all
three approaches captured these patterns, the carbon balance approach
yielded the highest estimates of BNPP, while minirhizotronbased estimates
were consistently low. Variation in BNPP estimates differs among approaches and is influenced by key environmental parameters.
WHITE, CARLETON S.,* DOUGLAS I. MOORE and JOHN A. CRAIG.
Biology Dept., MSCO3 2020, University of New Mexico, Albuquerque,
New Mexico. Regional-scale drought increases potential soil fertility in
semiarid grasslands.
Soils from a semiarid grassland on the Sevilleta National Wildlife Refuge
(NWR) were sampled from 1989 to 1998 for field-extractable nitrogen (N)
and potentially mineralizable N (PMN, incubations at near-optimal soil
moisture and temperature to measure potential soil fertility). These forms
of soil N were highest in collections from 1989 and 1995, which were
periods of regional-scale drought. Although previous work has suggested
that drought increases soil fertility in arid and semiarid systems, a direct
link between soil fertility in field samples and weather fluctuations has not
been documented. Whereas previous soil collections on the Sevilleta NWR
were made for different reasons, this study collected semiarid grassland
soils in June 2002 near the peak of a regional-scale drought to test the
hypothesis that potential soil fertility was increased by drought. An additional semiarid grassland site, the Bernalillo Watershed, was sampled to
extend the spatial extent of the analysis. The 2002 collections had soil PMN
near the highest at both sites, thereby supporting the hypothesis. The Palmer Drought Index (PDI) was used as the measure of regional-scale drought
and correlated with PMN at both sites. The correlation was significant (r
5 -0.533, P , 0.05) for the Sevilleta soils with higher PMN during periods
of drought over the 13-year period. A similar trend was shown over the 8year period at the Bernalillo Watershed, but the correlation was not statistically significant. Previously published results on mineralizable N from
another semiarid grassland site on the Sevilleta NWR were highly significantly correlated with the PDI (r 5 -0.723, P , 0.01), indicating that a
three-year wet-to-drought oscillation had significant effects on ecosystem
processes.
WHITE, DAVID L.* and JOAN L. WALKER. Southern Research Station,
USDA Forest Service, Clemson, SC, USA. Structure and composition
of 7080 yearold Fraser fir forests on Roan Mountain.
Roan Mountain and other Southern Appalachian sprucefir forests have
been heavily impacted by the Balsam Wooly Adelgid (Adelgis piceae
Ratz.), historical grazing, logging, fires and other disturbances. Roan
Mountain is also the only National Forest area where Fraser fir (Abies
fraseri(Pursh) Poir) seed and seedlings are harvested for the Christmas tree

industry. To address concerns about the condition of fir forests on Roan,

forest structure and composition were examined in 2000. Thirteen, 20320
m permanent plots were established in 7080 yrold firdominated stands
on southfacing slopes (el.17501850 m). Tree ($10 cm d.b.h.) basal area
of Fraser fir, red spruce (Picea rubens Sargent), all other species (OTHER),
and total dead species (DEAD) averaged 31.5, 11.8, 0.4 and 10.4 m2 ha-1,
respectively. Density for these same groups averaged 729, 229, 25 and 340
stems ha-1, respectively. Density of saplings (.1.4 m tall and ,10 cm
d.b.h.) of fir, spruce, OTHER and DEAD were 1971, 185, 2491 and 852
stems ha-1, respectively. Rubus species accounted for a majority of OTHER
stems. Its abundance in 30% of the plots was associated with increased
canopy openness. Stand disturbance, measured by canopy openness and
tree mortality, varied considerably across plots, with dead trees comprising
4 to 61 % of total (live + dead) basal area. Fir seedling abundance (stems
ha-1) of the smallest size class (unbranched) averaged 141,000 and varied
from 0 to 350,000. Regression analysis of the smallest size classes of fir
seedlings indicated that total dead saplings (+), understory herb cover (+),
and total live saplings (-) were significant (p,.01) predictors of seedling
abundance. The variation in size class distribution patterns found in this
study are similar to the range found in the Great Smoky Mt. NP, where
differences were attributed to time since adelgid-induced disturbance
events.
WHITE, ETHAN P.,1,* PABLO A. MARQUET2 and JAMES H. BROWN.1
1
University of New Mexico, Albuquerque, NM; 2 Pontificia Universidad
Catolica de Chile, Santiago, Chile. Linking the scaling of body size distributions and the species-area relationship.
The scaling of body size distributions (BSD) in mammals, from nearly loguniform in local communities to the traditionally observed right-skewed
modal distribution at continental scales, has received considerable attention
over the last decade. It can be shown that the scaling of the body size
distribution and the slopes of the species-area relationships (SAR) for the
different body-size classes are directly connected. The slope of the SAR
must be steepest at the modal body size class and must continue to decrease
monotonically to either side of the mode in order for the observed pattern
to exist. We compare the BSD scaling of birds to that previously observed
for mammals and show that in birds the BSD is scale invariant and that
the slope of the SAR does not change regularly with mass. This implies
that there are taxonomic differences in the scaling of BSDs that are potentially due to some combination of differences between birds and mammals
in dispersal limitation and/or assembly rules. The implications of these
differences for elucidating the underlying processes of the SAR are discussed.
WHITE, JENNIFER A.* and DAVID A. ANDOW. University of Minnesota, St. Paul, MN. Should I stay or should I go? Parasitoid foraging
for intermittently concealed hosts.
Many insect herbivores feed in concealed locations and thus limit their
susceptibility to foraging natural enemies. However, some herbivores move
into and out of concealment many times over the course of their development, creating windows of greater vulnerability to attack. Parasitoids that
specialize on such hosts face an interesting foraging decision: having located an inaccessible host, how long should the parasitoid wait for the host
to become accessible? We explored this question with a simple foraging
model and found that parasitoids should take one of two strategies, depending on three parameters. Parasitoids should leave immediately upon
discovering a host is inaccessible if 1) the probability of locating another
host is high, 2) the proportion of hosts that are accessible at any point in
time is high, and 3) the probability that an inaccessible host will emerge
from hiding is low. Conversely, the parasitoid should wait indefinitely for
hosts to emerge from concealment when the opposite conditions hold. We
then estimated these parameter values for the European corn borer, Ostrinia
nubilalis, and its specialist parasitoid, Macrocentrus grandii. From these
values, we predicted that M. grandii should adopt a "staying" strategy,
persisting in the vicinity of an inaccessible host until the host becomes
accessible. We confirmed that M. grandii does have a strong tendency to
persist near inaccessible hosts in laboratory assays. However, leaving rates
are higher under field conditions where environmental conditions and other
disturbances often terminate foraging bouts, suggesting that external factors
are important in determining the realized foraging behavior of M. grandii.

WHITE, JOSEPH D.,1,* KEVIN J. GUTZWILLER,1 WYLIE C. BARROW2 and LORI RANDALL.2 1 Baylor University, Waco, TX; 2 U.S. Geological Survey, National Wetlands Research Center, Lafayette, LA. Modeling vegetation community response to climate change and CO2 increases in the Chihuahuan Desert.
Future climate changes coupled with increases in atmospheric CO2 may
favor increased expansion and growth of woody shrubs in historically
grass-dominated desert communities. We developed a spatially explicit
mechanistic model to explore physiological and landscape processes associated with this potential conversion. The model predicts plant growth
and allocation for different functional types; reproductive dispersal through
varying pathways; and fire spread and effects. This model was applied to
a region including Big Bend National Park to assess climate change and
CO2 increases on community composition, above and belowground production, and fire frequency over the next 50 years. Average annual temperature for this area is expected to increase by 2 to 48C with slight decreases in annual precipitation. Simulation results showed that potential
future plant communities are dominated by woody shrub species as a combined effect of C3 photosynthesis physiology, greater drought tolerance,
longer foliar retention and activity, and more reproductive output. Grasses
decline in dominance because while their C4-type photosynthesis is enhanced with increased atmospheric CO2, their phenology is tightly coupled
to soil moisture and precipitation which decrease with climate change.
Overall primary productivity declines in the future scenario with a greater
proportion of photosynthate allocated to belowground structures. Although
fire fuels increase with climate change due to more shrub cover, fire frequency and return intervals decrease with less fine fuels contributed by
grasses. In addition, reduced productivity decreases average vegetation canopy cover, leading to patchier fuel distributions and lower probability of
fire spread. Spatial variation expressed in terms of both vegetation community and productivity decrease with climate change as increased shrub
dominance and reduced fire frequencies result in a more homogeneous
landscape.
WHITE, PETER S.,1,* MEGHAN MCKNIGHT1 and JOAN L. WALKER.2
1
University of North Carolina at Chapel Hill, Chapel Hill, North Carolina,
USA; 2 US Forest Service, Clemson University, Clemson, South Carolina,
USA. Endemism, hot spots of diversity, and ecosystems in the Southeast: the template on which climate change will act.
The Southeast has many narrow endemics, rapid geographic turnover, and
continental high points of diversity in a number of groups, including fish,
amphibians, turtles, freshwater mollusks, land snails, cave organisms, and
some groups of insects and plants. Rather than occurring in the same places
and ecosystems, however, the endemics and hot spots of diversity are widely scattered across the region. The distribution of ecosystems is also complex, with gradients of water and nutrient availability creating locally
changing compositions and structures. We compare the pattern of richness,
endemism, and globally rare species for different taxonomic groups in order
to better understand the vulnerabilty of this rich biological area to climate
change. For vascular plants, we show that the number of narrow endemics
decreases much more quickly northward than the decrease in total richness,
that endemism increases from west to east across the Southeast, and that
the average similarity is lower and the decay of similarity with distance is
higher across the Southeast than across the glaciated North. Glaciation has
evidentally left a long-lasting imprint on patterns of biological diversity in
eastern North America. These data suggest that conservation by ecosystem
type is disjunct from conservation of species diversity. Patterns in the the
Southeasts high levels of endemism suggest a unique and high degree of
vulnerability to climate change.
WHITE, SUSAN N.* and MERRYL ALBER. University of Georgia, Athens, GA. Will the marsh paradigm hold? Spartina distributions along
the length of the Altamaha River Estuary, GA.
In the Altamaha River estuary in Georgia, S. alterniflora and S. cynosuroides are the dominant plant species in salt and brackish marshes, respectively. The general paradigm for zonation in salt marshes is that stress
tolerance and competition combined drive vegetation patterns within a single marsh. However, it is unclear if the stress tolerance/competition para-

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357

digm can be applied along an estuarine gradient. Two field experiments

established in Spring 2001 examined factors that control S. alterniflora and
S. cynosuroides distribution along the Altamaha River estuary. Abiotic (salinity) and biotic (intra/interspecific competition) influences on Spartina
growth were investigated through reciprocal transplant studies coupled with
neighbor removal treatments (at salinity extremes) and experimental species removals (from a mid-salinity, mixed species area). Results from the
transplant experiments indicate that high salinities negatively affected S.
cynosuroides growth more than S. alterniflora (i.e., 20% S. cynosuroides
survival vs. 80% S. alterniflora survival, etc.). However, both plants
showed evidence of competitive effects. The presence of neighbors increased S. cynosuroides growth (i.e. avg. height with neighbors was
1.5(60.5) m vs. 0.85(60) m without neighbors in the high salinity zone,
1.45(60.73) m with neighbors vs. 0.97(60.59) m without neighbors in low
salinity zone) indicating the possibility of facilitation, whereas S. alterniflora performed poorly with neighbors present (i.e., avg. height with neighbors was 0.8(6 0.72) m vs. 1.2(60.75) m without neighbors in high salinity
zone, 0.3(6 0.45) m with neighbors vs. 0.69(6 0.14) m without neighbors
in low salinity zone), indicating a possible competitive effect. We interpret
these results to suggest that physiological tolerance to high salinity is the
primary control of species distribution but that competitive interactions are
important at lower salinities. These results provide support for the extension
of the stress tolerance/competition paradigm along the length of an estuary.
WHITLOW, LINDSAY.* Bowdoin College, Brunswick, ME. Historical
changes in native clam traits after crab invasion.
To determine whether native phenotypes change after invasion, native specimens collected over a range of dates before and after the invasion, and
over a range of locations, should be examined. In soft-sediment marine
ecosystems, many bivalve species burrow deeper in the sediment in an
attempt to avoid predators, and the deeper bivalves are less vulnerable to
many predators. The burrowing depths and siphon lengths of bivalves are
recorded in the pallial sinus length on the interior of the shell. In Maine,
USA, an introduced predator, the green crab, Carcinus maenas, spread
northward along the coast between 1900 and 2000 and contributed to declines in abundance of native soft-shell clams, Mya arenaria. Aspects of
shell morphology were quantified from clams collected before and after
the arrival of the crabs, and from multiple sites along the Maine coast.
Since the invasion increased the abundance of crab predators, shell morphologies were expected to reflect greater burrowing depths. Relative to
shell length, pallial sinus length, shell mass, and internal tissue scar length
all increased after green crab arrival, and the increases occurred earlier at
lower latitudes. Collections within more narrow latitudinal ranges (Casco
Bay and Mt. Desert Island) also indicate these morphological parameters
significantly increased after the crab invasion. These results suggest crabs
are the most likely factor responsible for these changes in clam shell morphology, rather than climatic conditions such as steadily rising seawater
temperatures. Pallial sinus length is correlated with burrowing depth and
siphon length, thus increased pallial sinus length indicates that the clams
may have burrowed deeper to evade predation or crabs may have consumed
more shallow clams. Shell mass relative to length increased after crab arrival, which may be due to slower growth rates of deeper burrowing clams
due to lower feeding efficiency.
WHITTAKER, KARA, TESSA B. FRANCIS,* JESSICA GRAYBILL,
APRIL MILLS and VIVEK SHANDAS. University of Washington, Urban
Ecology Program, Seattle, WA, USA. Patterns associated with the use
of best available science in critical area ordinances in western Washington State.
We investigated the factors affecting the use of Best Available Science in
ordinances protecting the critical areas of local jurisdictions in Washington
State. Recent state statutes mandate that the Critical Area Ordinances
(CAOs) should be updated and that these updates shall include best available science (BAS). We studied the CAO revision process for biological
critical areas in 9 counties and 30 cities in western Washington. The process
of incorporating scientific information, especially BAS, into public policy
is complex and opaque. We asked several questions regarding the inclusion
of BAS in policy: will the incorporation of BAS vary by jurisdiction and
if so, what are the factors inhibiting or aiding the use of BAS? Do the

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CAO update process and its outcomes vary by jurisdiction? What additional biophysical and socioeconomic factors are driving this policy process? Through this investigation, we aim to elucidate the process by which
policy makers use science in decision-making and thereby inform both
policy makers and scientists about the usability of science in applied settings.
WILLIAMS, ALISSA J.* and JOEL C. TREXLER. Florida International
University, Miami, FL. The effect of hydroperiod and productivity on
food-chain length in Everglades wetlands.
Theory predicts that food-chain length may shorten along gradients of disturbance or lengthen along gradients of increasing productivity; however,
in natural systems, these two environmental factors often vary simultaneously. We used stable isotopes to estimate food-chain length at 20 sites
in the Florida Everglades representing a range of productivity and hydroperiod. In the Everglades, hydroperiod gradients represent gradients of disturbance because drying events yield high mortality of fishes and macroinvertebrates. We characterized food webs at all sites during both the wet
and dry seasons. We recorded soil, floc, and periphyton TP, as well as
primary production of periphyton samples using light/dark bottle incubations as indicators of productivity. We analyzed 585 samples representing
basal, intermediate, and top trophic levels for d15N and d13C. Amphipods
and Seminole ramshorn snails were selected as the primary baseline detritivore and herbivore taxa. Eastern mosquitofish and riverine grass shrimp
represented intermediate trophic levels, and Florida gar were the top aquatic
carnivores at each study site. Our results indicate that more variation in
trophic position and food-chain length can be explained as a function of
joint productivity and disturbance descriptions than by single parameter
gradients. We observed marked variation in trophic relationships among
sites and between seasons. We also observed variation in d13C values of
primary consumers among our study sites, indicating spatial variation in
the isotopic value of carbon consumed at the base of the food web. After
correcting for spatial variation in isotopic signal, data revealed variation
within and among regions in both d13C and d15N. This indicates a possible
shift in the relative roles of detrital and algal carbon among sites. A comparison of intermediate consumers across sites, corrected for variation in
basal isotopic values, yielded little variation in d15N, but trophic position
of eastern mosquitofish did vary among sites.
WILLIAMS, CHRISTOPHER K.,1 TONY R. IVES1 and ROGER D. APPLEGATE.2 1 Department of Zoology, University of Wisconsin, Birge Hall,
430 Lincoln Drive, Madison, WI, USA; 2 Department of Wildlife and
Parks, 1830 Merchant Street, P.O. Box 1525, Emporia, KS, USA. Population dynamics across geographical ranges: Time-series analyses of
three small game species.
Little is known about how density-independent and density-dependent processes affecting population dynamics vary geographically across species
ranges. To address this problem for three vertebrate species (northern bobwhite, ring-necked pheasants, and Eastern cottontails), we examined longterm (1966-2001) seasonal (January, April, July, and October) time-series
data from the Kansas Rural Mail Carrier Survey, USA. All three species
have range boundaries in Kansas, with population abundances declining
towards the periphery of their ranges. We quantified the strengths of density-dependent and density-independent processes affecting the dynamics
of 10 populations of each species ranging from low to high mean abundance using first-order autoregressive models with measurement error. Peripheral populations with lower mean abundance tended to have greater
population variability. This pattern could potentially be explained by peripheral regions experiencing weaker density dependence or greater environmentally driven density-independent fluctuations in per capita population growth rates. In general, density dependence did not vary among geographic regions, although there was a trend for smaller, peripheral populations to exhibit stronger density dependence. Density-independent
variability in per capita population growth rates was higher in peripheral
populations. Furthermore, density-independent fluctuations in per capita
population growth rates were weakly correlated with temperature and precipitation, and were highest for the period October-January, identifying fall
as the period of greatest environmentally driven variability in population
dynamics. Per capita population growth rates fluctuated in moderate syn-

chrony among regions, especially for more abundant, non-peripheral populations in close geographical proximity. The strong density-dependent and
stronger density-independent processes in smaller, peripheral populations
suggest that the greater variability in peripheral populations densities are
caused by greater population sensitivity to environmental fluctuations. This
may make peripheral populations more likely to go extinct and leads to the
prediction that potential population declines will be accompanied by a contraction in their geographic ranges.
WILLIAMS, JOHN W.,1 ERIC SEABLOOM,1 DANIEL SLAYBACK,2
DAVID STOMS,1 SARAH SHAFER3 and PATRICK BARTLEIN.4 1 University of California Santa Barbara, Santa Barbara, CA; 2 University of
Maryland, College Park, MD; 3 US Geological Survey, Corwallis, OR;
4
University of Oregon, Eugene, OR. Have human activities altered productivity-diversity relationships in California?
Previous attempts to model the biogeographic factors of species richness
generally focus upon energetic, ecophysiological, and/or historical controls
on species distributions, ignoring the substantial impacts of humans on
species distributions and ecosystem functioning at regional to continental
scales. In this study we seek to understand how the relationship between
vascular plant richness, net primary productivity, and various abiotic indices of environmental favorability in California has been modified by
anthropogenic activities. The Californian vegetation is diverse (comprising
over 20% of US plant species), includes strong gradients in productivity
(.three orders of magnitude from the southeastern deserts to the coastal
mountains), and has been widely and variably impacted by human activities. Using plant species richness data from the CalFlora database, NPP
estimates for 1982-1999 from GLO-PEM, IPCC CRU 1961-1990 climate
data, StatsGo soil textural properties, and remotely sensed indices of land
cover conversion, we constructed and tested path models representing alternate hypotheses about a) the relationship between NPP, richness, and
exogenous environmental variables and b) the direct and indirect impacts
of human activity. The distribution of NPP and species richness was estimated for two alternate states representing pre-settlement and present-day
conditions; all other variables were assumed to be unchanged. Path models
that exclude anthropogenic influence are able to explain a high proportion
of the variance in both pre-settlement and present-day plant richness. However, additional variance in the present-day distribution of plant richness
corresponds to indices of land cover use.
WILLIAMS, KATHY S.* San Diego State University, kwilliams@
sunstroke.sdsu.edu, San Diego, CA. Using embedded assessment to enhance learning in college biology classes.
After years of teaching college biology, I remained frustrated at the poor
quality of answers I got on mid-term and final exams. Then, through Project
FIRST (Faculty Institutes for Reforming Science Teaching, funded by
NSF), I discovered the usefulness of embedded assessment to enhance
learning and communication skills in my biology classes. Now, in lieu of
traditional exams, I use small quizzes and writing activities in most class
meetings to learn what my students know and to help them improve their
understanding. Typically, immediately after the assignments are turned in,
I present a set of responses of different qualities that students evaluate
together. Most of the course grade comes from this embedded assessment,
rather than from a mid-term of final exam for which students often "cram."
To test the hypothesis that students have learned and retained the information as well as when they studied for exams, I have given unannounced
"recall exams," similar to mid-term exams I used in the past, in several
classes. Even though the "recall exams" were completely unexpected, student scores were typically a grade higher than when I taught the courses
in a more traditional manner. In addition, their writing skills were greatly
improved from previous classes. I will present outcomes from several college biology classes, and examples of rubrics I use that allow me to score
the frequent assignments efficiently. I also will discuss student attitudes
toward the frequent assessment methods.
WILLIAMS, KIMBERLYN,1,* BOB J. WILLIAMS2 and LAWRENCE J.
WESTRICK.1 1 California State University-San Bernardino, San Bernardino, CA; 2 Shasta County Cattlemens Association, Redding, CA. Interactions between oak regeneration and Rubus discolor, an invasive
blackberry, in Northern California.
Californias oak woodlands and savannahs have experienced a number of
changes over the past century or more. Low recruitment of blue oak (Quer-

cus douglasii) has been noted, and extensive research on oak regeneration
has focused on effects of grazing and browsing, fire suppression, and competition with introduced annual grasses. A second change in this system
has been the invasion of some areas by Rubus discolor (Himalayan blackberry) an invasive species forming dense thickets of canes. This invasion
has received less attention from ecologists, but more attention from cattle
ranchers who expend enormous efforts in trying to control its spread. Initial
observations of oak sapling distribution on a ranch in Northern California,
suggested that saplings were associated with blackberry thickets. To determine if blackberry thickets provided a favorable environment for oak seedling establishment, we planted 300 acorns of Quercus douglasii and 300
acorns of Quercus lobata in an oak savannah/berry thicket mosaic in December 2001. We planted one third of the acorns at the edge of thickets,
one third inside thickets (1 m from the edge), and one third in adjacent
grassland. The only live seedlings found the following spring were in the
grassland, and planting sites inside berry thickets showed signs of extensive
rodent activity. Given the rodent populations during the year of our study,
therefore, berry thickets did not provide a safe site for oak regeneration.
We speculate that the association of saplings with berry thickets may result
from protection of the rare seedling from browsing by large animals. However, those rare seedlings must have established during years with lower
rodent populations or in grassland before the blackberry thicket covered
the site.
WILLIAMS, NICHOLAS SG.,1,2,* MARK J. MCDONNELL1,2 and JOHN
W. MORGAN.3 1 Australian Research Centre for Urban Ecology, Royal
Botanic Gardens, Melbourne, Australia, Australia; 2 University of Melbourne, Melbourne, Australia, Australia; 3 Latrobe University, Melbourne,
Australia, Australia. The influence of plant species attributes on local
extinctions in urban and rural grasslands.
Native temperate grasslands in Australia are now extremely fragmented
and the persistence of many plant species depends on relict populations
confined to small, isolated remnants in urban and rural landscapes. To determine if there were differences between the persistence of plants in small
patches of native grassland in urban vs. rural landscapes in southern Victoria, we re-surveyed 31 grassland remnants that had undergone a comprehensive botanical survey between 1979 and 1990. A total of 330 (30%) of
the 1102 plant populations present in the 1980s were not relocated and
were presumed locally extinct. Rural patches lost fewer populations than
urban patches (26% vs 35%) Different suites of species were lost from
urban vs. rural grasslands, but species common in both landscapes had
consistently higher extinction rates in urban grasslands. Logistic regression
models were created to determine the plant attributes that influence the
probability that a population of a species will become locally extinct in
each landscape. Those species most likely to become locally extinct had
one or more of the following attributes: wind dispersed seed, growing buds
at the soil surface and rosette, partial rosette or mat-like growth form.
WILLIS, JONATHAN M.,1,2 MARK HESTER,2 KAREN MCKEE3 and
IRV MENDELSSOHN.4 1 Southeastern Louisiana University, Hammond,
LA; 2 University of New Orleans, New Orleans, LA; 3 National Wetlands
Research Center, Lafayette, LA; 4 Louisiana State University, Baton Rouge,
LA. Investigations of a coast-wide Spartina alterniflora marsh dieback
event: Preliminary results of environmental tolerance trials.
Spartina alterniflora marshes are a tremendously important component of
coastal ecosystems, providing benefits ranging from the dampening of
storm surges to habitat for larval marine species. Extensive dieback of
Louisiana S. alterniflora marshes was observed in 2001 toward the end of
a 3-year drought. Of great interest is that two other widespread plant species of Louisiana salt marshes, Juncus roemerianus and Avicennia germinans, did not demonstrate any large-scale dieback during this time and
remained green within stands of S. alterniflora stubble. An extensive, multifaceted approach to determining the specific causes of the S. alterniflora
dieback phenomenon has been undertaken by a team of Louisiana researchers. A major component of this effort is the elucidation of the tolerances
of S. alterniflora, J. roemerianus, and A. germinans to several environmental variables including salinity, moisture availability, pH, and interstitial
Al and Fe concentrations. Trials have focused on a series of single factor
experiments (e.g., salinity) that subject the three species to three treatments:

Abstracts

359

control (factor held at a constant non-lethal level), increased (factor increased until all species in have died), held (factor increased along with
increased treatment until stress detected, then held at this level). Results
from these trials are being utilized to design a multifactor experiment with
interacting variables most likely to have contributed to the sudden marsh
dieback event. Salinity trials indicate species tolerances to be A. germinans
. S. alterniflora . J. roemerianus. pH trials indicate species tolerances to
be J. roemerianus . S. alterniflora . A. germinans. Moisture availiability,
Al, and Fe trials are close to completion, at which point the multifactor
experiment will commence.

was an effect of density and a marginal effect of habitat quality when

survivorship of the entire target assemblage was considered as a whole. I
conclude that competitive equivalence in combination with variable settlement rates into a heterogeneous reef environment help explain how these
species coexist.

WILLSON, CYNTHIA J.,1,* HAFIZ MAHERALI,2 PAUL S. MANOS1

and ROBERT B. JACKSON.1 1 Duke University, Durham, North Carolina;
2
University of Guelph, Guelph, Ontario, Canada. Interspecific variation
in hydraulic architecture in Juniperus (Cupressaceae) species in a phylogenetic context.

In this study we investigate the impact of an invasive crayfish, Orconectes

rusticus, on the food web structure of lake littoral zones in northern Wisconsin. The rusty crayfish is invasive across the upper Midwest and northeastern United States, as well as into parts of Canada. In northern Wisconsin lakes, O. rusticus often dominates benthic biomass, reaching trap abundances from 2 - 20 times the abundance of native species. High densities
of rusty crayfish are negatively correlated with snail, macrophyte and macroinvertebrate abundances. O. rusticus obtains a size refuge from most
predatory fishes at carapace lengths . 35 mm. We hypothesized that, at
high densities, larger rusty crayfish represent the top of the near-shore benthic food web, forcing littoral fishes to switch to pelagically-derived food
resources. To test this hypothesis, we assessed trophic structure in 5 highdensity rusty crayfish lakes and 5 low-density rusty crayfish lakes in northern Wisconsin. We used stable isotopes (delta-13C and delta-15N) to measure
trophic position and prey source (littoral vs. pelagic) for a trophic range of
littoral fishes (smallmouth bass, bluegill, johnny darter, and yellow perch),
crayfish, macroinvertebrates, snails and mussels. We also assessed ambient
snail and macroinvertebrate populations using a benthic suction sampler in
both cobble and sand habitats. Initial results suggest significantly fewer
macroinvertebrates and snails in lakes with high densities of rusty crayfish;
we expect that these differences will be reflected in strong shifts in trophic
structure throughout the littoral food web.

A range of resistance to drought-induced xylem cavitation exists among

species. In interspecific comparisons, correlations among xylem cavitation
and other hydraulic traits can identify potentially adaptive relationships or
tradeoffs. We constructed a molecular phylogeny for 14 Juniperus species
to investigate adaptive hypotheses relating interspecific variation in hydraulic architectural traits to each other and to climatic variables and to
examine tradeoffs between cavitation resistance and such variables as hydraulic conductivity and wood density. Stem and shallow root (n54-6)
samples were collected in the summers of 2000, 2001, and 2002. Vulnerability curves were constructed using the centrifugal force method to generate cavitation at known tensions. The P50 value is a useful comparative
value that represents the tension inducing a 50% loss in hydraulic conductivity. A phylogeny based on sequence variation of the internal transcribed
spacer (ITS) of the 18S-26S nuclear ribosomal DNA (rDNA) repeat supports the subdivision of Juniperus section Sabina into species with and
without serrate leaf margins. The serrate-margin group, which is roughly
composed of the junipers in the western U. S., was more cavitation-resistant, with a mean stem P50 of -10.3 MPa and mean annual precipitation
(MAP) approximately half that of the non-serrate group. The non-serratemargin group, which is roughly composed of the junipers in the eastern U.
S. and the Caribbean, was less cavitation-resistant, with a mean stem P50
of -7.2 MPa. Across species, the P50 of both stems and roots showed a
significant linear relationship with MAP and a moisture index. Exploring
the effects of environment and phylogenetic history on interspecific variation in hydraulic architecture in Juniperus species offers insight into adaptation, tradeoffs, and the factors important in determining their distributions.
WILSON, JACQUELINE A.* University of Florida, jwilson@zoo.ufl.edu,
Gainesville, FL. Competitive ability of two marine gobies: How do they
coexist?
Historically, competition theory has emphasized the importance of niche
divergence among co-occurring species. More recent theoretical work, in
contrast, has emphasized the importance of similarity as a means to promote coexistence. Central to both viewpoints is the relative competitive
ability of competing species. Competitive ability can be decomposed into
competitive effect (ability to suppress other individuals) and competitive
response (ability to avoid being suppressed). In this study I investigated
the competitive effect and response of two species of marine gobies (Gobiosoma evelynae and G. prochilos) using a target/neighbor design in which
I varied the density and age-class of neighbors and monitored survival of
targets. I also conducted the experiment along a gradient of habitats differing in quality. Targets consisted of three G. evelynae and three G. prochilos settlers. Thus, every environment was experienced by settlers of both
species. These two species were competitively equivalent; neither species
nor age-class group differed significantly in their response to, or their effect
upon, one another. I also modeled survival of settlers of both species using
different models built on the Weibull distribution that sequentially incorporated the effects of species, age-class, density and habitat quality. I obtained maximum likelihood estimates of mean survival for each species
and selected the best-fitting model using Akaike Information Criteria.
When survival of the two species in the target assemblages was considered
separately, there was no effect of density or habitat quality. However, there

360

Abstracts

WILSON, KAREN A.,1,* THEODORE V. WILLIS2 and M. JAKE VANDER ZANDEN.2 1 Carleton College, Northfield, MN; 2 Center for Limnology, University of Wisconsin-Madison, Madison, WI. Impacts of invasive
crayfish on trophic structure in lake littoral zones.

WIMBERLY, MICHAEL C.* Warnell School of Forest Resources, University of Georgia, Athens, GA. Simulating ecological responses to static
versus dynamic landscape patterns.
The discipline of landscape ecology has traditionally emphasized the effects
of static spatial patterns on species dynamics. In many systems, such as
age-structured mosaics of forest habitats, landscape patterns themselves are
intrinsically dynamic. Succession inevitably changes microhabitat characteristics within forest patches, and disturbance continually reinitiates succession across the landscape. Does the influence of spatial pattern on ecological processes differ in static versus dynamic landscapes? To address
this question we incorporated a generalized metapopulation model into the
Landscape Age-Class Dynamics Simulator (LADS) to simulate the responses of hypothetical species to random landscape patterns. Static landscape patterns with a given proportion of habitat (p) were compared with
dynamic patterns in which the expected proportion of habitat was equal to
p. Values of p ranged from 0.05 to 0.95 at intervals of 0.05. Early-successional and late-successional habitat responses were tested. Species occupancy in both static and dynamic landscapes exhibited threshold responses
to changes in p. The location of these thresholds was sensitive to both
species characteristics and initial landscape conditions. When forest management strategies maintain habitat amounts near these thresholds, certain
species and ecological processes may be sensitive to relatively small shifts
in the rate of disturbance. In most cases, species occupancy was lower in
the dynamic than the static landscapes. However, when p was close to the
critical threshold, species occupancy was sometimes actually higher in the
dynamic landscapes. In these cases, patch dynamics maintain higher connectivity over space and time than is present in the static landscapes.
WIMP, GINA M.,* WILLIAM P. YOUNG, SCOTT A. WOOLBRIGHT,
GREG D. MARTINSEN, PAUL S. KEIM and THOMAS G. WHITHAM.
Northern Arizona University, Flagstaff, Arizona. Host plant genetic diversity affects arthropod diversity at multiple scales.
We examined the hypothesis that genetic diversity in a dominant tree has
higher-level effects on the structure and diversity of the dependent arthropod community. Plant hybridizing systems are ideal systems in which we

can test these predictions because they represent some of the most genetically diverse systems where gene flow is occurring. Hybridizing systems
therefore provide an important step between inter- and intraspecific host
plant diversity in determining the level at which arthropod communities
might be responding to genetic differences among their host plants. We
chose a hybrid cottonwood system (Populus angustifolia x P. fremontii),
and used clones of known genetic composition in a common garden to
address individual tree-level effects, combined with stand-level comparisons in the wild to address landscape-level effects of host plant genetic
diversity on arthropod communities. In a common garden analysis of 66
different arthropod species, we found that genetic diversity among different
cottonwood species and hybrid types affected the structure of arthropod
communities via changes in arthropod composition. We therefore predicted
that changes in arthropod composition among cottonwood species and hybrid types would lead to the greatest levels of arthropod diversity where
cottonwood genetic diversity was also greatest. When we tested this prediction using an analysis of 11 different cottonwood stands and 210 arthropod species, our results supported this prediction. We found a significant, positive relationship between cottonwood stand genetic diversity and
arthropod diversity such that genetic diversity among cottonwood stands
accounted for 59% of the variation in arthropod diversity. Because these
findings demonstrate the community-level consequences of genetic diversity, they argue for the consideration of genetic-based approaches to the
theory of community organization and conservation biology.
WINFREE, RACHAEL.* Department of Ecology and Evolutionary Biology, Princeton University, Princeton, NJ, USA. High offspring survival
in a novel habitat for an invasive species, the brown-headed cowbird.
Species with a life history that balances high fecundity with low offspring
survival may be predisposed to invading areas outside their native range,
if the factor(s) limiting offspring survival are absent in the new environment. This study measured egg-to-fledgling survival for the brown-headed
cowbird (Molothrus ater), an invasive, fecund brood parasite, in two habitats. The first habitat, old fields, contains host species that have probably
co-existed with the cowbird for much of its 0.5 million year history in
North America. The second, mature deciduous forest, is an evolutionarily
novel habitat for cowbirds (300 years). The study was done in the Shawnee
National Forest, southern IL and data came from 748 host nests in two
landscapes. Cowbird egg-to-fledging success was higher with forest hosts
for 15/15 species in landscape 1 and 7/8 species in landscape 2. Intrinsic
reproductive rate, R0, for cowbirds breeding in field habitat was 0.17
0.19, whereas in the forest R0 was 0.61 2.38. This suggests that increased
offspring survival in the novel forest habitat contributed to the invasion
success of the cowbird.

suggest, however, that this slight difference may have been sufficient to
shift the outcome of plant-microbial and plant-plant competition for nutrients in favor of the fast-growing annual. Contrary to expectation, exotic
earthworm activity does not appear to accelerate nutrient turnover, but may
influence plant species diversity by enhancing growth of fast-growing annuals relative to slower-growing forbs and grasses. Thus, functional differences between native and exotic earthworm species may be an important
component in the restoration and conservation of native grassland ecosystems.

WINSOR, JAMES A.,1,* BRADLEY D. LEYSHON,2 STEVEN E. TRAVERS1 and ANDREW G. STEPHENSON.1 1 The Pennsylvania State University-Altoona, Altoona, PA, USA; 2 The Pennsylvania State UniversityUniversity Park, University Park, PA, USA. Herbivores, herbivore-vectored disease, and inbreeding in Cucurbita pepo ssp texana.

Preliminary studies indicated that inbred plants experience higher populations of herbivores and greater herbivore damage than do outbred plants.
We wished to determine the relationship among the level of inbreeding,
incidence of herbivore infestation, herbivore damage, and infection by herbivore-vectored disease. A garden experiment was conducted using inbred
and outbred plants in which herbivores were excluded from half the plants
(by spraying with a commercial pyrethroid insectide) while the remaining
plants were exposed to herbivores (unsprayed). We recorded presence of
herbivores, incidence of herbivore-vectored disease (e.g., cucumber mosaic
virus, bacterial wilt), and a suite of measures of male and female plant
fitness. The spraying treatment significantly reduced aphid infestation
(p,0.001), damage by diabroticite beetles (p,0.001), and infection by mosaic viruses (p,0.001) in both inbred and outbred plants. Inbred plants
were more likely to sustain beetle damage (p,0.05) but were no more
likely to be infested with aphids (p.0.7). However, inbred plants were
more likely to be infected by viruses, whether or not they were sprayed
(p,0.0001). Plants from which herbivores were excluded by spraying produced more staminate flowers (p,0.0001), faster growing pollen tubes in
vitro (p,0.005), more pistillate flowers (p,0.0001), and larger fruits
(p,0.004). Similar trends were observed in comparisons between outbred
and inbred plants. Significant correlations were observed between virus
infestation and pollen tube growth rate (r25-0.42, p,0.0001), fruit number
per plant (r25-0.24, p,0.0001), and fruit volume (r25-0.15, p,0.015).
These results demonstrate that a plants probability of exposure and its
resistance to disease are affected by its level of inbreeding.

WINSOME, THAIS,1,2,* WILLIAM R. HORWATH,1 LYNN EPSTEIN1

and PAUL F. HENDRIX.2 1 University of California, Davis, CA; 2 University of Georgia, Athens, GA. Functional differences between exotic and
native earthworm species in a California grassland.

WINTERS, MONICA S.1,* and DAVID LIPSON.2 1 San Diego State University, San Diego, CA; 2 San Diego State University, San Diego, CA. The
effects of an invasive plant community on the coastal sage scrub soil
microbial community.

Exotic species are predominant in earthworm communities in managed

grasslands in California, but their relative role in ecosystem function remains unknown. Based on laboratory studies, we hypothesized that a common exotic species, Aporrectodea trapezoides (EX), would enhance C and
N mineralization rates and accelerate their turnover relative to a native,
Argilophilus marmoratus (NA). We applied 13C15N-labeled grass litter to
plots with earthworm treatments consisting of EX, NA, a 50/50 EX/NA
mixture, and a control with no earthworms (CO). The plots were seeded
with Lolium multiflorum, a fast-growing annual grass. We then followed
plant growth and turnover of litter-derived C and N through soil pools and
plant biomass. The EX and EX/NA treatments developed Lolium monocultures due to faster and more complete germination, while plant species
diversity was greater in NA and CO treatments. Relative to the control,
only EX increased total plant biomass and plant C and N, but all 3 earthworm treatments increased the percent recovery of litter-derived N in plant
biomass and followed the pattern EX 5 EX/NA . NA . CO (F519,
P,0.001). Shoot C-N ratio was significantly greater in CO and EX compared to NA and EX/NA. In contrast, there were few differences between
treatments in soil C and N pools, beyond a slight decrease in EX in microbial biomass and in the percent recovery of litter-derived N. Our results

Previous studies suggest the division between grassland and shrubland has
several causes involving herbivory, allelopathic responses, and climate
(Mooney 1988). The differences between grassland soils and coastal sage
scrub (CSS) soils from this study indicate that the soil microbial community
may have a role as well. In this study, soils were tested from sites covered
with the predominant coastal sage scrub shrub, Artemisia californica with
varying levels of invasive non-native grasses. Different enzyme analyses
(cellulase, ligninase, protease) and substrate-induced respiration (SIR) analyses of different substrates (glycine, glucose, salicylate) were used to elucidate a difference in the structure of the microbial community. An analysis
of soil characteristics (pH, soil moisture content, organic matter, and microbial biomass) was made to determine how the amount of invasive plants
directly affects the soil. A higher pH was found under bare ground and
non-native grasses than under A. californica. With increases in invasive
non-native grasses, a decrease in soil moisture content and organic matter
was observed. Results of the enzyme and SIR analyses concur with a difference in microbial community structure between grassland and shrubland
communities. These findings indicate changes in the structure of the CSS
soil microbial community with increasing levels of invasive grasses.

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361

WIPF, SONJA,1,2,3 CHRISTA P.H. MULDER,2 VERONIKA STOECKLI1

and BERNHARD SCHMID.3 1 Swiss Federal Institute for Snow and Avalanche Research SLF, Davos, Switzerland; 2 University of Alaska, Fairbanks, AK; 3 Insitute for Environmental Sciences, University of, Zurich,
Switzerland. Winter climate change in snow-rich environments: Reactions of arctic and alpine plant communities.
Arctic and alpine regions are particularly affected by global climate change.
Winter climate has been especially subject to major changes: in the two to
three last decades, the measured temperature rise in the American Arctic
and the European Alps was greatest during winter, and the snow cover
extent decreased in many regions. However, climate change research mainly focuses on the effects of increased summer temperatures, CO2 concentrations and nutrient availability. Previous studies on snow-vegetation interactions suggest that changes in snow cover duration (which determines
the start and length of the growing season) and changes in snow depth and
composition (which define the subnivean temperatures and the spring water
and nutrient input) have differential effects on the phenology, the population dynamics and the composition of arctic and alpine plant communities.
We (1) experimentally disentangle the two factors: duration of snow cover
and snow depth, (2) quantify their effects on phenology, population biology
of single species and community composition in an arctic and an alpine
study site, and (3) illuminate the mechanisms which may lead from changes
in phenology to changes in community structure and composition. While
an advanced snow-melt generally results in an advanced vegetation development, our results of snow manipulation experiments in alpine and subarctic habitats reveal that increased temperature conductance of the snow
cover and subsequent decreased subnivean temperatures can delay the development of early flowering plant species. In our experiments, we control
the interactions between start of the growing season and winter temperature
regime and explore their effect on phenology, productivity and reproduction
of tundra species.
WITHINGTON, CYNTHIA L.* and ROBERT L. SANFORD. University
of Denver, cwithing@du.edu, Denver, CO, USA. Nutrient uptake and
decomposition in the forest-alpine tundra ecotone of the Colorado
Front Range.
Nitrogen (N) and phosphorus (P) availabilities strongly influence alpine
plant productivity, yet much uncertainty exists about the nutrient status of
trees in the forest-alpine tundra ecotone. We hypothesize that nutrient availability decreases with elevation in this ecotone, thus contributing to stunted
tree growth (krummholz) and the inability for trees to grow above treeline.
Engelmann spruce (Picea engelmannii) root nutrient (N and P) contents
were determined and root N and P uptake bioassays were performed for
use as a comparative index of nutrient availability in surface soils at Mt.
Goliath, Rogers Peak, and two transects at Niwot Ridge, Colorado. Contrary to our hypothesis, we found no significant differences in P content
between krummholz (1.16 mg P g root-1 6 0.06 SE) and closed forest (1.16
mg P g root-1 6 0.09 SE) roots. Similarly, there were no significant differences in P uptake between krummholz (0.483 mg P g root-1 15 min.-1 6
0.091 SE) and closed forest (0.471 mg P g root-1 15 min.-1 6 0.063 SE)
roots. These results are surprising because previous studies of Engelmann
spruce needles have shown less phosphorus in krummholz than upright
trees. Nitrogen content in spruce roots and N uptake results are compared
as well. Because nutrient availability is believed to vary with organic turnover and decomposition, these results are interpreted in the context of a
year-long decomposition study with N and P treatment effects using an
experimental design in the field with cotton strip assays.
WITMAN, JON D.,* SALVATORE J. GENOVESE, JOHN F. BRUNO and
ANDREW H. ALTIERI. Marine masting: episodic prey recruitment
and the synchrony of community dynamics on large spatial scales.
A massive subtidal recruitment of the mussel, Mytilus edulis, across the
SW Gulf of Maine (GOM), USA provided an opportunity to test the hypothesis that the episodic increase in food resources had a bottom up effect
on mussel consumers. Surveys across a 120 km marine landscape revealed
that juvenile mussels covered 33 to 91 % of the bottom (8 to 12 m depth)
from October 1995 to June 1996. The cover of mussel prey explained
significant variation in the density and biomass of mussel predators (sea

362

Abstracts

stars, rock crabs) at 16 to 17 sites 10, 13 and 23 months after mussel

recruitment, consistent with hypothesized bottom up effects. Coupling between bottom up and top down control occurred 12 to 14 months after
mussel recruitment when predation by sea stars, crabs and sea urchins
eliminated extensive beds of juvenile mussels across the region. The elimination of mussel prey triggered cannibalism in the sea star Asterias vulgaris, which contributed to density-dependent sea star population declines
from 1996 to 1997. An assumption of the consumer reproduction and recruitment response, that larvae produced by adult stocks of consumers
could be retained in the study region, was supported by investigations of
the potential larval dispersal of sea stars and crabs in a simulated 3D flow
field of the SW GOM. In this study, massive prey recruitment stimulated
coupling between bottom up and top down forces, resulting in broad scale
changes in local consumer populations that persisted after the prey recruitment signal was removed, underscoring the importance of episodic events
in marine community dynamics. A review of long term data suggests that
episodic, large recruitment of prey species is an important feature of benthic
marine ecosystems. Like the phenomenon of mast seeding in terrestrial
ecosystems, it has the potential to synchronize population and community
dynamics across broad spatial scales.
WOJCIK, NATHAN J.,1,* ROBERT L. SANFORD,1 THOMAS W.
MCGINNIS,2 JON E. KEELEY,2 MATT BROOKS3 and JAYNE BELNAP.4
1
University of Denver, Denver, CO; 2 U.S. Geological Survey, Biological
Resources Discipline, Western Ecological Research Center, Sequoia-Kings
Canyon National Park, Three Rivers, CA; 3 U.S. Geological Survey, Biological Resources Discipline, Western Ecological Research Center, Las Vegas Field Station, Las Vegas, NV; 4 U.S. Geological Survey, Biological
Resources Discipline, Forest and Rangeland Ecosystem Science Center,
Moab, UT. Effects of fire on soil nutrients following cheatgrass invasion
of Sierra Nevada yellow pine forests.
Prescribed fire has greatly increased the invasion of cheatgrass (Bromus
tectorum L.) in old growth yellow pine forest of Kings Canyon National
Park, California. The potential changes in fuels and soil chemistry caused
by cheatgrass invasion have important implications for management. By
manipulating several variables in 5x5 m forest test plots (n5420), we determine the aspects of fire that have the most effect on cheatgrass, native
plants and soil chemistry. This paper focuses on the changes in soil phosphorus (P) and nitrogen (N). Late fall burns in 2001, with light fuel loads
and high relative humidity, caused non-significant changes in most forms
of soil P and soil N. Soil temperatures 5 cm belowground in the 2001 burn
reached a mean of only 188C. For the summer and fall 2002 burns we
added fine fuels in the form of a 5 cm-deep layer of pine needles. Fuel
additions caused a significantly greater heat flux belowground, and this
increase in soil temperature caused significant changes in both NH4-N and
NO3-N soil levels (p , 0.05). NH4-N in fuel treated plots increased to 1.3
mg N/g soil post burn, in comparison to control plots with 0.2 mg N/g soil
(2 cm depth). NO3-N decreased significantly after burning for both fuel
treated and for control plots (0.47 mg N/g soil and 0.35 mg N/g soil respectively), with the largest decrease in fuel treated plots. We predict that
changes in plant available soil P and N levels increase following fires that
heat soils to moderate temperatures (100 3008C), ultimately increasing
the productivity of invasive cheatgrass. We compare our predictions with
additional samples from the 2003 growing-season.
WOJDAK, JEREMY M.1,2,* 1 Michigan State University, East Lansing, MI;
2
W. K. Kellogg Biological Station, Hickory Corners, MI. Species richness
effects on ecosystems depend on ecological context: An experiment with
aquatic grazers.
Species richness may influence the functioning of ecosystems, but in spite
of intense interest in the subject ecologists know little about where and
when to expect particularly strong or weak richness effects. Ecological
context (biotic and abiotic conditions) determines the abundance and behavior of species, and should in part determine the degree to which species
partition resources or niche space. If functional complementarity among a
guild of species depends on ecological context, so should the effects of
species richness. I used a mesocosm experiment to address the hypothesis
that species richness effects on system properties will depend on ecological
context. A manipulation of aquatic snail species richness and composition

(each of three species in every combination of 1, 2, and 3 species) was

crossed with a factorial manipulation of nutrient availability (ambient/enriched) and invertebrate predator presence/absence. Thus I could look for
species richness effects on system properties (i.e. snail, periphyton, epiphyton, and macrophyte biomass, sedimentation, system productivity and
respiration) in four unique environmental contexts. Species rich mesocosms
had higher snail biomass, regardless of context. Overall, epiphyton and
periphyton were less abundant at higher levels of snail richness; however,
the presence of predators ameliorated the richness effect on epiphyton and
richness effects on periphyton were only apparent at high productivity.
Macrophyte biomass and whole system respiration also depended on richness and context, but in complex interactive ways. Some ecologists remain
skeptical that the effect of species richness on ecosystem properties can be
of a biologically relevant magnitude. This experiment demonstrates that for
many response variables the magnitude of species richness effects can be
equal to or greater than those of predator and nutrient manipulations.
WOLF, AMELIA A.,1,* PATRICK MEGONIGAL,1 BRUCE A. HUNGATE,2 FRANK P. DAY3 and BERT G. DRAKE.1 1 Smithsonian Environmental Research Center, Edgewater, MD; 2 Northern Arizona University,
Flagstaff, AZ; 3 Old Dominion University, Norfolk, VA. Effects of elevated CO2 on microbial respiration in a Florida scrub oak ecosystem.
We measured microbial respiration over soil depth profiles in order to examine the effects of elevated CO2 on the soil labile C pool in a Florida
scrub oak ecosystem. The study consisted of replicated (n58) open-top
chambers exposed to ambient or twice-ambient atmospheric [CO2], as well
as non-chambered control plots. Triplicate 1-m deep cores were taken from
each chambered plot and 1 core was taken from each non-chambered plot.
Microbial respiration was measured at 4 depth intervals in a laboratory
incubation over a 32-week period. Microbial respiration at the soil surface
(0-10 cm) was consistently higher by 7-19% in the ambient chambers than
in the elevated chambers, though this difference was not statistically significant. Previous studies have either observed a significant stimulation of
microbial respiration by elevated CO2, or a non-significant trend for such
an effect. At depths of 10-30 cm and 30-60 cm, elevated CO2 had no effect;
however, when a spodic horizon (60-90 cm) was present, there was a trend
of higher rates of microbial respiration in the elevated CO2 plots. Soils at
all depths from the elevated chambers were significantly more depleted in
13
C-CO2 than soils from ambient chambers and non-chambered plots, reflecting the highly depleted 13C content of the CO2 added to the elevated
chambers. Collectively, these data lead us to speculate that elevated CO2
has stimulated the transport of labile soil carbon from the soil surface to
deeper horizons, perhaps by reducing transpiration.
WOLF, EVAN C.* and DAVID J. COOPER. Colorado State University,
Fort Collins, CO. Climate- and beaver-induced hydrologic variation of
small streams: Implications for willow persistence in Yellowstone National Park.
Causes for a significant shift in riparian community composition, as indicated by a decline in willow abundance and stature in northern Yellowstone
National Park since 1900, are poorly understood. The goals of this study
are to investigate causes of willow decline by determining 1) the influence
of fluvial processes on willow establishment along small streams and 2)
the historic range of variability in hydrologic regime as controlled by climate and beaver activity. Establishment dates for a stratified random sample
of 80 excavated riparian willows show fairly continuous recruitment from
1920 to present with two peaks in abundance and one abrupt change in the
elevation of establishment. The first broad peak is from 1930 to 1950,
when willows established up to 2.6 m above the current thalweg, possibly
on sediments from abandoned beaver ponds. The second peak in abundance
is a significant cohort of willows that established in 1989, across an elevation range of 0.7 m to 2.4 m above the current thalweg and only at sites
where burning occurred in 1988, suggesting a direct influence of fire. Willow establishment after 1974 (1989 cohort excluded) is restricted to surfaces ,0.8 m above the current thalweg. The shift in the elevation of
establishment may indicate a stream downcutting event and associated drop
in water table beginning 1974. Preliminary dendroclimatic reconstructions and floodplain sediment analyses indicate that the hydrologic regime
during the 20th century is consistent with that of the past 500 years.

WOLF, JOY J.* University of Wisconsin - Parkside, Kenosha, WI, USA.

A comparison of restoration strategies in oak openings.
In seeking to understand the patterns we see in the largely human-altered
landscapes of today, it is often difficult to separate the natural processes
from those shaped by recent human activity. Although oak openings are
increasingly rare in Wisconsin, little is known in this plant community.
Age structure, species distribution and understory development have not
been fully determined, especially in disturbed areas. This study is based
on assessing the effects of restoration techniques for maximum efficiency.
Currently, varying levels of prescribed fire, mowing, tree removal, and
replanting native species are used in oak savanna restoration, yet monitoring these manipulations is limited. Dendrochronology, soil analysis, and
spatial field data collection were used to compare oak savanna structure in
varying degrees of management. Results show a shift in the age structure
from a bimodally aged young cohort in the most managed site to even but
higher aged structure in the unmanaged site. The point pattern analyses
show that Quercus trees, saplings and seedlings in unmanaged were not
clumped in 5m quadrats (n560, F51.09). However, managed sites were
clumped (n580-241, F51.40-3.0, p50.05). These data may indicate mortality rate in unmanaged areas. Q. macrocarpa dominated all diameter classes. Point center quarter and grid data show smaller, even sizes in less
managed sites, and uneven sizes with more restoration strategies. Although
Q. macrocarpa had the highest important value in managed sites (82 and
116), dogwood (97 compared to 35 for oak) had highest importance in the
unmanaged site. Basal area was higher in oak than other species for all
sites, but black cherry had values 60% of oak in the unmanaged site. These
results may reflect other woody undergrowth competing for resources. At
present, the species composition in the managed site reflects an oak savanna
assemblage; however, the species composition in the unmanaged site resembles a woodland assemblage. This study provides insight into how altered disturbance regimes can influence oak distribution, implications for
exotic invasion control and native species survival and contributes to the
future decision-making by land use managers in Wisconsin natural areas
and midwestern oak savannas.
WOLF, JULIE,1,* RAMESH MURTHY2 and GREG BARRON-GAFFORD.2 1 Northern Arizona University, Flagstaff, AZ, USA; 2 Columbia
Universitys Biosphere 2 Center, Oracle, AZ, USA. Elevated CO2 increases sporocarp production and intraradicle sporulation of one AMF species at Biosphere 2.
We quantified the growth of arbuscular mycorrhizal fungi (AMF) within
the Intensive Forestry Mesocosm (IFM) at Biosphere 2 (Oracle, AZ, USA).
The IFM is permanently partitioned into three bays, within which atmospheric carbon dioxide (CO2) has been maintained at concentrations of 420,
800, or 1200 parts per million since 2000. From 2000 to 2002, ninety-four
cottonwoods of similar genotype and age (all started from cuttings in 1998)
were grown under the CO2 concentrations within the three bays. In 2002,
multiple soil and root samples were taken from each bay and combined,
so that one composite sample represented a thorough spatial sampling of
the area within each bay. Spores of the same six species of AMF were
present in all three bays. Spore abundances of five of these species were
similar in all three bays, but the sixth species (an unidentified Glomus
species, producing Sclerocystis-type sporocarps), was increasingly abundant under increased CO2 concentrations (9, 11, and 16 sporocarps per 100
g soil under 420, 800, and 1200 parts per million, respectively). The size
of the sporocarps did not vary with CO2 treatments. This pattern corroborates the findings of the two other studies in which AMF spore communities were quantified under elevated CO2; in both, spores of certain species
within the genus Glomus were more abundant under high CO2 while the
other species present did not change in abundance. A qualitative change in
sporulation was also observed under the elevated CO2 treatments at Biosphere 2. Sporocarps of the above-mentioned Glomus species, which are
normally found in bulk soil, were also observed inside live roots growing
under elevated CO2 (0, 0.25, and 2.5 intraradicle sporocarps per 100 g
soil at 420, 800, and 1200 parts per million, respectively). There is no
previous documentation of sporocarps of this type developing intraradically. These quantitative and qualitative changes in sporulation provide further
evidence that elevated CO2 has important species-specific effects on arbuscular mycorrhizal fungi. These effects may impact plant and soil communities.

Abstracts

363

WOLF, MOSHEH,1,* JORGE SALAZAR-BRAVO2 and MIKE

FRGGENS.3 1 University of Illinois, Urbana-Champaign, Illinois; 2 Texas
Tech, Lubbock, Texas; 3 University of New Mexico, Albuquerque, New
Mexico. Morphological divergence in populations of the plains pocket
mouse Perognathus flavescens in adjacent habitats.
Speciation has historically been thought to occur mostly in isolated populations; however, a number of recent studies of tropical birds and reptiles
have shown that morphological divergence and speciation may occur in
adjacent habitats, supporting a model of parapatric speciation. Small mammal species may also show morphological divergence and speciation in
adjacent habitats, because of the scale of their home ranges and movements.
To test this, we studied a common heteromyid rodent, Perognathus flavescens, in two adjacent habitats within the Sevilleta LTER site in central New
Mexico: plains sand scrub (PSS), and chihuahuan desert scrub (CDS). We
tested the hypotheses that morphological differences exist between adult
populations of this rodent in these habitats. We found morphological differences on a number of levels. Mice from the PSS site had significantly
larger ears, tails, and feet than mice from CDS. We also observed differences in sexual dimorphism between sites. Males from CDS had larger ears
and tails than females, while no such difference was observed in PSS. Mice
from the CDS site also showed sexual dimorphism in allometry: females
showed correlations between body length and foot, tail, and ear lengths,
while males did not. In contrast, in mice from the PSS site these allometric
measurements did not differ between sexes. These morphological differences occur in populations separated by less than 5 km, separated by no
major barriers, and are most likely related to habitat differences alone,
because external variables, such as precipitation, temperature, or exposure
cannot be invoked as determinant factors. We are in the process of investigating differences in the genetic make-up of these populations; however,
preliminary findings indicate that parapatric speciation may be as pervasive
in small desert rodents as it is in tropical birds and reptiles.
WOLFE-BELLIN, KELLY S.,* JIN-SHENG HE and F. A. BAZZAZ. Harvard University, Cambridge, MA. Nonlinear responses of tree seedlings
to gradients of superambient air temperature and atmospheric CO2.
Increasing global air temperature and atmospheric CO2 concentration are
expected to have considerable impact on the function of forest ecosystems,
yet there is great uncertainty in the future predictions of each. Given the
sizeable range in these predictions, it is important to understand how forest
ecosystems may respond to a range of air temperatures and CO2 concentrations. We conducted a study to investigate how gradients of both elevated
air temperature and atmospheric CO2 affect seedling recruitment of three
species that co-occur as dominant canopy trees in northeastern hardwood
forests. Seedlings of gray birch, red oak, and red maple were exposed to
gradients of either elevated air temperature or a combination of elevated
air temperature and atmospheric CO2 for one growing season. The environmental gradients were created in greenhouse tunnels. All seedlings were
exposed to an air temperature gradient that ranged from ambient to approximately 38C above ambient during daylight hours. Additionally, half
the seedlings were exposed to a CO2 gradient that ranged from 370 to 810
ppm. Nonlinear responses to the gradients were evident at the assemblage
and species-specific levels. Total biomass at the end of the first growing
season exhibited little response to the temperature gradient alone, but exhibited a strong curvilinear (parabolic, concave-side down) response to the
combination of elevated temperature and CO2. The biomass of individual
species exhibited a range of responses to the temperature gradient alone,
while each species exhibited the same curvilinear response to the temperature and CO2 gradients evident in the total biomass response. These results
demonstrate that tree seedling recruitment may exhibit important nonlinear
responses to gradients of both environmental factors, and that the combination of elevated air temperature and atmospheric CO2 to which forests
may be exposed in the future should be considered when conducting experiments designed to predict ecosystem responses to global change.
WOLOSIN, MICHAEL S.,* JAMES S. CLARK, SHANNON LADEAU
and MICHAEL C. DIETZE. Duke University, Durham, NC. Canopy area,
growth, and fecundity in a mixed-hardwood forest.
Forest simulators predict community dynamics based on models of speciesspecific growth, fecundity, and mortality rates that vary with tree size. In

364

Abstracts

most, neighbors interact only through exploitative competition for light.

While several field studies have examined the changes in growth and mortality rates with resource availability for seedlings and saplings, the demographic responses and trophic trade-offs of larger trees are less well
known. These relationships are explored using a combination of remote
sensing and field data from four hectares of mixed pine-hardwood forest
in the North Carolina Piedmont. High-resolution aerial videography and
subsequent three-dimensional canopy reconstructions provide measurements of height and exposed canopy area (ECA) of individual trees at a
level of detail impossible from the ground. Additional field measurements
of total canopy area (TCA), diameter at breast height (dbh), and dbh growth
have been made for each tree over 20 cm dbh of several hardwood species,
including Acer rubrum,, Carya spp., Liquidambar styraciflua, Liriodendron
tulipifera, and Quercus alba. Seed trap data collected over three years and
inverse modeling of dispersal provide fecundity estimates of individual
trees. Regression analyses are used to estimate allometries between canopy
measurements, height, and diameter for each species. Initial results show
that the relationship between canopy area (a measure of light availability
and hence photosynthesis) and growth and fecundity do vary between species, providing evidence that large trees do experience trade-offs. Canopy
area is generally more predictive of growth at lower values. The difference
between observed canopy area for an individual and the predicted canopy
area based on dbh and height allometries provides an index of tree vigor;
the ability of this index to predict individual level growth and fecundity is
explored.
WONHAM, MARJOIRE J.,* TOMAS DE CAMINO-BECK and MARK
A. LEWIS. Centre for Mathematical Biology, CAB 632, University of Alberta, Edmonton, AB, Canada. Invasion model for West Nile virus.
The North American epidemic of the West Nile (WN) arbovirus was first
detected in New York State in 1999 and has spread each subsequent year.
The virus is typically amplified in natural transmission cycles between vector mosquitoes and host birds. The current epidemic has been particularly
damaging to corvids such as the American crow. Infected horses, humans,
and other mammals may suffer mortality, but are generally considered incidental hosts. We develop a model of WN cross-infection between birds
and mosquitoes to explore the dynamics of disease transmission and control. We extend the basic SIR model for malaria transmission (Anderson
and May 1991) to a system of differential equations for susceptible, infected, and recovered birds, and larval, susceptible, exposed, and infected
mosquitoes. We illustrate the WN transmission dynamics with numerical
simulations parameterized using literature values for American crow, Corvus brachyrhynchos, and mosquitoes, Culex pipiens. Using stability analysis of the disease-free equilibrium, we calculate the basic reproductive
rate of the epidemic, R0. By setting R0 5 1, we obtain a simple formula
for the critical susceptible mosquito population size, above which the virus
will invade. We develop a crude approximation to mosquito seasonal dynamics and outline a graphical method to evaluate invasion conditions for
WNV during a season where mosquito populations cycle. Emerging WN
control strategies include spraying after surveillance reports of WN-positive
mosquitoes or birds, or after confirmed reports of WN infection in horses
or humans. We use our model to examine the implications of these strategies, and discuss inter-annual and spatial extensions of the model.
WOOD, STANLEY R.* Human Needs and Ecosystems Services: How
Are Ecosystems Coping?
Improving the capacity of ecosystems to sustain themselves and, in turn,
human populations requires a clear understanding of their current state and
the growing pressures they face. This presentation describes the work of
the Conditions and Trends group of the Millennium Ecosystem Assessment
(MA) in examining the increasing capture of ecosystem services and alteration of ecosystems by humans. On a global scale there are large differences in the nature and distribution of ecosystems, in the type and level
of services each can provide, and in the need for and ability of humans to
capture those services. MA natural and social scientists are attempting to
describe, quantify, and map this complex web of linkages and dependencies. One task involves examining the evolution and status of ecosystem
service provision (biodiversity, food, fresh water, nutrient cycling, carbon
storage, recreation, and so on). Another is providing an ecosystem per-

spective of service provision, including the alteration and use of specific

ecosystems for example forest conversion to agriculture, the expansion
of urban areas, and the increasing modification of coastal zones. Global
evidence will be summarized by region, ecosystem and ecosystem service,
attempting to provide innovative ways of presenting this multidimensional
information. Attention is focused on developments since the mid 20th century. A key MA objective is to compile a comprehensive baseline of ecosystems and services in 2000 to serve as a platform for future comparative
studies.
WOOD, TANA E.* and DEBORAH LAWRENCE. University of Virginia,
Department of Environmental Sciences, Charlottesville, VA. Determinants of leaf litter nutrient cycling in a tropical rainforest: fertility
versus topography.
We were interested in the influence of landscape-level variation of soil
fertility and topographic position on leaf litter nutrient dynamics in a neotropical rain forest in Costa Rica. We sampled across the three main soil
types (ultisol slope, ultisol plateau, and inceptisol) to determine the effect
of soil nutrients on leaf litter nutrient concentrations while controlling for
topography, and to examine topographic effects while controlling for soil
nutrients. Both leaf litter macronutrient (phosphorus (P), nitrogen (N), sulfur (S), calcium (Ca), potassium (K), magnesium (Mg)) and micronutrient
concentrations were examined over a four-year period. All of the leaf litter
macronutrients examined, with the exception of N, varied significantly
among soil types. Variation in leaf litter [P] and [S] was explained by soil
fertility alone. In contrast, [Ca] and [Mg] were influenced by topographic
position, while [K] varied significantly with both soil fertility and topographic position. Macronutrients (P, K, Mg, S, Ca) were much less variable
than less limiting nutrients such as Fe and Al. Lower variability in essential
plant nutrients suggests a great deal of plant control over the amount of
nutrients resorbed before senescense. Leaf litter macronutrient concentrations varied significantly over the 4-year sample period; however, the pattern did not differ among the three soil types as anticipated. Hence, while
the magnitude of nutrient fluxes may be controlled by local factors such
as soil fertility and topographic position, temporal changes are likely regulated by a common environmental variable such as precipitation.
WOOLEY, STUART C.* and TIMOTHY D. PAINE. University of California, Riverside, CA. Mycorrhizal fungi may increase plant resistance
to herbivory, but at a fitness cost.
Mycorrhizal fungi associate with almost all land plants and contribute positively to plant nutrient status but constitute a substantial drain on photosynthate. Additionally, mycorrhizal plants are more resistant to insect herbivory, in a number of systems, as measured by herbivore abundance, performance and concentrations of plant defensive compounds. Because few
studies have examined these interactions using plants with nitrogen-based
defenses, we wanted to determine if host-plant resistance increased (herbivore abundance decreased) on non-mycorrhizal Nicotiana rustica compared to mycorrhizal N. rustica. We inoculated plants (n5118) with live
mycorrhizal inoculum and treated 40 plants with Benomylt to kill resident
fungi. Insect populations were recorded weekly for 10 weeks. We found
that mirid (Tupiocoris notatus) nymph abundance did not differ between
plants colonized by one species of mycorrhizal fungi (p . 0.86) but did
differ significantly (p ,0.0001) when plants were inoculated with three
fungal species. Adult populations decreased on plants colonized by multiple
fungi (p 50.024) but not by single species (p .0.05). Additionally, nonmycorrhizal plants were more heavily damaged than mycorrhizal plants,
by visual estimate. Plant chemistry (nitrogen, phosphorus, total carbon) did
not differ between treatments (p .0.2). Interestingly, while insect abundance decreased on mycorrhizal plants, mean seed weight (SW) (x5 0.201
6 0.238) and mean fruit mass (FM) (x5 0.252 6 0.316) was lower by 2.7
times and 2.0 times, respectively, than on non-mycorrhizal plants (SW
x50.538 6 0.587; FM x50.502 6 0.523), after 15 weeks of growth (p
,0.001). These results suggest that herbivore response to single mycorrhizal fungus colonization differs significantly compared to multi-species
inoculated plants. Mycorrhizal colonization appears to decrease host-plant
suitability but at an apparent significant fitness cost. Ongoing work is to
elucidate mechanisms and more clearly understand the role defensive
chemicals may play in this multi-trophic level interaction.

WRIGHT, ALLEN,1,2,* RAMESH MURTHY,1,2 GREG BARRON-GAFFORD,2 KATIE GRIEVE,2 TERENCE PAIGE,2 RANDY PATTERSON2
and KEVIN GRIFFIN.1 1 Columbia University, Lamont-Doherty Earth Observatory, Dept. of Earth and Environmental Sciences, Palisades, NY, USA;
2
Biosphere 2 Center, Columbia University, Oracle, AZ, USA. Biosphere2
Center - A large scale controlled environment facility for experimental
science.
Experimental Climate Change science provides information to researchers
investigating scientific questions through measurements obtained from manipulative experiments. In contrast to observational science, it uses controlled experiments to test hypotheses leading to mechanistic understanding
of biosphere processes. Through its capacity to support large-scale experimental research in complex mini-ecosystems (mesocosms) under controlled conditions, the Biosphere 2 Laboratory is a unique facility for accelerating our understanding of the causes of, responses to, and consequences of global climate change in marine and terrestrial ecosystems (Osmond 2002). We describe the use of several large (.1200 m2)
environmentally controlled mesocosms at the Biosphere 2 Laboratory facility. The five mesocosms currently available for research are rainforest,
ocean, desert, savannah, and agriforest. The agriforest mesocosm is divided
into three chambers of 500 m2 each. Temperature, absolute humidity, CO2,
and water can be controlled in each mesocosm. Additionally, Biosphere 2
is capable of mass and isotopic balance, replication in time, accelerated
testing, process studies, model validation, canopy access, and technology
calibration. These mesocosms provide model systems within which one
can study the effects of well-controlled single factors or a suite of factors
on entire stands of trees. We have outlined the technical aspects of this
facility, the types of experiments that have been undertaken and how such
a facility could complement research efforts in other large-scale experiments.
WRIGHT, JESSICA W.,* MAUREEN L. STANTON and ROSA SCHERSON. University of Califonia, Davis, CA, USA. Relationships between
soil chemistry and floral morphology in Collinsia sparsiflora, a native
California annual.
Serpentine soils are a challenging place for plants to grow. They have low
levels of nutrients, a skewed Mg:Ca ratio and high levels of heavy metals.
But rather than being barren wastelands, these soils support a diverse flora
that, particularly in California, acts as a refuge for a high number of native
species. One of those species is Collinsia sparsiflora, a native California
annual which has the ability to grow both on and off of serpentine soils.
At the McLaughlin Natural Reserve, C. sparsiflora plants growing on nonserpentine soils have small white flowers while plants growing on serpentine soils have much larger, showier purple flowers that appear to be adapted for outcrossing. Both ecotypes are capable of automatic self-fertilization,
however, suggesting that if this divergence in floral form is genetically
based, it may be driven in part by adaptation to edaphic factors. A reciprocal transplant experiment was used to study the relationship between
source population effects and those of the local environment on three floral
characteristics: flowering date, flower color and flower size. Flowering date
was very strongly influenced by the local environment- plants flowered
earlier on serpentine soils, regardless of where they came from. Flower
color was not influenced by the local environment. For example, white
flowered non-serpentine plants did not produce purple flowers on serpentine
soils. Flower size showed a combination of source population and environmental effects- plants produced the longest flowers on their home soil
type. Crosses between populations support the conclusion that flowering
date is strongly influenced by the environment, while flower color is under
strong genetic control, and flower size is influenced by a combination of
factors. These results suggest that the relationship between soil chemistry
and floral morphology in C. sparsiflora is highly complex and that the
response to selection on these traits could be highly dependent on the
effects of the local environment.
WRIGHT, JUSTIN P.1,* and SHAHID NAEEM.1,2 1 University of Washington, jpw5@u.washington.edu, Seattle, WA; 2 Columbia University, New
York, NY. Does functional biodiversity matter: A null-model analysis
of functional diversity ecosystem function relationships.
Recent investigations of the relationship between biodiversity and ecosystem functioning have focused on the importance of functional diversity,

Abstracts

365

most often measured as functional group richness (FGR). A number of key

experiments have shown that plots with higher FGR tend to exhibit higher
levels of ecosystem function, leading the investigators to conclude that
mechanisms associated with the complementarity of functional groups they
selected are likely to be what governs significant relationships between
biodiversity and ecosystem functioning. These studies have been primarily
plant-based and have used a priori schemes for classifying plant species.
For example, plant species are typically classified as grasses, non-leguminous forbs, or legumes. Such studies have not considered that a single a
priori functional group represents only one of an extremely large number
of possible ways for classifying species by function. Using data from six
key studies of grassland diversity and ecosystem functioning in Cedar
Creek, Minnesota and the BIODEPTH plots across Europe, we assigned
species to functional groups using Monte Carlo methods. We evaluated the
performance of a priori functional group classifications relative to the performance of randomly classified functional groups by comparing the former
with the distribution of R2 values from linear regressions between FGR
and ecosystem functions. These functions included above- and belowground biomass and soil N. Although there were statistically significant
relationships between a priori FGR and ecosystem functions in 21 out of
the 26 regressions, in only 6 cases did they explain more of the variance
than randomly assigning species to functional groups. Furthermore, neither
site nor ecosystem function predicted when a priori classification outperformed random classification. These results suggest that significant relationships between FGR and ecosystem function do not necessarily support
complementarity mechanisms implied in a priori classification schemes.
WRIGHT, MEREDITH S.* Colorado State University, Fort Collins, CO.
Relative importance of bacteria and fungi in freshwater shrimp feeding
preference and leaf decomposition.
Bacteria and fungi provide a critical link in lotic food webs, yet their
relative importance in tropical streams is not well understood. I compared
the roles of bacteria and fungi in providing energy and nutrients to two
species of freshwater shrimp in a tropical headwater stream. I first tested
four microbe treatments in both leaf decomposition and shrimp feeding
trials: bacteria only, fungi only, both bacteria and fungi, and neither bacteria
nor fungi present. I measured leaf decomposition using fine and coarse
mesh bags for two common riparian species: Cecropia schreberiana and
Dacroydes excelsa in laboratory microcosms and a first-order headwater
stream in the Luquillo Experimental Forest, Puerto Rico. I next tested feeding preference for leaf type and microbe treatment in microcosms using
Xiphocaris elongata, a shredder, and Atya lanipes, a scraper/filterer. I measured change in leaf surface area and the amount of particles generated. D.
excelsa and C. schreberiana had comparable leaf decay rates, but shrimp
preferred D. excelsa over C. schreberiana. Shrimp preference for leaves in
the microbe treatment suggests that the importance of bacterial food resources is underestimated in tropical stream food-web studies.
WRONA, AMANDA B.,1,* MICHAEL E. SHEPPARD,2 RICHARD F.
LEE3 and RICHARD G. WIEGERT.1 1 University of Georgia, abwrona@msn.com, Athens, GA; 2 Savannah State University, Savannah,
GA; 3 Skidaway Institute of Oceanography, Savannah, GA. Prevalence of
parasitic Hematodinium sp. in Georgia blue crabs (Callinectes sapidus):
a result of habitat degradation?
Blue crab (Callinectes sapidus) populations in Georgia are in severe decline. Factors that may be responsible include overfishing, habitat degradation, and disease. For aquatic estuarine species like blue crabs, habitat
degradation may include changes in water salinity to values that cause
increased physiologic stress and decreased survival. As salinities in Georgia
estuaries have been increasing over the past few years due to drought conditions, Hematodinium sp., a lethal parasitic dinoflagellate that infects blue
crabs, has been found in a large percentage of Georgia crabs (as high as
50% of crabs sampled in some areas). High prevalence of this parasite is
thought to coincide with high estuarine salinity. In January 2002, we began
to monitor for Hematodinium within the Sapelo Island National Estuary
Research Reserve, GA in order to investigate the relationship between Hematodinium infection in blue crabs, water conditions, and crab abundance.
Monthly samples of crabs taken from commercial traps and by trawl were
tested using a neutral red assay technique. The parasite was found in crabs

366

Abstracts

from 78 to 165 mm CW in size and most often occurred in males (21%)

vs. females (9%). Infection is positively correlated with both water salinity
and temperature and negatively correlated with commercial catches. Peak
infection (33% prevalence) occurred in August, when water temperatures
and salinity were greatest and prevalence was lowest in January, (3%)
following several weeks of cold temperatures and heavy rain that lowered
salinity and water temperatures to 16 ppt. and 7.6 8C.
WURZBURGER, NINA,1,* ANTHONY S. HARTSHORN2 and RONALD
L. HENDRICK.1 1 University of Georgia, Athens, GA; 2 University of California, Santa Barbara, CA. Ectomycorrhizal fungal community structure across a bog-forest ecotone in southeast Alaska.
We examined the ectomycorrhizal fungal community across a bog-forest
ecotone in southeast Alaska. The bog and edge were both characterized by
poorly drained Histosols and vegetation including and a continuous layer
of Sphagnum species, ericaceous shrubs, Carex species, and shore pine
(Pinus contorta Dougl. ex Loud. var. contorta). The forest had better
drained Inceptisols and Spodosols and a tree community comprised of
western hemlock (Tsuga heterophylla (Raf.) Sarg.), yellow cedar (Thuja
plicata Donn ex D. Don.), sitka spruce (Picea sitchensis (Bong.) Carr.) and
shore pine and an understory of ericaceous shrubs and herbs. Ectomycorrhizal root tip density (ECM tips cm-3) was significantly different in the
forest (6.04) compared to the bog and edge (0.05, 1.92) and ectomycorrhizal colonization was significantly different in all three plant communities
(bog 53, edge 88 and forest 99 %). The below ground ectomycorrhizal
fungal taxa were analyzed using molecular techniques (PCR-RFLP and
nucleotide sequencing) and the taxa observed are typical of boreal forests
worldwide. Three ECM fungal taxa, Suillus tomentosus (Kauffman) Singer,
Cenococcum geophilum Fr.:Fr, and a Russula species differed in relative
frequency, yet were among the four most frequent in all three plant communities. Differences in ECM fungi across the ecotone are likely due to
the composition of host trees and soil characteristics. Although differences
in ECM fungal richness were observed across plant communities, comparisons were confounded by unequal sampling of ECM roots due to root
density and colonization differences. We discuss challenges associated with
sampling and estimating richness in ectomycorrhizal fungal communities.
WYATT, JULIE L.* and MILES R. SILMAN. Biology Department, Wake
Forest University, Winston Salem, NC. Population regulation in two
rainforest dominant trees: Distance dependence depends.
Plant-animal interactions are crucial for maintaining diversity in tropical
tree communities in both equilibrium and non-equilibrium theory. The relative importance of these interactions can be expected to vary depending
on the composition of animal communities. We found distance-dependent
recruitment in two numerical dominant tree species: Iriartea deltoidea and
Astrocaryum murumuru (Arecaceae) was dependent on the population dynamics of Tayassu pecari, the largest, most widespread, and highest biomass seed predator in Neotropical forest communities. Areas with altered
mammal communities lacking T. pecari had decreased parent-offspring distances and up to 6000% increases in viable seeds on the forest floor. This
trend held up spatially at sites with different levels of human impact, and
temporally when T. pecari was locally extinct at the Cocha Cashu Biological Station, Peru. Recent reviews have concluded that invertebrates and
fungi, but not vertebrates, cause distance and density dependent seed and
seedling mortality conforming to the predictions of the Janzen-Connell
model. We suggest that the discrepancy between this and prior studies is
because the majority of previous studies have been conducted in altered
ecosystems lacking T. pecari, precluding evidence of distance or density
dependent seed mortality by large mammals. Empirical evidence and population projection analyses suggest the coupling of these two dominant
canopy trees to the highest biomass seed predator can cause revolutionary
changes in the structure of tropical tree communities.
XI, WEIMIN,1,* ROBERT K. PEET1 and DEAN L. URBAN.2 1 University
of North Carolina at Chapel Hill, Chapel Hill, NC, USA; 2 Duke University,
Durham, NC, USA. Hurricane disturbances, tree species diversity, and
succession in North Carolina Piedmont forests.
Windthrow has been hypothesized to play a critical role in maintaining
species diversity in temperate deciduous forests. Recent work by Beckage

and Clark has shown that large canopy gaps maintain tree diversity in
southern Appalachian forests. However, few long-term data are available
for assessing the importance or generality of disturbance-maintained tree
species coexistence. We have used long-term tree (stems .1 cm dbh) demographic data to assess the effect of Hurricanes Hazel (1954) and Fran
(1996) on tree species diversity and tree replacement in North Carolina
Piedmont forests. In addition, we used understory sapling (stems . 1m tall
and , 1 cm dbh) growth and survival data to project post-hurricane changes in canopy composition. Our results support the hypothesis that large
wind disturbances help to maintain local tree species diversity. Although
there is often an immediate drop in diversity following hurricane damage,
species diversity of saplings quickly increases to levels that typically exceed those prior to the disturbance.This typically leads to an increase in
tree species diversity (sometimes by as much as a factor of two) in substantially damaged stands. Nonetheless, time-series analysis showed that
hurricanes significantly decrease the dominance of several important,
shade-intolerant canopy species such as oaks and hickories owing to lack
of advanced regeneration in the understory, while increasing the dominance
of pre-established, shade-tolerant species such as red maple. We conclude
that large, infrequent wind disturbances help to maintain local tree diversity, but also accelerate the increase in dominance of red maple. This pattern may widely apply in the Piedmont region and adjacent temperate forests regions of the eastern United States.
XU, CHENGYUAN,1,2,* GUANGHUI LIN,1 KEVIN L. GRIFFIN2 and
RAYMOND SAMBROTTO.2 1 Biosphere 2 Center of Columbia University, Oracle, AZ; 2 Lamont Doherty Earth Observatory of Columbia University, Palisade, NY. Respiratory CO2 is 13C-enriched in five C3 plants.
We compared the carbon isotope ratios between respiratory CO2 and leaf
organic components (soluble sugar, water soluble fraction, starch, protein
and bulk organic matter) in 5 C3 plants grown in green house and Biosphere 2, including one species, Populus deltoides, in 3 different growth
CO2 concentrations. For all species or CO2 treatments, respiratory CO2 was
13
C-enriched in comparison with all leaf organic components, but the
amount of enrichment does not depend on the signature of the organic
components examined. We also found that d13C of leaf organic components
are correlated to d13C of leaf respiratory CO2. The amount of 13C enrichment was significantly influenced by the type of organic component examined, but not by species or CO2 treatments. We argue that in normal
growth condition, non-statistical carbon isotope distribution in sugar may
determine the signature of respiratory CO2 dominantly. These results suggest that 1) 13C-enriched respiratory CO2 is widespread in C3 plants and
2) we can predict d13C of respiratory CO2 for ecosystem scale study by
combining the average 13C-enrichment in respiratory CO2 and the signatures of leaf organic components.
XU, TAO,1,* DAFENG HUI,2 YIQI LUO2 and LUTHER WHITE.1,* 1 Department of Mathematics,601 Elm Avenue,University of Oklahoma, Norman, Oklahoma, USA; 2 Department of Botany and Microbiology,University of Oklahoma,770 Van Vleet Oval, Norman, Oklahoma, USA.
Modeling net ecosystem exchange using auto-regression model with
exogenous inputs.
Improving the prediction accuracy of net ecosystem exchange (NEE) in
terrestrial ecosystems is an important task in ecosystem modeling. Considering that the present time NEE not only depends on the current and historical climatic variables, but also correlates with its own past history, we
proposed an approach by using a time series analysis model. The model
can be best described by the ARX (auto-regression with exogenous inputs)
model in the form of A(q)y(t)5B(q)u(t)+e(t), where y(t) is output NEE,
u(t) is multiple-input including photosynthetically active radiation, air temperature, relative humidity, vapor pressure deficit etc. and e(t) is model
error. Weekly averaged NEE and climatic variables from year 1992 to year
1999 in Harvard Forest were used in analysis. Compared to multivariate
linear regression model, ARX model greatly improved the NEE estimation.
The variation in NEE explained by the ARX model increased on the average to 65% from 52.3% given by multiple regression when the autoregressor had time delay from 1 to 20, and that was increased to more than
71% on the average if the auto-regressor had time delay ranging from 45
to 55. This indicates that the NEE pattern of the present season, while

being influenced by the current environmental factors, is either closely

auto-correlated with its most recent history or, at another extreme, closely
auto-correlated with the patterns of the same season of the previous year.
While the former has an obvious explanation, the later strongly suggests
that it is possible to use historical patterns of NEE of a specific site, coupled
with the current/predicted environmental factors, to generate more accurate
current/predicted NEE at that specific site.
YAGER, LISA Y.,1,* JEANNE JONES2 and DEBORAH L. MILLER.3
The Nature Conservancy, Camp Shelby Field Office, Camp Shelby, MS,
US; 2 Mississippi State University, Mississippi State, MS, US; 3 University
of Florida, Milton, FL, US. Cogongrass (Imperata cylindrica) encroachment into upland pine communities on Camp Shelby Training Site
(CSTS), Mississippi.

Historically, uplands on CSTS were dominated by longleaf forests with an

open understory of scattered hardwoods and bluestem grass-dominated
groundcover. Many areas may still be characterized this way. However, fire
exclusion over time has resulted in shrub and hardwood encroachment and
reduced herbaceous abundance in pine forests on much of CSTS. Differences in light availability and vegetative structure, composition and abundance between these 2 community types (pine/bluestem, pine/shrub) may
affect their susceptibility to invasion by cogongrass. Introduced in the early
1900s from Asia, cogongrass is spreading throughout the southeast and
can displace native vegetation and alter fire regimes and soil processes in
native plant communities. Quantification of rates of spread and possible
factors related to rates of spread into pine/bluestem and pine/shrub communities is needed to develop appropriate control strategies. To compare
rates of vegetative spread (tiller extension), 20 cogongrass patches growing
into pine/bluestem or pine/shrub were selected for each community type.
In April 2002, a 1 m section of patch edge selected so that no tillers
extended further into the habitat of interest was marked with stakes for
each patch. Soil pH, nutrient status and texture for samples collected at
these edge plots did not differ between the two community types. Although
light availability (PAR) and percent cover of grasses, forbs, shrubs and
vines at the edge of the patches differed between the two community types,
cogongrass tiller extension measured in October 2002 from the edge plots
was not significantly different. Tiller extension from patch edges ranged
from 0 to . 0.5 m for both habitat types. Results of fall and spring tiller
extension rate and the relationship of spread to environmental variables
(PAR, dominant vegetation by type and species) will be evaluated to determine community susceptibility to vegetative encroachment by cogongrass and potential factors influencing its growth.
YANAI, RUTH D.,1 STEVEN P. HAMBURG,2 MARY A. ARTHUR3 and
JOEL D. BLUM.4 1 SUNY-ESF, Syracuse, NY; 2 Brown University, Providence, RI; 3 U of Kentucky, Lexington, KY; 4 U of Michigan, Ann Arbor,
MI. Whether repeated harvest and acid rain threaten calcium supply
in northern hardwood forests may depend on apatite weathering.
Acid rain and repeated forest harvest are thought to threaten the supply of
calcium to forested ecosystems. Ecosystem budgets for New Hampshire
northern hardwoods show that young stands (,30 y old) mobilize Ca at
rapid rates (3.3 to 4.7 g Ca m-2 y-1), based on the rates of Ca export in
streamwater and accumulation in vegetation and the forest floor. In contrast,
older stands are in approximate steady state (losing 1 g Ca m-2 y-1). If
young stands tap a pool of soil Ca unavailable to older stands, then repeated
harvest will not result in rapid Ca depletion, as has been predicted based
on rates of export and weathering in older stands. If, on the other hand,
rapid Ca uptake by young stands hastens the depletion of readily available
pools, sustainable forestry may be threatened. We have identified apatite
(Ca phosphate) at depth in soils at the Hubbard Brook Experimental Forest
(65 g Ca/m2 in E and B horizons and 6 g Ca/m2 per cm of depth in the C
horizon), based on sequential extractions and P:Ca and Ca:Sr ratios. We
propose that apatite weathering is accelerated in young stands, perhaps
because P is more limiting than N shortly after forest disturbance. We have
analyzed soils and plant material from 10 additional sites in New York,
Pennsylvania, and Maine to determine the importance of this previously
unappreciated Ca source to forests of different ages.

Abstracts

367

YANG, LOUIE H.* University of California, Davis, Center for Population

Biology, Davis, CA. 17-year periodical cicadas as a resource pulse:
Effects on soil nitrogen availability.
Many natural systems experience resource pulses: occasional, sudden and
brief changes in resource availability over time. While resource pulses appear to be a common feature of many natural systems, the ecological importance of these rare events of exceptional magnitude remains largely
unexplored, especially in a belowground context. Here, I present the results
of a manipulative field experiment conducted to examine the effects of
superabundant 17-year periodical cicadas (Magicicada spp.) as a resource
pulse in eastern North American forests. In these experiments, supplementation of cicada carcasses substantially increased the availability of soluble
soil nitrogen over the summer growing season. I examine potential indirect
consequences of this resource pulse for both belowground and aboveground
communities and consider some emerging generalities for the ecological
consequences of resource pulses in belowground systems.
YANOSKY, THOMAS M.* and DANIEL E. KROES. US Geological Survey, Reston, VA, USA. Impact of saline intrusion upon growth of loblolly pines, Blackwater National Wildlife Refuge, Maryland USA.
Rings of loblolly pines (Pinus taeda L.) were studied to determine the
effects of saline intrusion upon radial growth and survivability within parts
of the Blackwater National Wildlife Refuge, and to estimate the approximate onset of intrusion. The sparse forest is even-aged and began to grow
about 1940. Increment cores were collected at breast height from 24 living
trees along an intrusion gradient; cross sections also were cut from 11 dead
trees. Surface elevations were determined near the trunk of each pine. Each
ring was crossdated, measured to the nearest 0.01 mm, and converted to
area (in cm2). Mean-annual sensitivity (MAS) values (a measure of ringto-ring variability and thus of growth responses to environmental variables)
of all 35 trees ranged from 0.18 to 0.29 for 13 unimpacted trees, and from
0.34 to 0.72 for 22 impacted living and dead trees. MAS values for unimpacted trees from 1960-1980 generally were greater or did not differ
from those from 1980-present; for most impacted trees, however, MAS
values were significantly greater after 1980 than during 1960-1980, suggesting that a variable other than climate (presumably saline intrusion)
became an important factor controlling tree growth after 1980. Although
the radial growth of all study trees declined during droughts of the mid
1990s, unimpacted trees subsequently resumed prior growth trajectories and
continue to grow vigorously at present; impacted living trees, however,
have not recovered and growth since the mid 1990s remains suppressed.
The 11 dead trees died between 1994 and 2002 following declines in 2-7
consecutive years during which ring areas were less than approximately
1.0 cm2. Based on these findings, it seems likely that surviving trees will
die in the near future as the forest continues to convert to marshland.
YANOVIAK, STEPHEN P.,1,2,* L P. LOUNIBOS,1 S C. WEAVER2 and R
B. TESH.2 1 University of Florida, Florida Medical Entomology Lab, Vero
Beach, Florida; 2 University of Texas Medical Branch, Galveston, Texas.
Diversity of phytotelm fauna along a deforestation gradient in the Peruvian Amazon.
Deforestation potentially alters the distribution and abundance of phytotelmata and their resident fauna. We surveyed phytotelm habitats along a
deforestation gradient near Iquitos, Peru, as part of a larger study of arbovirus ecology. Mean phytotelm density was greater in cultivated areas
("chacra"; 174 6 33 SD per hectare) than in secondary forest ("purma";
28 6 36) and primary forest ("bosque"; 25 6 17). Fallen leaves and plant
axils were the most abundant microhabitats present at all sites. Half (47%)
of the colonizable microhabitats contained macroinvertebrates and mosquitoes were the most abundant occupants. In addition to the microhabitat
survey, replicated phytotelm analogs (water-filled bamboo sections) were
placed along the gradient and destructively sampled every two weeks.
Mean species richness (ca. 3 per bamboo) and abundance (ca. 50) were
similar among chacra, purma and bosque. Trichoprosopon digitatum was
the most abundant macroinvertebrate in the bamboo sections across all
sites. However, species composition of top predators differed, with Toxorhynchites spp. occurring more often in chacra and odonates exclusively
inhabiting bosque. These differences may have cascading effects on the

368

Abstracts

abundance and composition of mosquito prey along the deforestation gradient.

YARNES, CHRISTOPHER T.* and WILLIAM J. BOECKLEN. New Mexico State University, Las Cruces, NM. Synergistic abiotic factors and
host plant quality: Trophic consequences of variable nitrogen and elevation.
Theory surrounding the structure and dynamics of trophic relationships in
terrestrial ecosystems has emphasized the need for empirical studies that
describe and quantify the function of biotic and abiotic heterogeneity. Abiotic heterogeneity can have significant consequences for natural populations both directly and indirectly through the production of biotic heterogeneity. Plant species are known to exhibit heterogeneity in host quality
for insect herbivores across multiple spatial and temporal scales, from within-leaf and intraplant variation to population and intraspecific variation.
This heterogeneity may serve as a defense strategy per se for plants with
long generation times by serving as a means to contend with a multitude
of rapidly-evolving insect herbivores exhibiting rapid generation times. To
date, relatively few empirical studies have examined the synergistic effects
of multiple sources of abiotic heterogeneity on spatio-temporal host quality
variation and the effect on the relative roles of bottom-up and top-down
control on herbivore populations. We examined preference-performance relationships and mortality of two species of leaf-mining Lepidoptera, Phyllonorycter sp. and Camareria sp. (Lepidoptera: Gracillariidae and Nepticulidae), associated with host-quality of Quercus gambelii (Nutt.) along an
elevation gradient under two nitrogen amendment levels in the San Mateo
mountain range of west-central New Mexico, U.S.A. during 2001 and 2002.
Foliar nitrogen was heterogeneous at multiple spatial and temporal scales
and was significantly related to elevation, nitrogen availability and ambient
environmental parameters. Elevation and seasonal variation in foliar nitrogen had significant effects on herbivore densities. Herbivore emergence and
parasitoid success was significantly related to nitrogen treatment. Herbivore
performance and parasitism rates appear to be related to the magnitude of
within-tree spatio-temporal heterogeneity in foliar nitrogen. Extreme
drought during the 2002 growing season contributed to massive levels of
unexplained mortality and will be discussed.
YEAKLEY, J. A.,1,* D. C. COLEMAN,2 B. L. HAINES,3 B. D. KLOEPPEL,2 J. L. MEYER,2 W. T. SWANK4 and S. STEINER.2 1 Environmental
Science, Portland State University, Portland, OR, USA; 2 Institute of Ecology, University of Georgia, Athens, GA, USA; 3 Department of Plant Biology, University of Georgia, Athens, GA, USA; 4 Coweeta Hydrologic
Laboratory, USDA-Forest Service, Otto, NC, USA. Hillslope and streamwater nutrient dynamics following upland riparian canopy tree windthrow in the southern Appalachians.
We investigated effects of natural blowdown of canopy trees on nutrient
export to streams in the southern Appalachians. Transects were instrumented on adjacent hillslopes in a first order watershed at the Coweeta
Hydrologic Laboratory (35o03 N, 83o 25 W). Dissolved organic carbon
(DOC), K+, Na+, Ca2+, Mg2+, NO3--N, NH4+-N, PO43--P, and SO42- were
measured for two years prior to disturbance. In October 1995, Hurricane
Opal uprooted nine canopy trees on one of the hillslopes, downing 81% of
total woody biomass. During three years following disturbance, soilwater
NO3--N on the storm-affected hillslope showed persistent 500-fold increases, groundwater NO3--N increased by fourfold, and streamwater NO3--N
doubled. Significant changes also occurred in soilwater pH, DOC, SO42-,
Ca2+ and Mg2+. By the end of the sixth year following disturbance, streamwater levels of NO3--N had returned to pre-disturbance levels, although
soilwater NO3--N concentrations remained slightly elevated. Our results
suggest nutrient uptake by canopy trees is a key control on NO3--N export
in upland riparian zones, and that streamwater nutrient responses to localized riparian canopy tree windthrow in upland forested watersheds may last
as long as half a decade.
YEE, DONALD A.* and STEVEN A. JULIANO. Illinois State University,
Normal, Illinois, U.S.A. Does microorganism productivity influence
macroinvertebrate richness in aquatic detritus-based communities?
The food web of water-filled tree holes is an ideal system to test alternative
hypotheses for relationships between productivity and species richness.

Tree hole communities are dependent on exogenous inputs of detritus,

which support a microorganism assemblage, which in turn supports numerous macroinvertebrate species. We conducted an 11 month field study
in artificial tree holes to determine if: a) different initial inputs of detritus
(senescent leaves, dead invertebrates) affect microorganism productivity
and abundance; 2) the rate of subsequent input affects microorganism productivity and abundance; and 3) microorganism productivity or abundance
influence richness or abundance of macroinvertebrates. We established 6
replicates of 4 initial input treatments (0.25x, 0.5x, 1x, 4x observed field
detritus inputs) that were open to colonization for one month. Following
that period, 2 containers of each initial input treatment were assigned one
of 3 subsequent input treatments: None (no additional detritus); Once (all
detritus at the beginning); and Weekly (equal detritus additions over 6
weeks). Both Once and Weekly subsequent inputs ultimately yielded a
doubling of the initial detritus amount. Every 3 weeks we measured microorganism productivity as respiration rate (mL of O2 hr-1) of fluid and
leaf samples, microorganism abundance, and abundance and richness of
macroinvertebrate species in each container. During the initial phase, the
two highest detritus treatments produced significantly higher diversity and
abundance. There were non-significant trends of decreased leaf-associated
productivity and increased fluid-associated productivity as total detritus increased. Richness was significantly higher in containers receiving detritus
all at once compared to other addition rates. Abundance was affected by
both initial and subsequent input treatments. In all sampling periods, there
were significant positive relationships between richness and abundance.
This is one of the first studies of the effects of microorganism productivity
on richness and abundance of macroinvertebrate species over an extended
period.
YEE, SUSAN MH.* University of Chicago, Chicago, IL, 60420. Spatial
structure affects dynamics in a treehopper - ant mutualism.
Although mutualisms are ubiquitous in nature, formal models of mutualisms have seen limited development and application. Simple models suggest mutualisms are prone to extinction, making it difficult for mutualisms
to become established from small numbers of individuals. Reaction-diffusion models indicate spatial structure may be a stabilizing mechanism in
mutualism dynamics. It is unclear, however, whether space is a mechanism
preventing extinction in natural populations. I used a reaction-diffusion
model of mutualist dynamics to explore how aspects of spatial structure,
including dispersal ability and initial spatial distribution, are likely to affect
species interactions. Predictions were tested experimentally in a naturally
occurring ant-treehopper mutualism. I assessed the effects of dispersal ability on treehopper dynamics within a metapopulation of treehopper colonies
tended by a single ant colony. I manipulated treehopper movement between
plants in concert with ant access to treehopper colonies. Results indicate
that a reduction in treehopper dispersal significantly reduces both treehopper population growth rate and the number of plants within each metapopulation occupied by treehoppers. Effects of treehopper dispersal were strongest when ant access to each colony was low. Clearly, the ability of treehoppers to colonize plants and the ability of ants to locate treehopper colonies are important determinants of the dynamics in this mutualistic
interaction. The stability of mutualisms in natural systems likely depends
on the colonization abilities of each species, as well as the initial spatial
distribution of each species and the availability of habitat.
YERMAKOV, ZHANNA* and DAVID E. ROTHSTEIN. Michigan State
University, East Lansing, Michigan. Recovery of soil processes after
wildfire in jack pine forests.
We studied how biological, chemical and physical factors interact to influence the recovery of soil processes after wildfire in jack pine (Pinus banksiana) forests. We measured changes in soil organic matter (SOM) quality,
quantity and mineralization rates across a chronosequence of eleven wildfire-regenerated jack pine stands in the northern Lower Peninsula of Michigan. The stands ranged in age from 1 to 72 years. Changes in C and N
mineralization and SOM all conformed to a gamma function
(Y5AXbecxd+E), in which all three processes initially declined to a minimum by 15 years since wildfire and later increased to an asymptote. Although both N mineralization and SOM quantity increased with age, the
relative proportion of total soil N mineralized decreased, suggesting a de-

cline in SOM quality. Changes in microclimate with stand age also influenced the recovery of soil processes after wildfire. In particular, the magnitude of seasonal fluctuations in C and N mineralization was greater in
younger stands compared to older stands. Our results show that the recovery of C and N mineralization is predictable after wildfire and that the rate
of recovery is controlled by the interaction of SOM quality and quantity,
as well as microclimate.
YODER, CARRIE L., WILLIAM J. PLATT* and L. STANTON. Louisiana
State University, Department of Biological Sciences, Baton Rouge, LA.
Multiple disturbances and the dynamics of Southeastern coastal ecosystems.
Large-scale disturbances that occur in close association may produce effects different from those of isolated disturbances. Novel effects on important species may result in long-term changes, especially in ecotones
between plant communities. Although disturbances are often hypothesized
to structure communities, predictions of combined effects of different types
of disturbances do not exist for communities and ecotones in most landscapes. We develop concepts for interactive effects of hurricanes, fires, and
sea level rise as the most prominent disturbances in current southeastern
coastal landscapes. We initially summarize studies conducted in isolation,
focusing on effects of wind damage during hurricanes, fire damage, prolonged flooding and increased salinity from sea level rise. We then combine
the known effects to explore potential effects of interactions among these
disturbances. Throughout we focus on predictions of short- and long-term
effects of single and multiple disturbances that influence coastal fresh-water
marshes and bottomland forests, as well as the ecotone between these communities. We apply these concepts to a gradual topographic gradient from
a soft bottom marine community along a tidal creek through an open estuarine marsh, fresh water marsh, into bottomland and mesic hardwood
forest in the Weeks Bay National Estuarine Research Reserve in Baldwin
County, Alabama. We explore relationships between disturbances and their
effects on dominant species in the ecotone between fresh-water marsh and
bottomland hardwood forest, focusing on the dominant herbaceous species,
Cladium jamaicense, and the dominant tree species, Acer rubrum. We apply
the concepts regarding potential interactions between natural disturbances
to predict community and landscape patterns expected at Weeks Bay National Estuarine Research Reserve.
YOO, HO JUNG S.* Purdue University, yoo@entm.purdue.edu, West Lafayette, IN, USA. Factors determining spatial distribution of an herbivorous insect: A model of patch density-dependent rates of parasitism vs. dispersal mortality.
The western tussock moth, Orgyia vetusta, is a poor disperser in patchy
coastal habitats. Only first instar larvae are capable of significant betweenpatch movement, which is accomplished by passive wind dispersal. At a
site in central California, however, moth abundance is greater on isolated
patches of its host plant, Lupinus chamissonis, despite the likelihood that
dispersal mortality is higher in areas of low patch density. Empirical studies
have previously shown that 1) mortality in crawlers (instars II to VI) is
density-dependent, and 2) fecundity, pupal predation, and pupal parasitism
by a wasp are significantly correlated with local patch density. Only pupal
parasitism, however, varies in a manner consistent with the observed abundance pattern. To determine whether variation in pupal parasitism is sufficient to explain the distribution of moths, a spatially-explicit agent-based
simulation model was developed. Within-patch dynamics included all four
significant relationships in demographic processes mentioned in 1) and 2)
above. Dispersal between patches was localized. A simulation experiment
was run in a 2 x 2 factorial design to qualitatively assess the independent
and joint effects of local (vs. global) dispersal and the spatially varying
(vs. fixed) birth and death rates. The results indicated that while local dispersal alone causes highest abundances in areas of high patch density, the
effect is essentially reversed when variability in pupal parasitism by the
wasp is included; areas of high patch density attain the lowest moth abundances. This result was robust to a realistic range of parameter values for
dispersal distances, emigration rates, and larval death rates. Thus, attack
patterns by a parasitoid appear to have a greater influence than does dispersal ability on moth distribution.

Abstracts

369

YOSHIDA, TAKEHITO,1 LAURA JONES,1 STEPHEN P. ELLNER,1

GREGOR F. FUSSMANN2 and NELSON G. HAIRSTON.1,* 1 Department
of Ecology and Evolutionary Biology, Cornell University, Ithaca, New
York, USA; 2 Institute fuer Biochemie und Biologie, University of Potsdam,
Potsdam, Germany, Germany. Rapid evolution drives ecological dynamics in a predator-prey system.
The accumulating evidence for rapid evolution indicates that ecological and
evolutionary dynamics can occur on similar time-scales. Theoretical studies
have predicted that rapid evolution can affect ecological dynamics but predictions remain inconclusive: for example, depending on assumptions, prey
evolution can stabilize or destabilize predator-prey dynamics. Using laboratory microcosms, we studied the effect on predator-prey (rotifer-alga)
cycles of rapid prey evolution in response to oscillating predator density,
in order to test explicit predictions of a model for this system allowing
prey evolution. Here we report an evolutionary tradeoff between algal competitive ability and defense against consumption. We then demonstrate its
effect on cycle dynamics by manipulating the evolutionary potential of the
prey population. Single-clone algal cultures (lacking genetic variability)
produced short cycle periods and typical quarter-phase lags between prey
and predator densities, while multi-clonal (genetically variable) algal cultures produced long cycles with prey and predator densities nearly out of
phase, exactly as predicted. These results confirm that prey evolution can
dramatically alter predator-prey dynamics, hence attempts to understand
population oscillations in nature cannot ignore potential effects from ongoing rapid evolution.
YOUNG, LAUREN M.* and SVATA M. LOUDA. University of Nebraska,
Lincoln, Nebraska. Differences in survival, traits, and herbivory of
seedlings from native and invasive congeners.
Naturalized plants often appear more vigorous in their non-native range. A
hypothesized mechanism underlying such increased vigor by plants in the
non-native range is reduced herbivore pressure. Under this hypothesis
plants in their non-native range are expected to allocate more to growth
and biomass (enhanced competitive ability) and less to defense (absence
of native enemies). We examined size, frequency of herbivory, and survival
of invasive thistle seedlings, Cirsium vulgare, in North America. Seedlings
of C. vulgare were grown in the field from seed using seed from exotic
and naturalized seed sources. We compared the exotic seedlings to those
of a native North American congeneric thistle, Cirsium altissimum. The
proportion of seedlings surviving was greatest for the native C. altissimum,
followed by C. vulgare from naturalized seeds, and then from exotic seed
origin. Native C. altissimum and C. vulgare seedlings of exotic seed origin
had greater rosette area than did C. vulgare seedlings of naturalized seed.
Unexpectedly, C. vulgare seedlings from naturalized seed were more frequently damaged by July, than were those of C. altissimum, the native
thistle. The greater survival of C. vulgare seedlings from naturalized seed
than from exotic seed is consistent with the hypothesis of potential adaptive
change in this weed. Additionally, the greater frequency of herbivore feeding on Cirsium vulgare seedlings from naturalized seed than the native C.
altissimum suggests that native herbivores could be providing resistance to
invasive spread by C. vulgare in tallgrass prairie areas of eastern Nebraska.
YOUNGSTEADT, ELSA K.* University of Wisconsin-Milwaukee, Milwaukee, WI, USA. The effects of ants on the carnivorous caterpillar
Feniseca tarquinius and the role of chemical camouflage in the Feniseca-ant interaction.
The majority of caterpillars in the butterfly family Lycaenidae are myrmecophilous, that is, they provide ants with nutritional secretions in exchange for protection from natural enemies. However, the lycaenid subfamily Miletinae is exceptional in its relationship to both ants and Homoptera.
Miletine larvae are carnivores of ant-tended homopterans, yet lack the secretory organs crucial to myrmecophily in other lycaenids. This raises questions of how miletines have invaded the ant-homopteran symbiosis, and
how ants affect miletine larval success. Here I address these questions
through a study of the only New World miletine, Feniseca tarquinius. I
present evidence that chemical camouflage enables F. tarquinius caterpillars to go undetected by ants, and that caterpillars may benefit from the
presence of ants. Gas chromatographic analysis of cuticular extracts indi-

370

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cated that F. tarquinius caterpillars bear a cuticular hydrocarbon profile

similar to that of its aphid prey, and that the profile varies with prey species.
Bioassays in the field confirmed the behavioral relevance of the extracts
analyzed. Field censuses of 98 caterpillars in 2001 suggested F. tarquinius
larvae benefit from the presence of some ant species. Caterpillars in the
2nd through 4th instars were less likely to conceal themselves under silken
webs if the ants Camponotus pennsylvanicus or Formica obscuriventris
were tending the aphid prey, and pupation rate was greater in the presence
of C. pennsylvanicus than in the absence of ants. These effects were not,
however, reproduced by 11 pairs of caterpillars in an ant exclusion experiment in a screened garden in 2002.
YURKONIS, KATHRYN A.* and SCOTT J. MEINERS. Eastern Illinois
University, Charleston, IL. Invasion by Lonicera japonica impacts species turnover in a successional system.
Invasive plant species are often associated with reductions in species richness within native communities. Reductions in species richness can either
occur through changes in species extinction rates or colonization rates within native communities. Documenting species turnover in long-term studies
provides an opportunity to identifying the mechanistic basis for reductions
in species richness associated with exotic species invasions. One such study
that has been documenting exotic species invasions in abandoned agricultural fields is the Buell-Small Succession study. Over the past 44 years,
species abundance and coverage has been recorded for 480 plots in 10
fields differing in last crop and type and season of abandonment. During
a ten year time window from 5 to 15 years since abandonment, coverage
of the exotic species Lonicera japonica increased. Within this time period,
increasing Lonicera japonica cover is associated with decreases in species
richness. Additional analyses indicate increases in coverage of Lonicera
japonica have little effect on community extinction rates, whereas increasing Lonicera coverage results in a reduction in community colonization
rates. In association with reduced colonization rates, variation exists in
dominant species and lifeform responses to invasion. These findings suggest that species richness declines seen with invasion result from reduced
community colonization rates and not competitive species displacement as
is often hypothesized.
ZAFIRIS, ANGELIKIE A.,* EVELYN E. GAISER and MICHAEL S.
ROSS. Southeast Environmental Research Center, Florida International
University, Miami, FL. Tracking effects of salt-water encroachment on
community migration in coastal South Florida wetlands using mollusks.
Coastal wetland communities of South Florida have been cut off from
freshwater sheet flow for decades and are contracting and migrating landward due to salt-water encroachment. We conducted a paleoecological
study to determine the effect of salt-water encroachment on the location of
boundaries between fresh- and salt-water ecotones in the southeastern saline Everglades. Wetland soils were cored to bedrock at 8 locations in two
transect perpendicular to the coast. Transitions from basal marl soils to peat
were evident throughout the transect, with a deepening of the peat layer
toward the coast indicating increased production and interior expansion of
the fringing mangrove forest. Mollusks were abundant throughout the cores
and the 24 collected taxa served as useful paleoecological indicators. Modern distributions among 86 sites in the same wetland were used to determine local habitat affinities, which were then applied to infer past settings
from the sequence of sedimented mollusks. Sites located between the drainage canal and the coast showed signficant upcore increases in the ratio of
marine to freshwater taxa, while sites to the interior of the canal showed
the opposite trend. Terrestrial taxa have also increased in the interior sites,
indicating a transition from shallow gramminoid marsh to the current
shrub-forest community. Together with historic accounts and aerial photograph archives, the paleoecological data are showing an interior migration
of the fringing mangrove ecotone within the past 60 years, replacement of
a mixed gramminoid-mangrove zone by a dense monoculture of dwarf
mangroves, and a confinement of the freshwater gramminoid marsh to landward areas between urban developments and drainage canals.

ZAK, DONALD R.,1,* WILLIAM E. HOLMES,1 ADRIEN C. FINZI,2

RICHARD J. NORBY3 and WILLIAM H. SCHLESINGER.4 1 University
of Michigan, Ann Arbor, Michigan; 2 University of Boston, Boston, Massachusetts; 3 Oak Ridge National Laboratory, Oak Ridge, Tennessee; 4 Duke
University, Durham, North Carolina. Soil Nitrogen Cycling Under Elevated CO2: A Synthesis of Forest FACE Experiments.
The extent to which greater NPP will be sustained as the atmospheric CO2
concentration increases will depend, in part, on the long-term supply of N
for plant growth. Over a two-year period, we used common field and laboratory methods to quantify microbial N, gross N mineralization, microbial
N immobilization, and specific microbial N immobilization in three freeair CO2 enrichment experiments (Duke Forest, Oak Ridge, Rhinelander).
In these experiments, elevated atmospheric CO2 has increased the input of
above- and belowground litter production, which fuels heterotrophic metabolism in soil. Nonetheless, we found no effect of atmospheric CO2 concentration on any microbial N cycling pool or process, indicating that greater litter production had not initially altered the microbial supply of N for
plant growth. Thus, we have no evidence that changes in plant litter production under elevated CO2 will initially slow soil N availability and produce a negative feedback on NPP. Understanding the time scale over which
greater plant production modifies microbial N demand lies at the heart of
our ability to predict long-term changes in soil N availability, and hence
whether greater NPP will be sustained in a CO2-enriched atmosphere.
ZEGLIN, LYDIA H.,* DEANA D. PENNINGTON, ROBERT R. PARMENTER, JAMES R. GOSZ and JAMES W. BRUNT. Department of Biology, University of New Mexico, Albuquerque, NM, USA. Assessing
semi-arid grassland recovery from grazing using multispectral imagery.
The Sevilleta National Wildlife Refuge (Sevilleta NWR) in central New
Mexico was established in 1974 when former ranchland was donated to
the U.S. Fish and Wildlife Service. In 1989, the site became a Long Term
Ecological Research (LTER) site. A recurring issue has been the extent to
which the semi-arid grasslands of the Sevilleta NWR reflect the legacy of
past livestock grazing. To address this question, grazing exclosure plots
were constructed in 1992 on ranchland just north of the Sevilleta boundary.
Three exclosure plots of 300 m by 300 m were established with analogs
within the Sevilleta boundary. In addition, three continually grazed plots
were located in the grazing land close to the exclosures. These nine plots
were studied using TM imagery and GIS analysis. We measured NDVI at
the nine plots for thirteen years beginning in 1989, three years before the
establishment of the exclosures. The rate and extent to which NDVI values
converge between the exclosures and the Sevilleta NWR plots serve as an
estimate for grazing recovery time on the Sevilleta NWR. Analysis of
NDVI proves a useful tool to address vegetation responses to grazing and
to analyze rates of recovery following removal of domestic grazing animals.
ZELLMER, AMADA J.* and TIMOTHY F.H. ALLEN. Department of
Botany, Birge Hall, University of Wisconsin, 430 Lincoln Drive, Madison,
WI, USA. Evaluating ecological models: Encoding, decoding and complementarity.
We have built a unified treatment of several bodies of ecological theory
from hierarchical complex systems analysis and general systems theory
(GST). Our framework cleaves cycles of model-building from cycles of
material emergence, but allows them to intersect in individual structures,
such as organisms. In one cycle a model is formed from the observation
of a structure, and its assignment to an equivalence class or type. The class
may be a level of organization. That model is verified by application to
other structures through scaled observation, to see if the other structures
meet the criterion for equivalence. The other cycle is a self-correcting system of realization of a structure from a context (e.g. though DNA for
organisms or election for a US President), followed by modification of that
context by the realized structure (e.g. though natural selection in biology,
or changes in attitude after Presidential impeachment). These analogous
cycles of reinforcement are the basis of our device for recognizing observer
presence in observation and models. With this framework, we can assist
environmental practitioners in keeping track of the layers of subjectivity

embedded in their actions. When models cease to be verified by application

to new structures (e.g. in fixed levels of chemical benchmarks accepted to
indicate toxicity), change in the other cycle leaves the model behind (e.g.
increased tolerance in natural populations). Now that we have uncovered
this pattern, we attempt to bring it into the arena of environmental application. Although the ultimate goal is application to concrete management
situations, we must first work out how one takes these abstractions and
applies them to more tangible circumstances.
ZETTLER, JENNIFER A.,1,* BRADFORD A. ROBINSON,2 CRAIG R.
ALLEN3 and TIMOTHY P. SPIRA.4 1 Department of Biology, Armstrong
Atlantic State University, Savannah, GA, USA; 2 Department of Biology
Instruction, Clemson University, Clemson, SC, USA; 3 U.S. Geological
Survey, S. C. Cooperative Fish and Wildlife Research Unit, Clemson University, Clemson, SC, USA; 4 Department of Biological Sciences, Clemson
University, Clemson, SC, USA. Ants reduce oviposition success of gulf
fritillaries on Passiflora incarnata.
Extrafloral nectaries on maypops, Passiflora incarnata, attract many ant
species including the red imported fire ant, Solenopsis invicta, and the
native ant, Crematogaster lineolata. Ants attracted to these nectaries are
known to encounter and remove herbivores. In a greenhouse located in
Clemson, South Carolina, we compared the effects of S. invicta and C.
lineolata on oviposition success and larval development in gulf fritillaries,
Agraulis vanillae, on their chief food plant, P. incarnata. We placed individual potted plants in containers with colonies of S. invicta or C. lineolata. Plants placed in empty containers served as controls. Fifty A. vanillae
pupae from a commercial breeder were allowed to emerge inside the greenhouse where we observed ant-butterfly behaviors. We found that the presence of either ant species significantly (P5 0.004) reduced the number of
eggs deposited on P. incarnata. Although not statistically significant, fewer
eggs were deposited on plants tended by S. invicta than C. lineolata. Twenty days post-oviposition, 51% of 67 eggs in the control group, 0% of 30
eggs in the C. lineolata group, and 0% of 1 egg in the S. invicta group
developed into larvae. Therefore, ant presence also reduced caterpillar survival. Because S. invicta populations are most likely to occur in fallow
fields and roadsides where P. incarnata abounds, this ant species may be
especially deleterious to gulf fritillary populations.
ZHANG, CHI,1,* HANQIN TIAN,1 SHUFEN PAN,1 JERRY MELILLO2
and DAVID KICKLIGHTER.2 1 The University of Kansas, Lawrence, KS
66045; 2 Marine Biological Laboratory, Woods Hole, MA 02543. Coupled
cycles of carbon and phosphorus in tropical scosystems: A modeling
study.
Phosphorus (P) is an important nutrient element for ecosystem production
that affects carbon (C) exchange between terrestrial ecosystems and the
atmosphere. Evidences from field experiments and observations have indicated that P is the limitation factor for terrestrial ecosystems in their late
succession period, especially for tropical rain forests growing on intensively weathered soil. This has been suggested to be caused by the low
content of soil P as well as the low availability of P fixed in soil. Our
study on 2,400 soil pedons shows that the average soil P density of the top
50 cm soil is only 830 g/m3. And the spatial distribution of P is uneven,
and the tropical region has a low supply of soil P. The P storage is also
threatened by land conversion, which could cause a mean P loss of about
2 t/ha from the terrestrial ecosystems. To study the interaction of P supply
and C sequestration processes, we have developed a submodel of the P
cycle for the Terrestrial Ecosystem Model (TEM). The P cycle is coupled
with C process by limiting gross primary production with the available P
supply, and by the feedback of net primary production on resource allocation to the P uptake. To address the problem of tropical P limitation, the
model is specially calibrated against several intensively studied tropical
rainforest sites. Then we apply the model to investigate coupled cycles of
P and C in tropical terrestrial ecosystems, and further to assess how the P
loss induced by tropical deforestation could influence ecosystem production
in tropics.

Abstracts

371

ZHANG, WEIJIAN,1,2,* JIANSHENG WU,1 CONG TU1 and SHUIJIN

HU.1 1 North Carolina State University, Raleigh, NC; 2 Nanjing Agricultural
University, Nanjing, Jiangshu, China. Divergence in microbial community structure and activities in managed ecosystems: Implications for
soil carbon dynamic.
Carbon sequestration in agricultural soils may have the potential to significantly reduce the agricultural contribution to atmospheric carbon dioxide.
Nevertheless, the magnitude of this C sequestration is unclear because the
resulting effects of alternative management practices on soil microbes and
microbially mediated processes are not well understood. We investigated
the effects of different management regimes on soil microbial community
structure and activities in six ecosystems near Goldsboro, NC. The six
adjacent and replicated ecosystems studied included a successional ecosystem, a plantation forestry-woodlot, an integrated crop-animal production
system, an organic production system and two row cropping systems with
conventional tillage or no-tillage. After 5 years of continuous treatments,
the soil microbial communities diverge, with microbial biomass, activities
and diversity being significantly higher in the ecosystems with intermediate
levels of disturbance and chemical inputs than in the ecosystems with high
or low levels. Fungal contribution to the total microbial biomass increased
as disturbance decreased except in the crop-animal production system. Increased fungal dominance coincided with a decrease in microbial respiration in ecosystems with high disturbance.These results suggest that reducing human disturbance and increasing the complexity of agroecosystems
may facilitate the development of microbial communities that are conducive to C sequestration in agricultural soils.
ZHANG, YU,1,* ANGELA M. BAKER2 and WILLIAM G. WILSON.1
1
Duke University, Durham, NC; 2 University of Oulu, Oulu, FI. The influences of ecological processes on stigma-height dimorphic species.
Species exhibiting stigma-height dimorphism are composed of two floral
morphs differing in stigma-height. In long-styled plants (L-morph), the stigma is exserted beyond the stamen, whereas in short-styled plants (Smorph), the stigma is lower. It is commonly believed that a stigma-height
dimorphism is established when each morph is more proficient at pollinating the other morph than plants of its own type and the two morphs do
not differ in the proficiency of autogamous or geitonogamous pollination.
These mating patterns will result in an isoplethic (1L:1S) equilibrium
morph ratio; however observed morph ratios in many stigma-height dimorphic species range widely from isoplethy to populations fixed for the
L-morph. We explored how the morph ratio of stigma-height dimorphic
species is affected by factors such as selfing rate, intra- and intermorph
pollination, pollen limitation, ovule fertilization probability, and population
size by using both analytical and simulation models of plant population
dynamics in an ecological context. Comparisons between our simulation
results and empirical data of self-sterile Narcissus assoanus and self-compatible N. dubius supported several predictions. For the self-sterile N. assoanus, population size-dependent L-biased morph ratios are due either to
a greater degree of disassortative mating in the L-morph and pollen limitation in large populations, or greater ovule fertilization probability in Lmorph and pollen limitation in small populations. For the self-compatible
species N. dubius, strongly L-biased morph ratios independent of population size may be due to lethality of homozygous S-morph.
ZHENG, LEI,1,* R. J. STEVENSON1 and CHRISTOPHER CRAFT.2 1 Department of Zoology, Michigan State University, East Lansing, MI;
2
School of Public & Environmental Affairs, University of Indiana, Bloomington, IN. Changes in benthic algal attributes during salt marshes
restoration and use in bioassessment.
Although salt marsh restoration has been used to mitigate loss of habitat
and ecosystem function, few effective rules and indicators have been developed to determine success of ecological restoration in salt marshes. To
assess attributes of algal assemblages as indicators of salt marsh restoration,
we chose 8 pairs of salt marshes in North Carolina, each pair with one
restored marsh (from 1 to 28 year old) and a nearby existing salt marsh.
Algae on both macrophytes and sediments were collected in each marsh
during spring and summer 1998 for assaying algal biomass (dry mass
(DM), ash free dry mass (AFDM), chl a content, algal biovolume), algal

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Abstracts

species composition and diversity, and gross primary production. An attribute restoration ratio was calculated by dividing attribute values from each
restored marsh by values from a paired reference marsh. The organic matter
restoration ratio of sediments increased with age of restored marshes in
both spring and summer. The algal biomass restoration ratios of epiphytes,
calculated with algal biovolume and chl a, increased with restored marsh
age in summer but not during spring. Biomass of sediment algae was not
related to marsh age. The species diversity of sediment algae in summer
showed an asymptotic relationship with sediment nutrient concentration.
The similarity of diatom species composition between paired restored and
reference sites increased with age of restored marshes during spring and
summer. Primary production by epiphytic and sediment algae in summer
showed site-specific changes and did not change consistently with marsh
age. Algal biomass, algal diversity, and diatom species assemblages during
summer were positively correlated with sediment nitrogen and phosphorus
concentration. We concluded that overall structural and functional development of restored wetlands, especially nutrient storage in sediments, regulates algal community structure and function, which can be used to evaluate marsh restoration.
ZHU, WEIXING* and TAO ZHANG. Binghamton University, Binghamton, New York, USA. Nitrogen retention and hydrological loss in intact
soil cores during a 24-week lab incubation.
Forests have been found retaining N inputs from human activities in many
parts of the world. Studies in the past decade suggest that soil is the largest
N-sink, through possible mechanisms of microbial immobilization, organic
matter accumulation, and abiotic retention. We tested N retention in intact
soil cores collected from forest patches along an urban-rural gradient in the
New York City metropolitan area. We hypothesized that higher nitrification
found in urban soils would lead to higher N-loss and less N-retention. We
also tested the hypothesis that leaf litter with different C-to-N ratios would
retain N differently, by adding sugar maple and red oak litters to respective
soil cores. Cores were leached weekly with a 2-cm simulated rain containing 2 mg/L NH4-N and NO3-N. Inorganic-N export increased linearly during the first 12 weeks and soils quickly shifted from net N-sinks to net Nsources. By the end of the 12th week, mean NH4-N in leachate reached
4.4 mg/L and mean NO3-N 23.9 mg/L. Urban soils had much higher NO3
export than the rural soils. We found, however, no litter effort on N retention. From weeks 13 to 20, we added 100 mg/L labile carbon to 60% of
the soil cores, and in the last 4 weeks, created temporarily hydrological
retention on these C-amended cores. We found neither carbon addition, nor
hydrological retention, had affected N-export. The cumulative net N-exports in this 24-week lab incubation spanned from 0.5 to 15.1 g m-2 and
were positively correlated to the previous field measurements of net N
mineralization and nitrification rates. The large hydrological loss of inorganic-N from these intact soil cores suggests that retention mechanisms
based on short-term soil studies need to be examined carefully in the context of a functional ecosystem containing live plants and plant-soil interactions.
ZHUANG, QIANLAI,1 JERRY M. MELILLO,1 DAVID W. KICKLIGHTER,1 RONALD G. PRINN,2 PAUL A. STEUDLER,1 DAVID A. MCGUIRE,3 BENJAMIN S. FELZER1 and SHAOMIN HU.1 1 The Ecosystems
Center, Marine Biological Laboratory, Woods Hole, MA; 2 Massachusetts
Institute of Technology, Boston, MA; 3 University of Alaska Fairbanks,
Fairbanks, AK. Modeling methane consumption and emission between
the terrestrial biosphere and the atmosphere.
Methane is an important greenhouse gas that has major natural sources and
sinks associated with the activities of soil microbes. As climate changes it
is possible that the magnitudes of these sources and sinks will change
dramatically. To explore these possible changes we have modified our global biogeochemistry model, the Terrestrial Ecosystem Model (TEM 5.0).
The modifications include a revision of our water balance and soil thermal
modules, and the development of a new methane dynamics module. The
revised water balance model enables us to consider the dynamics of ecosystem hydrology, including water fluxes, soil water content, and water
tables on a daily rather than a monthly time step. The revised soil thermal
module enables us to more accurately simulate various aspects of soil thermal dynamics in northern ecosystems including soil temperatures, active

layer depths, and permafrost dynamics on a daily time step. The new methane dynamics module has been developed to simulate the processes of
methanotrophy and methanogenesis, along with the physical and biological
transport mechanisms of this greenhouse gas. Our new version of TEM has
been run at a daily time step for a variety of sites, from tundra to tropical
forests, where methane flux measurements have been made. Preliminary
studies show good agreement between the simulated and measured fluxes
at most sites. In the future we plan to use the model to examine the methane
responses to climate change (e.g. warming and permafrost melting in the
Northern Hemisphere) and land-use and land-cover changes (e.g. agricultural activities in Brazil) at regional and global scales. We will also run
the model within the MIT Integrated Global System Model (IGSM) framework to examine the feedbacks of methane dynamics between terrestrial
ecosystems and the climate system.
ZIMMERMANN, CRAIG R.* Warnell School of Forest Resources, University of Georgia, Athens, GA. The Interaction of invasion sequence
with a productivity gradient in assembled protist communities.
Laboratory microcosm experiments in community assembly have shown
that permutating the order of species invasion can significantly impact subsequent community structure. To date, the influence of varying environmental regimes on the effects of colonization history, however, has received
only limited attention. Here, I explore the interaction of invasion sequence
with a productivity gradient by assembling multitrophic detritus-based protist communities in a laboratory setting. Ordered species introductions were
carried out under two alternative schedules against a productivity gradient
consisting of five discrete nutrient levels. Censusing of community composition was performed at periodic intervals for a total of 64 days. In
addition, a suite of chemical variables was quantified for each microcosm
at the conclusion of the experiment to assess the impact of invasion sequence on media properties. The following results were observed: 1) the
degree of overall variation in final community structure exhibited across
productivity levels was dependent on invasion order, 2) alternative community states resulting as a function of invasion sequence were only observed at the three highest productivity levels, 3) a total of five unique
community states was found to arise from the interaction of invasion sequence and productivity level, 4) the degree of variation in community
composition en route to the final state was dependent on invasion order
with the greatest variation in both sequences observed at the highest productivity levels, 5) invasion sequences were found to produce unique
changes to media chemistry.
ZIMPFER, JEFF F.,1,* DIANE WAGNER2 and STANLEY D. SMITH.1
Department of Biological Sciences, University of Nevada, Las Vegas, NV;
2
Biology and Wildlife Department, University of Alaska, Fairbanks, AK.
Effects of N deposition, increased summer rains and trampling on nitrogen fixing ability of biological crusts in the Mojave Desert.

Biological crusts play an important role in the Mojave Desert by reducing

soil erosion and fixing atmospheric nitrogen. Changing patterns of climate
and land use are likely to affect the species composition and metabolic
activity of crusts. In a 3-way factorial experiment at the Mojave Global
Change Facility (MGCF), we increased rates of nitrogen deposition (0, 10,
and 40 kg N ha-1 yr-1), summer rains (7.5 cm yr-1 irrigation), and physical
disturbance (trampling) on 96 14 x 14 m plots. After two successive years
of treatment, we measured effects on percent cover by the dominant Nfixing lichen (Collema), nitrogenase activity (via acetylene reduction, hereafter AR) and soil chemistry. Summer irrigation and trampling significantly
reduced lichen cover, whereas N addition had no effect. Irrigation decreased
soil organic matter and increased both soil and mineral N. Disturbance had
no significant effect on soil chemistry, and N addition increased mineral
N but had no effect on soil organic matter. Control plots had a mean AR
rate of 13.2 n moles cm-2 h-1. Both 40 kg of added N ha-1 and trampling
significantly decreased the rate of AR. Irrigation had no effect on AR rate,
possibly because the positive effects of increased soil moisture on AR rates
offset negative effects due to the irrigation-related decrease in areal coverage of N-fixing lichens. Our results suggest that (1) high levels of N
deposition and increased physical disturbance will decrease rates of N fixation in the Mojave Desert and (2) cover by N-fixing lichens will decrease
in response to increased summer precipitation.

ZITZER, STEPHEN F.1,* and STEVE R. ARCHER.2 1 Desert Research

Institute, stephen@dri.edu, Las Vegas, NV, USA; 2 University of Arizona,
Tucson, AZ, USA. Influence of variation in spatial and temporal light
distribution on the dynamics of leaf area biomass and nitrogen of
woody plants and their relationship to community dominance in subtropical savanna.
Leaf area biomass and nitrogen relationships can vary greatly within and
between species occurring in the same plant community. Determining environmental factors that affect these leaf relationships and the amplitude of
variation may suggest mechanisms for ecological success. We tested the
hypothesis that species with the greatest plasticity in leaf area biomass and
nitrogen characteristics will be the most successful in terms of having the
greatest frequency within the plant community. Leaves of the seven dominant woody growth-forms from a south Texas subtropical savanna were
collected from four canopy positions within vertical transects of four landscape types in summer and in winter. We measured specific leaf area (cm2
leaf area/g dry weight leaf), mean leaf biomass (mg dry wt/leaf), mean leaf
size (cm2 area/leaf) and leaf nitrogen as a function of leaf area and weight.
Zanthoxylum fagara, a drought-deciduous evergreen with compound coriaceous leaves, had the greatest leaf plasticity and the greatest frequency.
Conversely, evergreen sclerophyllous Mahonia trifoliolata and evergreen
coriaceous Schaefferia cuneifolia had the least leaf plasticity and the lowest
frequencies. Among the intermediate species, leaf plasticity decreased from
Prosopis glandulosa, through Celtis pallida, Condalia hookeri and Diospyros texana and was also correlated with decreasing frequency. The data
did not suggest a significant relationship between leaf phenology and genetic plasticity. Canopy position, season and landscape type were equally
important relative to significantly affecting leaf characteristics across all
species. Specific leaf area, mean leaf nitrogen content and specific leaf
nitrogen had the most variation, while leaf nitrogen concentration was the
most stable characteristic. Understanding dynamics in leaf biomass and
nitrogen relationships is prerequisite to estimating dynamics in foliar biomass at the ecosystem level.
ZOU, XIAOMING,1,2,* QINGYU HUANG3 and LUCY HOU.3 1 Kunming
Section, Xishuangbanna Tropical Botanical Garden, 88 Xuefu Road,
Kunming, Yunnan, P. C. China; 2 Institute for Tropical Ecosystem Studies,
University of Puerto Rico, P.O. Box 23341, San Juan, PR, USA; 3 Department of Biology, National Cheng Kung University, 1 University Road,
Tainan, Taiwan, R. O. China. Invasion of Pontoscolex corethrurus in a
tropical forest of Taiwan.
There are few tropical soil fauna as invasive as earthworm Pontoscolex
corethrurus. The invasion of P. corethrurus was widely reported in tropical
ecosystems under human influence. We found that P. corethrurus was progressively invading a tropical forest in the Najenshan National Park of
southern Taiwan. Earthworm density reached 400-600 individuals per
square meter in places with the exotic species as compared with less than
100 individuals per square meter in places without the exotic earthworms.
The invasion of P. corethrurus reduced the density and diversity of native
earthworms and enhanced the efflux of carbon dioxide from soil to atmosphere by 10-20%. Invasive P. corethrurus may alter tropical forests by
reducing soil faunal diversity and by elevating atmospheric carbon dioxide
levels.
ZYGMUNT, JENNIFER R.,* ROBERT K. BOOTH and STEPHEN T.
JACKSON. Department of Botany, University of Wyoming, Laramie, WY,
USA. Ecology of testate amoebae in Rocky Mountain peatlands and
their application to paleohydrological reconstruction.
Testate amoebae (Protozoa: Rhizopoda) are abundant and well-preserved
in Sphagnum-dominated peatlands, where their distribution is primarily
controlled by surface moisture. Recent work has demonstrated that testate
amoebae are sensitive paleohydrological indicators, and quantitative paleoclimate reconstructions with multi-decadal to centennial-scale resolution
have been obtained from North America in the western Great Lakes region.
However, baseline data describing testate amoeba ecology are needed to
support paleohydrological studies in other regions. We investigated the relationship between environmental conditions and testate amoebae in 139
samples collected from 15 Sphagnum-dominated peatlands of the central

Abstracts

373

Rocky Mountain region. Our primary objectives were to: 1) identify environmental controls on testate amoeba distribution in the region, 2) develop transfer functions to infer environmental conditions from fossil testate amoeba assemblages, and 3) compare testate amoeba assemblages and
species-environment relationships between the central Rocky Mountains
and western Great Lakes regions to assess biogeographical patterns. Our
results indicate that substrate moisture, measured as depth to the water
table, is the dominant control on assemblage composition. Assuming good
modern analogues, past changes in water-table depth can be reconstructed

374

Abstracts

from fossil assemblages with a mean error of 7.0 cm. Over 50% of the
taxa encountered in the western Great Lakes region are also found in Rocky
Mountain peatlands, and moisture preferences of these taxa are similar in
both regions. We use our calibration dataset to infer hydrological history
of a Sphagnum-dominated peatland in Yellowstone National Park. Our preliminary results suggest testate amoebae have great potential for generating
sensitive, high-resolution records of past moisture variability in the central
Rocky Mountain region.

AUTHOR INDEX
A
AARSSEN, LONNIE W. ......... 15, 268, 302
ABBOTT, ALBERT G. ...................... 183
ABDULLA, AMEER A. ....................... 5
ABER, JOHN D. ............................. 237
ABERNETHY, KATE..........................14
ABRAHAM, JOEL K........................... 5
ABRAMS, MARC D...........................35
ABU-ATTIA, FAIEZ......................... 330
ABUZEINEH, ALISA ....................... 229
ACHEE, NICOLE ............................ 279
ACKERLY, DAVID ...... 5, 32, 73, 184, 222,
302
ADAMS, HENRY D. ........................... 5
ADAMS, JULIAN B. ..........................55
ADDINGTON, ROBERT N. ............... 226
ADEMA, ERWIN B. ........................ 132
ADKISON, GREG .............................. 5
ADLER, LYNN S. .............................. 6
ADLER, PAUL R................................ 6
ADLER, PETER B. ............................. 6
AGARWAL, PANKAJ K......................66
AGRAWAL, ANURAG A. ....... 6, 170, 317
AHUMADA, JORGE A. ....................... 6
AIKIO, SAMI .................................... 7
AKINBOYEWA, OLAYENI................ 154
AKPIK, MAASAK ........................... 109
AKSHALOV, K. .............................. 255
ALABACK, PAUL .............................. 7
ALBANI, MARCO.............................. 7
ALBER, MERRYL................. 7, 235, 357
ALBERS, GAYLE L. ........................... 7
ALBERTI, MARINA ........................ 237
ALBERTSON, JOHN ..........................98
ALBRECHT, MATTHEW A................... 8
ALBRECHT, STEPHANIE ............ 51, 301
ALDEA, MIHAI .............................. 137
ALERIC, KATHERINE M. .................... 8
ALERS-GARCIA, JANICE.................... 8
ALEXANDER, HELEN M........ 8, 223, 241
ALEXANDROV, GEORGE A.............. 164
ALLAN, BRIAN.............................. 177
ALLAN, DAVE.................................31
ALLEN, ANDREW .......................... 100
ALLEN, CHRISTOPHER B. .................. 8
ALLEN, CRAIG D. ............................45
ALLEN, CRAIG R. ... 9, 100, 118, 194, 264,
332, 371
ALLEN, DAMIAN ........................... 200
ALLEN, ELIZABETH A. ...................... 9
ALLEN, JON C. .............................. 310
ALLEN, MICHAEL F..........................97
ALLEN, ROB B. ............................. 281
ALLEN, THOMAS R........................ 230
ALLEN, TIMOTHY F. H.................9, 371
ALLEN-GIL, SUSAN ....................... 109
ALLEY, KELLY .............................. 225
ALLFREE, LESLIE ............................12

ALLISON, GARY W............................ 9

ALLISON, STEVEN D. .......................10
ALLISON, VICTORIA J. .....................10
ALO, DOMINIQUE............................27
ALONSO, CONCHITA........................10
ALSUM, ESTHER M........................ 135
ALTHOFF, DONALD P...................... 283
ALTIERI, ANDREW H...................... 362
ALTIZER, SONIA.........................10, 82
ALTWEGG, RES ...............................10
ALVERSON, WILLIAM S. ..................11
ALWASH, AZZAM .......................... 322
ALWASH, SUZIE ............................ 322
AMBERMAN, KEITH D. ....................40
AMMANN, CASPAR........................ 346
AMMANN, REBECCA L.....................11
ANAND, MADHUR ...........................11
ANDELMAN, SANDY J......................41
ANDERSEN, ALAN N. ..................... 250
ANDERSEN, BRANNON .................. 205
ANDERSEN, CHRISTIAN ................. 229
ANDERSON, CHARLES W. ............... 261
ANDERSON, DEAN P. .......................11
ANDERSON, JIM E. ..........................58
ANDERSON, KRISTINA J. ..................11
ANDERSON, KURT......................11, 39
ANDERSON, LYNN L. .......................12
ANDERSON, M. REBECCA ................12
ANDERSON, P. D. ........................... 240
ANDERSON, ROGER C. ........ 14, 296, 312
ANDERSON, SEAN S. .......................12
ANDERSON, WENDY B. ........ 12, 21, 145
ANDOW, DAVID A....................140, 357
ANDREASSI, CHRIS .........................55
ANDREFOUET, SERGE .................... 284
ANDREWS, JEFFREY S. ....................13
ANDREWS, SUSAN S. .......................13
ANDRUSKIW, MARK C. ....................13
ANEKONDA, THIMMAPPA S. ........... 177
ANEKONDA, THIMMAPPA .............. 310
ANGERT, AMY L..............................13
ANHOLT, BRADLEY R. .....................10
ANHOLT, BRADLEY ....................... 126
ANSARI, SHAHIN.............................85
ANSLEY, R. JAMES ........................ 141
ANTHONY, NICOLA M. .....................14
ANTONOVICS, JANIS ..................8, 264
ANTONSEN, HILDE ..........................14
APPERSON, CHARLES S.................. 340
APPLE, JENNIFER L. ........................14
APPLE, MARTHA E. .........................14
APPLEGATE, ROGER D. .................. 358
AQUILINO, KRISTIN M. ....................14
ARBAB ZAVAR, M.H. ..................... 121
ARCHER, STEVEN R............ 15, 336, 373
AREF, SUSANNE ............................ 241
ARMITAGE, ANNA R. .......................15
ARMITAGE, KENNETH B. ............... 255

ARMSWORTH, PAUL R. ....................15

ARNOLD, HEATHER G. .....................15
ARNOLD, JENNIFER M. ....................16
ARNONE, JAY 16, 169, 271, 289, 343, 351
ARNOTT, SHELLEY E. .......... 16, 83, 326
ARP, CHRIS ................................... 254
ARROYO, MARY T.K. ..................... 247
ARTHUR, MARY A. ..................108, 367
ASCHENBACH, TODD A....................16
ASHMAN, TIA-LYNN ............ 58, 68, 320
ASHTON, ISABEL W. ........................16
ATHY, ERIN R..................................17
ATKINSON, CARTER ......................... 6
AUGSPURGER, CAROL K. ... 17, 207, 284,
292, 349
AUGUSTINE, DAVID.................161, 296
AUKEMA, JULIANN E.......................17
AULD, JOSH R.................... 17, 156, 217
AUSTIN, AMY T. ..............................18
AUSTIN, MIKE P. ........................... 341
AWADA, TALA ................................18
AYRES, DEBRA ............................. 326
AYRES, MATTHEW ................... 18, 152
B
BACHMANN, CHARLES M. ............. 292
BACKLUND, BRANT ...................... 303
BACKUS, VICKIE L. .........................18
BADE, DARREN L. .............. 18, 104, 283
BAER, NICHOLAS A. ........................19
BAIDYA ROY, SOMNATH ..................19
BAILEY, LARISSA L. ........................19
BAILEY, SCOTT W....................155, 210
BAKER, ANGELA M. ...................... 372
BAKER, MICHELLE........................ 254
BAKER, PATRICK J. .........................19
BAKER, PAUL W. .............................19
BAKKER, CHRIS ..............................20
BAKKER, LIESBETH.........................20
BALA, INDU ................................. 343
BALBACH, HAROLD E............... 20, 190
BALDWIN, IAN T. .......................... 180
BALICE, RANDY G...........................20
BALL, MARILYN C......................... 211
BALLANTYNE, ASHLEY P. ................20
BALLANTYNE, FORD .......................21
BALSER, TERI C. ........................... 233
BANAVAR, JAYNATH R. ................. 158
BANCROFT, G. THOMAS ...................21
BANKER, MICHELE G. ................... 165
BARBOUR, MARGARET M. ........ 42, 336
BARBOUR, MICHAEL G. ...........247, 315
BARD, ALICE M. ........................... 327
BARFIELD, MICHAEL..................... 187
BARNES, FAIRLEY J....................... 234
BARNES, GRENVILLE ......................34
BARNETT, DAVID T. .................309, 324
BARNEY, JACOB N...........................21

Author Index

375

BARRETO, CRISTINE...................... 111

BARRETT, J E. .................................21
BARRETT, KYLE..............................21
BARRETT, MARK A. .........................22
BARRON, STACY H. .........................22
BARRON, STACY........................... 207
BARRON-GAFFORD, GREG .... 22, 36, 78,
130, 192, 265, 348, 363, 365
BARROW, WYLIE C. .................229, 357
BARRY, MATTHEW J. ..................22, 41
BARTEL, REBECCA A.......................22
BARTLEIN, PATRICK ...................... 359
BARTON, KASEY E. .........................23
BARTUSKA, ANN .......................... 235
BARTUSZEVIGE, ANNE M.................23
BARTZ, TRACI M. ............................23
BARUCH, ZDRAVKO ........................23
BASHEY, FARRAH ...........................23
BASSIRIRAD, HORMOZ ..... 185, 251, 308
BASTOW, JUSTIN L. .........................24
BATCHELOR, MARGARET E. .............24
BATTAGLIA, LORETTA L. ............24, 69
BATTAGLIA, MICHAEL A..................24
BATZLI, JANET MCCRAY..................24
BAUER, LISA A. ..............................49
BAUERLE, TARYN L. .......................25
BAUERLE, WILLIAM L. ....................25
BAUKUS, ADAM............................ 127
BAXTER, COLDEN V. .......................25
BAZZAZ, FAKHRI A. .....25, 206, 324, 364
BEAL, ANGUS LH .......................... 243
BEARD, KAREN H..................... 25, 193
BEAUCHAMP, VANESSA B. ...............25
BECKAGE, BRIAN............................26
BECKSTEAD, JULIE .........................26
BEDFORD, BARBARA..................... 171
BEEDLOW, PETER.......................... 229
BEIER, COLIN M..............................67
BELAY, ASFAW.............................. 327
BELESKY, DAVID P......................... 136
BELK, MARK C. ....................... 26, 248
BELL, CHRISTOPHER J. ....................26
BELL, DANIELA L............................27
BELL, JOHN M. ............................. 239
BELL, SUSAN S. ............................ 318
BELL, THOMAS CD........................ 318
BELLOWS, A. SCOTT ..................... 230
BELMONT, JONATHON M. ............... 353
BELNAP, JAYNE.... 227, 278, 296, 298, 362
BELOTE, TRAVIS .............................27
BENAVIDES, MELISSA A...................27
BENBOW, M. ERIC ...........................27
BENDA, NICOLE D. ..........................27
BENDICK, ROBERT L. .................... 329
BENFIELD, FRED ........................... 118
BENJAMINS, MAIJA E. .....................28
BENNETT, ASHLEY B. ......................28
BENNETT, ELENA M.........................28
BENNETT, KYLE F. ......................... 231
BENNETT, T. M. BULL ......................28
BENNING, TRACY L. ................. 28, 294
BENNINGTON, CINDY ......................29
BENOIT-BIRD, KELLY J.....................29

376

Author Index

BENSON, GARY L. ...........................29

BERENBAUM, MAY R..................... 137
BERG, ED .......................................29
BERGEN, LYDIA ............................ 212
BERGER, CARRIE .......................... 301
BERGER, MICHELLE L. ....................30
BERGNER, BELLE............................30
BERISH, CORY ......................... 30, 273
BERKOWITZ, ALAN R. ........ 30, 109, 174
BERKSON, JIM ........ 30, 82, 132, 161, 227
BERLOW, ERIC ................................80
BERN, CARLETON R. ..................... 336
BERNAL, ROCIO..............................31
BERNHARDT, EMILY........................31
BERRY, JOSEPH A. .............. 31, 207, 338
BERTNESS, MARK D. ....31, 245, 266, 308
BESSEY, K. MICHAEL..................... 118
BETHERS, SUZANNE........................31
BETHMANN, CORY..........................12
BEVER, JAMES D...... 8, 32, 142, 345, 356
BEVIS, KEVIN P. ....................... 32, 333
BEYER, DEAN E. .............................11
BHASKAR, RADIKA .........................32
BIEBIGHAUSER, TOM .................... 285
BIERWAGEN, BRITTA G....................32
BIGELOW, SETH W. .....................32, 56
BIGGERS, ANDREW D. .....................33
BIGLER, WENDY .............................33
BIL, K Y. ...................................... 192
BILLOCK, ARLENE G. ......................33
BILYEU, DANIELLE M. .....................33
BINCKLEY, CHRISTOPHER A. ............33
BINFORD, MICHAEL W. ............. 34, 319
BINKLEY, DAN ................................34
BIONDINI, MARIO E. ...................... 237
BIRD, CHRISTOPHER E.....................34
BIRD, SUZANNE ..............................34
BISHOP, BLAIR B. ............................34
BISHOP, JOHN G. .............................14
BJORN, ANDREW M. ...................... 237
BJORNDAL, KAREN A. .....................34
BJRNSTAD, OTTAR N. ............160, 207
BLACK, BRYAN A............................35
BLACK, R. ALAN..........35, 110, 146, 259
BLACK, ROSS ............................... 320
BLAGODATSKII, S A. ..................... 192
BLAIR, LAUREN P. ...........................35
BLAIR, ROBERT B............................35
BLAKE, JOHN ............................... 105
BLANCHETTE, CAROL .............. 39, 275
BLAUSTEIN, ANDREW R............ 35, 298
BLEDSOE, CAROLINE S. ...........144, 275
BLISS, KRISTIN M............................36
BLOOD, ELIZABETH R......... 36, 153, 273
BLOSSEY, BERND .......................... 215
BLUM, JOEL D. ............................. 367
BLUM, MICHAEL J...........................36
BLUMENTHAL, DANA M. .................36
BOBICH, EDWARD G. ........... 22, 36, 130
BOCKHEIM, JAMES G..................... 208
BOECKLEN, WILLIAM J. ............ 37, 368
BOELMAN, NATALIE T. ............. 37, 130
BOERNER, RALPH EJ ................ 37, 310

BOETTNER, GEORGE H. ................. 261

BOGGS, JOHNNY L. ....................... 260
BOHANAN, ROBERT E......................37
BOHANNAN, BRENDAN J. M. ..... 57, 154
BOHLMAN, STEPHANIE A............... 221
HNING-GAESE, KATRIN................53
BO
BOLKER, BENJAMIN M. ...... 38, 224, 345
BOLLENBACHER, WALTER E. ......... 247
BOLSTAD, PAUL............................ 118
BOLTEN, ALAN B. ...........................34
BOLTON, GARY H............................38
BOND, BARBARA J. .................120, 242
BOND, WILLIAM J. ................... 75, 330
BOND-LAMBERTY, BEN P................ 103
BONELLO, PIERLUIGI .................... 195
BONNET, VERONIQUE H. ........... 38, 301
BOOTH, DONALD C. ........................75
BOOTH, ROBERT K. .................. 38, 373
BOOTS, MIKE................................ 158
BORER, CATHERINE H. ....................38
BORER, ELIZABETH.........................39
BORG, CHRISTOPHER .................... 189
BORNER, ANDREW........................ 347
BOROWICZ, VICTORIA A. .................39
BORRETT, STUART R. ............... 39, 262
BORSUK, MARK E. ..........................39
BOS, DARREN ............................... 265
BOSCO, G .......................................58
BOSSART, JANICE............................39
BOSWELL, GANTT...........................40
BOTHWELL, SARA G...................... 204
BOUCHARD, VIRGINIE ................... 122
BOUDELL, JERE A. ..........................40
BOULTON, APRIL M. ........................40
BOUNOUA, LAHOUARI .................. 162
BOURDAGHS, MICHAEL................. 171
BOURG, NORMAN A. .......................40
BOUTIN, ROBERT .......................... 197
BOUTTON, THOMAS W. ............. 15, 141
BOVA, ANTHONY S. ........................40
BOWEN, MARK ............................. 202
BOWERS, M. DEANE ........................23
BOWERS, RACHEL N........................41
BOWERS, RICHARD W. .....................41
BOWLES, CHRISTY M. .....................41
BOWLING, ANNA M. ........................41
BOWLING, DAVID R............. 31, 41, 212
BOWMAN CUTWAY, HEATHER ..........42
BOWMAN, WILLIAM D. ....................78
BOWMAN, WILLIAM P. .....................42
BOX, DAVID O. ............................. 136
BOYCE, CARLA J........................... 159
BOYCE, RICHARD L.........................42
BOYD, ROBERT S. .............. 42, 168, 239
BOYER, ALISON G. ..........................42
BOYER, KATHARYN E......................42
BOYLE, BRAD............................... 352
BRACHO, ROSVEL .................... 43, 271
BRADBURN, MEGAN .......................68
BRADFORD, JOHN B. .......................43
BRADLEY, KATE L. ..........................43
BRADSHAW, G. A.............................43
BRADY, VALERIE .......................... 251

BRAGG, DON C. ..............................43

BRAKE, ROBIN A.............................44
BRAKS, MARIETA A.H.................... 173
BRAMBLE, DENNIS M. .....................44
BRANDT, STEPHEN B. .................... 153
BRANDTBERG, TOMAS .................. 196
BRANTLEY, SANDRA ..................... 176
BRASSIL, CHAD E............................44
BRAULT, SOLANGE..........................16
BRAUNING, DANIEL W. .................. 221
BRAY, SARAH R. .............................44
BREEN, AMANDA N.........................44
BREMIGAN, MARY T. ..................... 317
BRESHEARS, DAVID D. 20, 45, 124, 169,
178, 234, 278, 314, 324
BRETT, MICHAEL T. .........................20
BREWER, CAROL A. ................. 46, 267
BREWER, CAROL ..................... 45, 314
BREWER, ELIZABETH A. ..................45
BREWER, STEPHEN .........................45
BREWER, STEVE .....................126, 152
BREWSTER, AMANDA L. ................ 289
BREZINSKI, BRIAN ..........................46
BRIDGHAM, SCOTT D. ................... 178
BRIGGS, CHERYL ..................... 80, 245
BRIGGS, JOHN M. ....... 130, 146, 252, 297
BRIGHT, KERRY L............................46
BRINKMAN, JENNIFER A. .......... 37, 310
BRINKMAN, ROBERT B. ...................46
BRISBIN, I. LEHR.............................46
BROCKWAY, DALE G. ......................46
BRODERSEN, CRAIG R. ....................47
BROECKELMAN, B M..................... 223
BROECKER, WALLACE S. ............... 197
BROITMAN, BERNARDO R. ........ 39, 298
BRONSTEIN, JUDITH L. .................. 250
BROOKER, ROB............................. 211
BROOKS, KATHERINE E. ..................47
BROOKS, MATTHEW L. ............227, 362
BROOKS, MELODY A. .......... 47, 83, 218
BROOKS, RENEE ........................... 229
BROOKS, ROBERT P. ...................... 349
BROTHERS, AMANDA .................... 175
BROUSSARD, DAVID........................90
BROWN, BETTY W. ........................ 247
BROWN, BRYAN L. ..........................47
BROWN, JAMES H. .......45, 168, 297, 357
BROWN, JOEL S. ...................... 51, 240
BROWN, JOHN B. ............................47
BROWN, KIM J. ............. 48, 63, 221, 336
BROWN, NINA A. ............................48
BROWN, REBECCA L. ......................48
BROWN, ROGER W. ....................... 336
BROWN, TERRY ......................171, 251
BROWN, THOMAS A. ..................... 249
BROWNE, DAVID R. .........................48
BRUFORD, MICHAEL .......................14
BRUGAM, RICHARD ...................... 343
BRUHN, JOHANN N..........................48
BRULAND, GREGORY L............. 49, 142
BRUNA, EMILIO M...........................49
BRUNKOW, PAUL E.............. 49, 94, 145
BRUNNER, JESSE L. .........................49

BRUNO, JOHN F. ..............42, 48, 50, 362

BRUNS, MARY ANN.........................53
BRUNT, JAMES W........................... 371
BRYANT, PABLO..............................63
BRYCE, THOMAS D. ...................... 328
BUBIS, JOSE ...................................23
BUCHMANN, STEPHEN L. ............... 262
BUCK, TRACY L. ........................... 226
BUCKLEY, DAVID .......................... 301
BUCKLEY, LAUREN B. .....................50
BUCKLEY, THOMAS N......................50
BUEHLER, DAVID .......................... 324
BUENO, RUDY ......................... 82, 143
BUI, LINDA................................... 288
BULL, C. MICHAEL..................316, 339
BULLEN, THOMAS......................... 270
BULTMAN, THOMAS L. ....................50
BUNKER, DANIEL E. ................. 50, 319
BURDON, JEREMY J....................... 311
BURGER, LOREN W..........................71
BURGESS, ANN ...............................24
BURKE, ANNA M.............................51
BURKE, INGRID C....... 6, 43, 51, 136, 137
BURKE, JOSEPH ............................ 292
BURKE, LAURETTA ....................... 284
BURKE, MARIANNE K. ................... 142
BURKEPILE, DERON E......................51
BURLEYSON, TRAVIS................ 51, 301
BURNS, SARAH ...............................94
BURTON, MICHAEL G. .....................51
BURTON, MICHELE L. .................... 209
BURTON, WILLODEAN .....................46
BUSHAW-NEWTON, KAREN L............52
BUTCHER, LADEAN M. ....... 52, 100, 236
BUTCHER, MARYN ..........................73
BUTLER, MARK J. ......................... 200
BUTTERFIELD, SCOTT .....................52
BYERS, JAMES E. ............................52
BYRD, KRISTIN ...............................53
BYRNE, LOREN B. ...........................53
C
CABEZAS, HERIBERTO................... 221
CABLE, JESSICA M. .........................53
CACERES, CARLA E.........................53
CADE, BRIAN S. ..............................54
CADE-MENUN, BARBARA .............. 346
CADENASSO, MARY L......... 54, 232, 302
CADOTTE, MARC W. ........................54
CALABRESE, JUSTIN M. ...................54
CALCOTE, RANDY.................... 55, 213
CALDWELL, MARTYN M. .. 201, 265, 293
CALEF, MONIKA P............................55
CALLAHAM, MAC A. .......................37
CALLAWAY, RAGAN M. ...... 55, 211, 251,
279, 335
CALLOWAY, RAY........................... 134
CAMACHO, FRANK A.......................55
CAMARGO, PLINIO ........................ 338
CAMERON, TOM C......................... 351
CAMILL, PHIL .................................55
CAMPBELL, DANIEL ...................... 288
CAMPBELL, PETYA E. .................... 236

CAMPBELL, PETYA K.......................73

CANDAU, JEAN-NOEL .................... 108
CANHAM, CHARLES D. ........ 32, 56, 259
` N .................58
CANO-SANTANA, ZENO
CAPLAN, J. S...................................56
CAPPUCCINO, NAOMI .................... 194
CARBONE, CHRIS .......................... 168
CARDELUS, CATHERINE............ 56, 350
CARDILLE, JEFFREY A. ....................56
CARDINA, JOHN............................ 316
CARDINALE, BRADLEY J.............14, 56
CARDON, ZOE G...........57, 118, 149, 154
CAREY, ANDREW B. ........................17
CARNEY, KAREN M. ........................57
CARPENTER, GEOFF ...................... 176
CARPENTER, STEPHEN R....... 18, 28, 56,
104, 120, 328
CARR, ANN B..................................90
CARR, DAVID E. ................. 57, 101, 164
CARR, MARK H. ..............................57
CARR, SUSAN ............................... 268
CARRIERE, YVES .......................... 309
CARRINGTON, MARY E. ...................57
CARROLL, AMY ............................ 347
CARSON, WALTER P. ..... 50, 69, 290, 319,
322
CARSTEN, L D.................................58
CARTER, DERB S. ............................58
CARTER, GREGORY A. .....................58
CARTER, JACOBY............................73
CARTER, ROBERT E. ...................... 307
CARVAN, MICHAEL J. .................... 334
CASE, ANDREA ...............................58
CASELLE, JENNIFER ........................57
CASLAVKA, CURT ...........................24
CASLER, CLARK L......................... 352
N .........58
CASTELLANOS-VARGAS, IVA
CASWELL, HAL............................. 312
CATLIN, DANIEL H. .........................59
CAVALCANTI, GUADALUPE G...........59
CAYLOR, KELLY K. .........................59
CHADWICK, OLIVER...................... 344
CHAI, NAMYI................................ 181
CHAINE, ALEXIS S...........................59
CHALCRAFT, DAVID R. ....................59
CHAMBERS, CHRIS..........................60
CHAMBERS, JEFFREY Q. ........... 60, 338
CHAN, FRANCIS ....................... 60, 233
CHANDY, SHIBI...............................60
CHANETON, ENRIQUE J............. 61, 222
CHANEY, MAXINE L. ..................... 191
CHANEY, PHILIP L. ..... 146, 209, 301, 326
CHANEY, PHILLIP .......................... 225
CHANG, YIN ................................. 106
CHAPMAN, CAROL ..........................61
CHAPPELKA, ART H.........................61
CHAPPELLE, EMMETT W. .......... 52, 100
CHARNOV, ERIC L. ........................ 297
CHARRON, ISABELLE ......................61
CHASE, JONATHAN M. .............. 61, 170
CHAUDHARI, H. K. ..........................62
CHAUDHARY, BALA ........................32
CHEDIACK, ARIADNA......................62

Author Index

377

CHEESEMAN, JOHN M.................17, 62

CHELIUS, MARISA......................... 242
CHEN, HAN ....................................62
CHEN, JANET..................................62
CHEN, JIAXIN ............................... 266
CHEN, JIQUAN ................................63
CHEN, XIN......................................63
CHEN, XIONGWEN ..........................63
CHENG, SONG.................................79
CHENG, YUFU.................................63
CHESSON, PETER L.... 63, 81, 220, 304, 314
CHIANG, JYH-MIN ...........................63
CHIARIELLO, NONA R.................63, 94
CHIAVELLI, DEBORAH A. .................64
CHICK, JOHN H. ....................... 64, 123
CHILDS, JAMES E. ......................... 292
CHILDS, JAMES............................. 277
CHIVERS, DOUG..............................35
CHMURA, GAIL L. ......................... 325
CHO, YONG-JOO......................182, 201
CHOCZYNSKA, JOANNA...................64
CHOI, HONG KEUN........................ 182
CHOI, YOUNG D. ........................... 124
CHOJNACKY, DAVID C. ....................64
CHOLER, PHILLIPE ........................ 211
CHOO, JOHANNA ............................64
CHRIST, MAJA M. .......................... 348
CHRISTENSEN, NORMAN L. ..............65
CHRISTIAN, ALAN D. .......................41
CHRISTIAN, ROBERT R................... 262
CHRISTY BOWLES, SANDY J.
ANDELMAN, ...................................99
CHU, YONG-KYU........................... 229
CHULUUN, TOGTOHYN .................. 255
CHUNG, HAEGEUN ..........................65
CIPOLLINI, DON ..............................65
CIPOLLINI, MARTIN L. .....................65
CIRTAIN, MARGARET C....................66
CIRUNA, KRISTINE ........................ 235
CISNEROS DOZAL, LUZ MARIA....... 338
CIVILLE, JANIE C. ...................332, 339
CIVILLE, JANIE ............................. 326
CLANTON, KEITH B. ........................66
CLARK, CHRIS .............................. 104
CLARK, DAVID B.............................66
CLARK, DEBORAH A. ......................66
CLARK, JAMES S. .. 66, 162, 194, 230, 364
CLARK, JAMES ...............................67
CLAUSNITZER, DAVE..................... 255
CLAY, KEITH...... 8, 67, 171, 190, 258, 279
CLELAND, ELSA E. ..........................67
CLEMENT, JEAN-CHRISTOPHE...........97
CLESCERI, ERIKA J........................ 127
CLEVELAND, CORY C. ................... 336
CLIFFORD, STEPHEN .......................14
CLINE, GEORGE ............................ 307
CLINTON, BARTON D.......................67
CLOUGH, JONATHAN S. ................. 260
COAD, BRIAN ............................... 322
COBB, NEIL ............................. 89, 176
COBBOLD, CHRISTINA A. .................67
COCHRAN-STAFIRA, LIANE ..............68
COCRHAN, KATHY ........................ 242

378

Author Index

COE, MICHAEL T. ............................56

COFFEY, KIM ..................................68
COHEN, WARREN B. ...................... 254
COLAUTTI, ROBERT I.......................68
COLBOURNE, JOHN K. .....................68
COLE, DENISE H..............................68
COLE, JON J.................................. 283
COLE, JONATHAN J................... 18, 340
COLE-CHRISTENSEN, DARRYL........ 135
COLEMAN, DAVID C. ........ 215, 263, 368
COLEMAN, FELICIA.........................76
COLEMAN, JAMES S. .......................16
COLEMAN, MARK D. ..8, 69, 75, 105, 324
COLEY, PHYLLIS D. .................. 69, 107
COLLIER, SHAWN............................69
COLLINGE, SHARON K. .................. 120
COLLINS, BEVERLY S. ..... 69, 75, 89, 92,
94, 300, 305
COLLINS, BEVERLY....................... 215
COLLINS, JAMES P. ..........................49
COLLINS, RACHEL J. .......................69
COLLINS, SCOTT L. ............ 70, 297, 312
COMAS, LOUISE..............................70
COMERFORD, NICHOLAS B. .............76
COMITA, LIZA S. .............................70
CONANT, RICHARD T. ............... 70, 251
CONARD, SUSAN........................... 315
CONNELL, JOSEPH H. .................... 129
CONNELLY, SCOTT J. ..................... 147
CONNER, CRISTIN A. .......................70
CONNOLLY, SEAN R..................... 5, 92
CONOVER, ROSS R. .........................71
CONTRERAS, THOMAS A................ 307
CONTRERAS, TOM A...................... 320
COOCH, EVAN G. .......................... 155
COOK, ED..................................... 264
COOK, JESSICA L. ...........................71
COOK, MICHELLE ......................... 268
COOK, ROBERT...............................71
COOKE, TREY .................................71
COOLEY, HEATHER S. ......................71
COOLEY, HILLARY C. ......................71
COOPER, DAVID J. .................... 33, 363
COOPER, ROBERT J..........................72
COOPER, SCOTT..............................39
COOPERBAND, LESLIE R. .................72
COPENHEAVER, CAROLYN A. .... 72, 115
CORBIN, JEFFERY D. ................... 5, 72
CORDELL, SUSAN ......................... 257
CORNELISSEN, TATIANA G. ..............72
CORNWELL, WILLIAM K. ............. 5, 73
CORP, LAWRENCE A. ..... 52, 73, 100, 236
COSGRIFF, ROBERT J. ......................73
COSTANZO, KATIE S. ................ 73, 180
COSTELLO, ANDRIA M.....................74
COSYLEON, GABRIEL .................... 302
COTHRAN, RICKEY D. ................... 353
COTTINGHAM, KATHRYN L. 64, 74, 284
COUGENOUR, MICHAEL B. ............. 315
COUILLARD, DAVID M. .................. 325
COULSON, TIM ...............................74
COVICH, ALAN P. .......................25, 74
COX, MARY.................................. 349

COX, ROBERT M. .......................... 231

COX, STEPHEN B........................74, 90
COYLE, DAVID R. ........................ 8, 75
CRABTREE, ROBERT L. .................. 237
CRAFT, CHRIS B. ........................... 178
CRAFT, CHRISTOPHER ................... 372
CRAIG, BRODERSEN ...................... 121
CRAIG, JOHN A. ............................ 356
CRAIG, RACHAEL G.........................75
CRAINE, JOSEPH M..........................75
CRAWFORD, CHRISTOPHER B. ..........75
CRENSHAW, CHELSEA L................. 109
CRIDDLE, RICHARD S. .............177, 335
CRISAFULLI, CHARLIE M. .............. 228
CRISMAN, THOMAS L. ................... 322
CROMARTIE, WILLIAM J. .................75
CROPPER, WENDELL P. ....................76
CROSS, ALISON...............................76
CROTEAU, MAXINE ....................... 338
CROW, THOMAS............................ 301
CROWDER, LARRY B. ......................76
CROWL, TODD A. .......................25, 74
CROWLEY, PHILIP H. ..................... 285
CROXALL, JOHN P. ..........................16
CROZIER, LISA G.............................76
CRUMP, BYRON C.......................... 173
CRUMRINE, PATRICK W....................76
CRUZAN, MITCHELL B. .................. 319
CRYER, GREG T...............................97
CULPEPPER, A. STANLEY .................51
CUMMING, BRIAN F. ...................... 200
CUMMING, GRAEME S. ...............28, 76
CUMMING, JONATHAN R.... 77, 178, 186,
240
CUPP, EDDIE W.............................. 340
CURRAN, LISA M. ......................... 226
CURRAN, MARY C.................... 77, 115
CURRIE, DAVID J........................... 304
D
DABUNDO, DENISE M. ....................77
DANTONIO, CARLA M................. 5, 72
DANTONIO, CARLA ................. 80, 144
DAVANZO, CHARLENE....... 77, 128, 332
DODORICO, PAOLO ........................98
DSA, JEAN V. .................................78
DA SILVA, ROSEANA P. ....................60
DAEHLER, CURT C. .........................85
DAIL, BRYAN................................ 237
DAILEY, THERESA B. .......................78
DAILY, GRETCHEN C... 209, 222, 291, 304
DALLING, JAMES W. ...................... 117
DAMM, MARY C..............................78
DAMSCHEN, ELLEN............ 78, 257, 287
DANE, LAURA ................................78
DANEHY, BOB.................................60
DANG, QING-LAI .............................79
DANG, QINGLAI ............................ 266
DANIEL, F. BERNARD..................... 131
DANZ, NICHOLAS P................... 79, 251
DAOUST, ROBERT J..........................79
DARCY, TARA L. .............................79
DARNELL, TRACI M.........................79

DARROUZET-NARDI, ANTHONY ........80

DAS, ABHIK....................................80
DASZAK, PETER..............................80
DAUFRESNE, TANGUY ................... 228
DAUGHERTY, MATT.........................80
DAVE, NILAM B. ........................... 151
DAVELOS, ANITA L..........................80
DAVEY, PHILLIP ............................ 200
DAVIDSON, ANA D. .........................81
DAVIDSON, CARLOS ........................81
DAVIDSON, ERIC .....................217, 338
DAVIDSON, JENNIFER M. .................81
DAVIDSON, R. SCOTT ......................81
DAVIES, KENDI F. ............................81
DAVIS, ANDREW K. ....................10, 82
DAVIS, ANTHONY M. ..................... 108
DAVIS, DAYTON K...........................82
DAVIS, HEATHER G. ........... 82, 326, 332
DAVIS, KARA A............................. 197
DAVIS, MARK A. .............................82
DAVIS, MICHELLE ......................30, 82
DAVIS, STEPHEN D. .......... 102, 166, 272
DAVIS-BORN, RENEE ..................... 212
DAWSON, AMY C. ...........................83
DAWSON, TODD E. ..........................72
DAY, FRANK P. ................. 258, 325, 363
DAY, JONATHAN F.......................... 210
DAYEM, KATHERINE E. .................. 124
DE CAMINO-BECK, TOMAS ............ 364
DE LUCIA, EVAN H..........................83
DE SZALAY, FERENC A. ...................41
DEANGELIS, DONALD L. ................ 263
DEATON, RAELYNN ............. 47, 83, 218
DEBELJAK, MARKO.........................83
DEBIASE, ADRIENNE E.....................84
DEBOER, GEORGE E. .......................84
DEBUSK, WILLIAM F. ..................... 272
DECHANT, TIMOTHY P. ....................23
DEFOREST, JARED L. .......................84
DEGLORIA, STEPHEN..................... 181
DEGROOT, KRISTIN .........................46
DEHAYES, DONALD H......................38
DEL GROSSO, STEPHEN J. ................84
DELGADO, CLAUDIA .......................34
DELGROSSO, STEVE ...................... 262
DELL, ANTHONY I. ..........................85
DELLINGER, THOMAS......................34
DELONG, ANGELIQUE T. ................ 327
DELPH, LYNDA F. ............................85
DELUCIA, EVAN....86, 108, 137, 187, 200,
241, 271, 318
DEMYANOVA, E G. ........................ 192
DENDY, SHAUNA P......................... 223
DENG, XIANGZHENG ..................... 259
DENNING, A. SCOTT ...................... 273
DENNIS, ANN................................ 247
DENNO, ROBERT F. ...... 85, 107, 124, 150,
158, 205
DENSLOW, JULIE S. .........24, 85, 87, 159
DENSLOW, NANCY D. .................... 147
DEPRIEST, TIMOTHY T. ....................85
DERMODY, ORLA ..................... 86, 137
DERNER, JUSTIN D. ....................... 269

DERR, KATHARYN D........................86

DERRICKSON, ELISSA M. .................86
DESROCHERS, RACHELLE E. ............86
DETLING, JAMES K..................128, 142
DEVANNA, KRISTEN........................86
DEVIVO, JOE ................................ 273
DEVOLDER, ANDREW......................29
DEWALT, SAARA J. ..........................87
DHONDT, ANDRE A........................ 155
DIAMANT, ADAM .......................... 112
DIAZ, ALEX .................................. 129
DIAZ, SANDRA................................87
DIBBLE, ERIC D. ........................... 310
DICK, CARL W. ............................. 229
DICKINSON, MATTHEW B.................40
DICKMAN, ELIZABETH ....................87
DICKSON, TIMOTHY L. ............. 87, 110
DIEFENBACH, DUANE R. ................ 221
DIEHL, SEBASTIAN.....................11, 87
DIEKMANN, LUCY O........................88
DIEMER, KATHERINA E....................88
DIETZE, MICHAEL C. ........... 66, 88, 364
DIEZ, JEFF M. .................................88
DIJKSTRA, FEIKE A. ........................88
DILUSTRO, JOHN J...... 75, 89, 92, 94, 325
DIRZO, RODOLFO..................... 89, 216
DITOMASO, JOE ............................ 306
DITOMMASO, ANTONIO ...................21
DITTBRENNER, BENJAMIN J. ............89
DIVITTORIO, CHRISTOPHER T. ..........89
DIXON, JEREMY..............................89
DIXON, KEN R. ..........................74, 90
DIXON, PHILIP ........................113, 263
DIXON, ROBERT M. .........................90
DOATY, SARAH ...............................52
DOBEL, HARTMUT G. ......................85
DOBLAS-MIRANDA, ENRIQUE...........90
DOBROWOLSKI, WENDY M. .............90
DOBSON, ANDREW P....... 6, 90, 155, 223,
303
DOBSON, F. STEPHEN.......................90
DODSON, STANLEY I. .................... 173
DOMINGUES, TOMAS F.....................91
DONAHUE, MEGAN J. ......................91
DONALDSON, JACK R. .....................91
DONOHUE, KATHLEEN................... 354
DONOVAN, ARTHUR J. ................... 262
DONOVAN, LISA A. .................280, 287
DORN, NATHAN J. ...........................91
DORNBUSH, MATHEW E...................91
DORNELAS, MARIA .........................92
DORSEY, ANN M............................ 132
DOS SANTOS, JOAQUIM ...................60
DOTT, CYNTHIA E. ..........................92
DOWNES, BARBARA J.................... 196
DOWNING, AMY L. ..........................86
DOZIER, HALLIE ........................... 288
DRAGOO, JERRY W. ....................... 290
DRAKE, B. G. ................................ 206
DRAKE, BERT G. ...43, 200, 258, 271, 296,
325, 363
DRAKE, DANA .............................. 181
DRAKE, JAMES A. ...........................92

DRAKE, JASON B.............................92

DRAKE, JOHN .......................... 99, 204
DRAKE, SARA J...............................92
DRENOVSKY, REBECCA E.................93
DRESS, WILLIAM J. .................. 93, 346
DREWA, PAUL B..............................93
DRIJBER, RHAE...............................43
DRINKWATER, LAURIE E..................93
DUBAYAH, RALPH O........................92
DUBERSTEIN, JAMIE A.....................94
DUDA, JEFFERY J. ............ 127, 190, 192
DUFFIE, CAROLINE V. ......................94
DUFFY, J. EMMETT ................... 42, 112
DUKES, JEFFREY S. .................. 94, 154
DUNCAN, LISA K................... 75, 89, 94
DUNCAN, RICHARD P..................... 265
DUNCAN, WILLIAM ....................... 235
DUNKER, KRISSY J. .........................94
DUNMORE, ROBYN A. ................... 332
DUNNE, THOMAS .......................... 217
DURBIN, KATIE...............................94
DUSEK, MARSHA ....................210, 356
DUSSOURD, DAVID E. .................... 147
DUURSMA, REMKO A. .....................95
DWIRE, KATHLEEN A.......................95
DYBZINSKI, RAY........................... 104
DYER, SUSAN A. .............................95
DZIALOWSKI, ANDREW R. ...............95
E
EASON, THOMAS ..................... 89, 244
EATON, JAMES M. ...........................95
EATOUGH JONES, MICHELE..............96
EBERT-MAY, DIANE .........................96
EBINGER, MIKE H. ........................ 278
ECHT, CRAIG S. ............................. 217
ECONOMO, EVAN P................... 96, 100
EDWARDS, J ...................................96
EDWARDS, KATRINA J. ....................96
EGAN, SCOTT P. ..............................97
EGERTON, JOHN J.......................... 211
EGERTON-WARBURTON, LOUISE M. ..97
EHLERINGER, JAMES R. ............ 91, 174
EHRENFELD, JOAN G. ......................97
EHRHART, LLEWELLYN M. .......268, 283
EHRLICH, PAUL R............. 209, 291, 304
EILTS, ALEX ...................................97
EISSENSTAT, DAVID....................96, 97
EITARO, WADA ............................. 150
ELLE, ELIZABETH ......................... 140
ELLIOTT, KATHERINE J. ...................98
ELLIS, ALICIA M. ............................98
ELLIS, JULIE C. ............................. 104
ELLISON, AARON M. ................ 98, 126
ELLNER, STEPHEN P. ................ 98, 370
ELSA E. CLELAND, KAREN CARNEY, ..............................................99
ELSER, JAMES J. ........................... 180
ELSHAFEI, GHDA .......................... 330
ELSKUS, ADRIA A. ........................ 285
EMANUEL, RYAN ............................98
EMERSON, DAVID ............ 123, 231, 250
EMERY, SARAH M. ..........................99

Author Index

379

EMLEN, JOHN M......................127, 192

ENDO, MASAHIRO...........................99
ENGEL, E. CAYENNE........................99
ENGEL, VICTOR C. ..........................99
ENGELBRECHT, BETTINA M.J.......... 193
ENGLE, DAVID M. ...................140, 179
ENGLISH-LOEB, GREG ................... 253
RAN V. ............... 99, 326
ENGLUND, GO
ENGSTROM, DAN R. ...................... 147
ENQUIST, BRIAN J. 45, 96, 100, 180, 224,
271, 273, 352
ENRIGHT, STEPHANIE ......................65
ENSIGN, WILLIAM E. ..................... 354
ENTCHEVA CAMPBELL, PETYA K.... 100
ENTCHEVA-CAMPBELL, PETYA.........52
EPPERSON, BRYAN K..................... 217
EPPERSON, DEBORAH M. ............... 100
EPPES, MARTHA C......................... 222
EPSTEIN, HOWARD.. 55, 98, 100, 106, 177
EPSTEIN, LYNN ............................. 361
ERICKSON, JOHN E........................ 101
ERICSON, LARS.......................311, 326
ERNEST, MORGAN ......................... 101
ERNST, W.H.O. .................................20
ERSKINE, JENNIFER A.................... 101
ERWIN, PATRICK M. ................101, 333
ESTES, BECKY L. ............................46
ETCHEMENDY, SHERI ....................... 9
EUBANKS, MICKY D. 101, 143, 160, 164,
168, 175, 327
EVANGELISTA, PAUL H. ................. 174
EVANHOE, L. ................................ 245
EVANS, CELIA ...........................47, 96
EVANS, CYNTHIA A....................... 102
EVANS, DANAE A.......................... 102
EVANS, EDWARD W. ...................... 281
EVANS, MIKE ............................... 322
EVANS, R.D. ...........................102, 298
EVINER, VALERIE T. ...................... 102
EWANCHUK, PATRICK J. ................ 266
EWE, SHARON ML......................... 102
EWERS, BRENT E. ......................... 103
EWERS, FRANK W. ..................166, 272
F
FADEN, MIKE................................ 103
FAGAN, MATTHEW E. .................... 103
FAGAN, ROXANE .......................... 311
FAGAN, WILLIAM F........ 14, 54, 103, 180
FAHEY, TIMOTHY J. .................. 74, 277
FAHRIG, LENORE .......................... 228
FALK, DONALD A.......................... 103
FALKOWSKI, PAUL G. ..............104, 208
FALL, ANDREW............................. 300
FANG, WEI ................................... 104
FANKHAUSER, CRYSTAL L. ............ 104
FARGIONE, JOE ............................. 104
FARINA, JOSE M............................ 104
FARLEY, JAQUELINE...................... 273
FARQUHAR, GRAHAM D. .................50
FARRIS, MARIANNE G.................... 105
FASTIE, CHRISTOPHER L. ............... 209
FASY, ELIZABETH ...........................35

380

Author Index

FATH, BRIAN D. ............................ 105

FAUSCH, KURT D. ...........................25
FAUTH, JOHN E. ....................... 88, 105
FAY, JOHN P. ................................. 252
FAY, PHIL A. ................................. 223
FEAGIN, R. A. ............................... 105
FEARON, MATTHEW ........................92
FEHMI, JEFFREY S. ..................151, 279
FEI, SONGLIN ............................... 105
FELDMAN, DAVID L....................... 105
FELDMAN, TRACY S. ..................... 106
FELDMANN, REINER...................... 299
FELDT, JOHN ................................ 273
FELGENHAUER, BRUCE E............... 330
FELLER, ILKA C. ........................... 102
FELS, JOHN E................................ 197
FELZER, BENJAMIN S. ................... 372
FEMINELLA, JACK W. ....... 216, 236, 239
FEMINELLA, JOHN W. .................... 146
FERAL, CHRISTIE .......................... 106
FERNER, MATTHEW C.................... 106
FERRARI, JAMES B. ....................... 106
FERRARI, MATTHEW J. .................. 106
FERTIG, WALTER F......................... 107
FESSENDEN, JULIANNA E............... 178
FETTIG, CHRISTOPHER J. .................75
FIELD, C. B. ....................................94
FIELD, CHRISTOPHER B. ......63, 67, 107,
233, 267
FIGUEIRA, WILL F..................... 76, 107
FIGUEIREDO, RICARDO.................. 217
FINE, PAUL V.A. ............................ 107
FINKE, DEBORAH L. ................. 85, 107
FINKELSTEIN, SARAH A. ................ 108
FINLAY, JACQUES C....................... 179
FINLEY, JAMES .......................105, 307
FINZI, ADRIEN ........... 108, 207, 241, 371
FIRESTONE, MARY K. ..............144, 332
FISCHER, TIFFANNY ........................65
FISHER, ANN ................................ 307
FISHER, STUART G. ....................... 317
FISK, MELANY C. ..................... 74, 173
FITZHUGH, ROSS D........................ 108
FITZPATRICK, JOHN W. .................. 178
FITZSIMMONS, KEVIN ................... 197
FLANAGAN, LAWRENCE B. ..............91
FLATHER, CURTIS H. ..................... 324
FLECKER, ALEXANDER S. .............. 229
FLEMING, ALLISON .........................34
FLEMING, RICHARD A. .................. 108
FLETCHER, ROBERT J. ................... 109
FOLEY, JONATHAN A. ......................56
FOLEY, MELISSA ........................... 233
FOLLSTAD SHAH, JENNIFER J. ........ 109
FONG, PEGGY .................................15
FORBES, ANDREW E. ..................... 109
FORD, CHELCY R. ......................... 109
FORD, JESSE ................................. 109
FORD, MARY E. ............................ 109
FORKNER, REBECCA E................... 110
FORMAN, REBECCA ...................... 147
FORNWALT, BRANDON .................. 329
FORRSETER, NAOMI ...................... 158

FORSETH, IRWIN N. ....................... 163

FORSGREN, TODD ......................... 110
FORSHAY, KENNETH J.................... 110
FOSTER, BRYAN .... 16, 87, 110, 175, 232,
245
FOSTER, DAVID R.......................... 345
FOSTER, KARI............................... 121
FOSTER, SARA K. .......................... 110
FOSTER, TAMMY E. .................111, 299
FOUFOPOULOS, JOHANNES ............ 111
FOWLER, NORMA L. ........... 24, 116, 214
FOWNES, JAMES H. ....................... 186
FOWX, CYNTHIA........................... 240
FOX, CHARLES W. ......................... 323
FOX, GORDON A. ....................111, 178
FOX, GORDON .............................. 248
FOX, JENNIFER A. ......................... 111
FOX, JEREMY W. ........................... 111
FOX, JESSICA A............................. 112
FOX, MARK .................................. 124
FRANCE, KRISTIN E....................... 112
FRANCIS, TESSA B. .................112, 358
FRANK, KENNETH......................... 191
FRANKLIN, JANET......................... 112
FRANKLIN, SCOTT B................. 66, 112
FRANKS, STEVEN J........................ 113
FRATERRIGO, JENNIFER M. ............ 113
FREEMAN, BYRON J. ..................... 354
FREEMAN, CARL........................... 127
FREEMAN, D. CARL.................190, 192
FREEMAN, MARY............. 113, 235, 354
FREIDENBURG, KEALOHA.............. 113
FRELICH, LEE E. ........................... 152
FREY, JENNIFER ............................ 176
FREY, SERITA D. .....................113, 122
FRGGENS, MIKE............................ 363
FRIDLEY, JASON D....... 48, 114, 127, 342,
355
FRIEDENBERG, NICHOLAS A. ......... 114
FRIESWYK, CHRISTIN .................... 171
FRIGGENS, MICHAEL T................... 114
FRITHSEN, JEFFREY B.................... 353
FRIZZELL, KATHY ......................... 202
FROST, CECIL .........................114, 197
FROST, CHRISTOPHER J.................. 115
FROST, JODY L. ............................. 115
FRY, BRIAN .................................. 247
FRYKHOLM, JEFF .......................... 344
FRYXELL, JOHN M. .........................13
FUHLENDORF, SAMUEL D.........140, 179
FUHRMAN, NICHOLAS E. ............... 115
FUJIWARA, KAZUE ........................ 115
FUKAMI, TADASHI ..........................54
FULL, WILLIAM E.......................... 315
FULLER, PAM ............................... 324
FULTON, MARK R.......................... 115
FUNK, JENNIFER L. ....................... 115
FUREDI, MARY ANN................116, 227
FUSINA, ROBERT A........................ 292
FUSSMANN, GREGOR F. ...........116, 370
G
GABBARD, BETHANY L.................. 116
GAFFORD-BARRON, GREG ............. 218

GAGE, DANIEL J............................ 149

GAGNON, PAUL R.......................... 116
GAINES, KAREN H. ........................ 116
GAINES, STEVEN D..................233, 297
GAISER, EVELYN E........... 117, 334, 370
GALATOWITSCH, SUSAN M. ........... 278
GALEANO, GLORIA ....................... 285
GALLAGHER, JIM J. ....................... 261
GALLANDT, ERIC R........................ 139
GALLANT, NATACHA ..................... 338
GALLEGO-FERNANDEZ, JUAN
BAUTISTA .................................... 117
GALLERY, RACHEL E. .................... 117
GALLOWAY, LAURA F. .....................27
GALLUZZI, LAURIE A. ................... 117
GAMON, JOHN A. ............. 117, 244, 309
GANGER, MIKE T........................... 117
GANNON, JILL J. .............................72
GANNON, WILLIAM L. .............130, 256
GANTAR, MIROSLAV ..................... 334
GAOUE, OROU G. .......................... 274
GARCIA, TIFFANY S....................... 298
GARCIA-NOVO, FRANCISCO ........... 117
GARDESCU, SANA......................... 343
GARDINER, NED ........................... 118
GARGAS, ANDREA ........................ 197
GARMESTANI, AHJOND S. .............. 118
GARNEAU, DANIELLE E. ................ 118
GARNERY, LIONEL ........................ 325
GARNIER, ERIC ............................. 344
GARRETT, HOLLY.......................... 205
GARRETT, KAREN A. ...............118, 223
GARTNER, TRACY B. ..................... 118
GARVEY, JAMES E. ........................ 218
GAUDINSKI, JULIA B. .................... 338
GAUTHIER, JACQUES A....................26
GAVIN, DANIEL G.......................... 119
GAYANILO, FELIMON .................... 269
GEBELEIN, JENNIFER..................... 284
GEBER, MONICA A. ....................... 239
GEDDES, NEOMA A. ...................... 119
GEIGER, DONALD R....................... 165
GEIGER, ERIKA L. ......................... 119
GEIGER, JOHN H. .......................... 119
GELLERSTEDT, PAUL A. ................. 115
GELWICK, FRAN ........................... 336
GENC, LEVENT ........................ 34, 319
GENDE, SCOTT M. ......................... 313
GENKAI-KATO, MOTOMI ................ 120
GENOVESE, SALVATORE J. ............. 362
GENTRY, DALE J. .......................... 343
GEORGE, KATE ............................. 120
GEORGE, SOPHIE B........................ 120
GERACI, CHRISTY J. ...................... 120
GERACI, CHRISTY JO..................... 278
GERBER, LEAH ............................. 223
GERHARDT, FRITZ......................... 120
GERLACH, JOHN P. ........................ 121
GERMINO, MATTHEW J. ..... 47, 121, 143,
149, 169, 215
GERVIN, JANNETTE C. ................... 187
GETZ, LOWELL L........................... 258
GHASSEMZADEH, FERESHTEH........ 121

GHERSA, CLAUDIO M. .....................61

GHOLZ, HENRY L. ......................... 319
GHOLZ, HENRY...............................34
GIARDINA, CHRISTIAN P. ............... 211
GIBBONS, J. W............................... 331
GIBBONS, J. WHITFIELD................. 263
GIBSON, DAVID J...................... 60, 121
GIL-WEIR, KARINE C. .................... 352
GILADI, ITAMAR ........................... 121
GILBERT, BENJAMIN D. .................. 122
GILBERT, JANICE........................... 122
GILL, DOUGLAS E. ..........................40
GILL, RICHARD A. ......................... 122
GILLEVET, PATRICK M. .................. 123
GILLIAM, FRANK S........................ 122
GILLOOLY, JAMES F. ......... 100, 122, 297
GINZEL, MATTHEW D. .............123, 276
GIPSON, PHILIP S. ......................... 283
GITTINGER, LORI S........................ 123
GIULIANI, RITA...............................48
GIVLER, KIM A. ............................ 123
GIVNISH, THOMAS ........................ 197
GLADWELL, RANDI R. ................... 123
GLASSON, GEORGE ....................... 344
GLEASON, FLORENCE K................. 217
GLEESON, SCOTT K. ................123, 285
GLEESON, SCOTT ............................. 5
GLENN, EDWARD .......................... 197
GLIDDON, JASON R. ........................75
GLOWACKI, GARY A...................... 124
GOASSMAN, BRYAN ...................... 235
GODDARD, KELLY L. ..................... 124
GODWIN, HARRY W. ...................... 136
GOERIZ, RACHEL E. ...................... 124
GOHEEN, JACOB R. .................124, 177
GOLDBERG, DEBORAH E................ 328
GOLDSTEIN, ALLEN ...................... 331
GOLDWASSER, LLOYD................... 129
GOLLADAY, STEPHEN W................. 249
GOLUBIEWSKI, NANCY E. .............. 124
GOLUBSKI, ANTONIO J. ................. 125
GOMEZ-SAPIENS, MARTHA ............ 125
GONELLA, MICHAEL P. .................. 125
GONZALEZ, ANDREW .................... 125
GONZALEZ, MARIA J. .................... 342
LEZ-MEGIAS, ADELA.............90
GONZA
GONZALEZ-MELER, MIQUEL A. ..... 125,
311, 331, 338
GOOD, WILLIAM ........................... 251
GOODELL, KAREN......................... 125
GOODIN, DOUGLAS G. ................... 148
GOODMAN, ROBERT M. ................. 233
GOPALANI, SARAH........................ 296
GORAN, WILLIAM D. .......................20
GORCHOV, DAVID L.................. 23, 245
GORDON, DORIA........................... 342
GOSLEE, SARAH C......................6, 126
GOSZ, JAMES R. ......................114, 371
GOTELLI, NICHOLAS J. ............. 98, 126
GOTTDENKER, NICOLE L. .............. 126
GOULD, ERNEST ........................... 158
GOULD, PETER J. .......................... 105
GOVERNO, ROBIN ......................... 179

GOVINDARAJAN, SATHISH ...............66

GOVINDARAJULU, PURNIMA .......... 126
GOWE, AMY K. ............................. 126
GOWER, STITH T. ....................103, 219
GOYER, RICHARD A. ..................... 316
GRABOWSKI, JONATHAN H. ........... 127
GRACE, JAMES B...................... 33, 127
GRAHAM, JOHN H. ........... 127, 190, 192
GRAMLING, JOEL ............... 99, 114, 127
GRANDY, A. STUART ..................... 128
GRANT, ALASTAIR ........................ 303
GRANT, BRUCE W. ...................128, 332
GRANT, NIKKI M. .......................... 128
GRANTHAM, BRIAN ........................60
GRATTON, CLAUDIO........................85
GRAVES, MICHAEL........................ 344
GRAY, DENNIS W. .......................... 115
GRAY, JANET B. ............................ 128
GRAYBILL, JESSICA....................... 358
GREATHOUSE, EFFIE A. ................. 128
GREAVER, TARA ........................... 129
GREEN, DOUGLAS M. .................... 129
GREEN, EDMUND P. ....................... 284
GREEN, G. .................................... 261
GREEN, WALTON A........................ 129
GREENBERG, CATHRYN H. .......129, 194
GREENBERG, DAVID B. .................. 129
GREENBERG, RUSSELL .................. 267
GREENE, JUDITH L. ....................... 263
GREENLEE, ERIN C. ....................... 256
GREENLEE, ERIN S. ....................... 130
GREGORY, PETER J. ....................... 171
GRENE, RUTH ............................... 244
GRIECO, JOHN .............................. 279
GRIES, CORINNA.....................130, 326
GRIEVE, KATIE ...... 22, 36, 130, 218, 265,
365
GRIFFIN, CELINE A. ....................... 130
GRIFFIN, KEVIN L. 37, 42, 130, 205, 291,
305, 348, 367
GRIFFIN, KEVIN ......... 185, 248, 336, 365
GRIFFIS, PAIGE A. ......................... 131
GRIFFIS-KYLE, KERRY L. ............... 131
GRIFFITH, MICHAEL B. .................. 131
GRIFFITH, TIMOTHY M. ................. 131
GRIGOROVICH, IGOR A. ...................68
GRISE, DAVID J. ............................ 132
GRISSINO MAYER, HENRI .............. 159
GRISWOLD, MARCUS W. ................ 131
GROFFMAN, PETER M.................... 132
GROGAN, WHITNEY N. .................. 132
GROOTJANS, AB P. ........................ 132
GROSHOLZ, EDWIN D. .............132, 339
GROSS, KATHERINE L. .............133, 313
GROSS, LOU ...................................26
GROSSMAN, GARY D. .................... 346
GROVE, J. MORGAN ...................30, 54
GROVE, MORGAN ......................... 349
GROW, DAVID E. ........................... 133
GRUBBS, SCOTT............................ 231
GRUENZWEIG, JOSE M. .................. 243
GRULKE, NANCY ............................90
GRUNER, DAN S............................ 133

Author Index

381

GRUNER, SUSAN V. ....................... 310

GUENTHER, DEBBIE A. .................. 174
GUIDO, DANIEL R.......................... 251
GUILLETTE, LOUIS J. ..................... 133
GULDIN, JAMES M...........................43
GUNAWARDENE, ESHANI U. ........... 133
GUNZBURGER, MARGARET S. ........ 134
GUO, DALI ................................... 134
GUR-EDEMAN, ELVAN ................... 174
GUREVITCH, JESSICA ....... 134, 162, 248
GURNEE, JULIE E. ......................... 134
GUTZWILLER, KEVIN J.............229, 357
M, MIKAEL ................... 134
GYLLSTRO
H
HA, KYONG .................................. 167
HAAS, CAROLA A.......................... 135
HADDAD, NICK M. ..........................78
HAEFNER, JAMES W. ...............202, 254
HAINES, B. L................................. 368
HAINES, BRUCE ............................ 135
HAIRSTON, NELSON G. ............111, 370
HAJDAS, IRKA .............................. 197
HALE, BRACK W. .......................... 135
HALE, SCOTT................................ 342
HALE, V. CODY ............................. 235
HALEY, LUKE ............................... 341
HALFACRE, ANGELA C. ................. 142
HALITSCHKE, RAYKO .................... 135
HALL, BOB ................................... 254
HALL, FORREST G. ........................ 187
HALL, MYRA C. ............................ 135
HALL, SHARON J........................... 136
HALL, SONIA A. ............................ 136
HALL, SPENCER R. ........................ 136
HALL, THOMAS J. ......................... 210
HALLETT, RICHARD A.................... 155
HALPERN, BENJAMIN ......................39
HALVERSON, MARK ANDERS ......... 136
HALVORSON, JONATHAN J. ............ 136
HAMBURG, STEVEN P. ................... 367
HAMERLYNCK, ERIK P. ............137, 222
HAMILTON, E WILLIAM ...........137, 198
HAMILTON, JASON ........... 137, 187, 241
HAMILTON, ROBERT................140, 273
HAMILTON, STEVE ..........................46
HAMMAN, SARAH T....................... 137
HAMMOND, DOUGLAS E. ............... 197
HAMPTON, STEPHANIE E. .............. 138
HAMRICK, JAMES H. ..................... 280
HAMRICK, JAMES L.................. 87, 138
HAN, BARBARA A. ..........................35
HANAN, NIALL P. .......................... 138
HANCOCK, THOMAS E. .................. 138
HANDLEY, L ................................. 192
HANKS, JOSEPH H. ................... 26, 138
HANKS, LAWRENCE M. ...... 28, 123, 194,
276
HANNON, SUSAN J. ....................... 139
HANOWSKI, JOANN .................. 79, 139
HANSEN, LEE D. .............. 177, 310, 335
HANSEN, LEE ............................... 328
HANSON, PAUL J. .......................... 139

382

Author Index

HANSSON, LARS-ANDERS .............. 134

HARAMOTO, ERIN R. ..................... 139
HARCOMBE, PAUL A. .................... 115
HARDESTY, BRITTA DENISE ........... 139
HARDING, LAWRENCE W................ 258
HARE, J DANIEL............................ 312
HARE, J. DANIEL...................... 96, 140
HARLAN, NICOLE P........................ 260
HARLEY, CHRISTOPHER D.G. .......... 140
HARMON, JASON P. ....................... 140
HARPER, STEVEN J........................ 305
HARPER, STEVEN.......................... 215
HARPOLE, DOUGLAS N. ................. 135
HARPOLE, STAN............................ 104
HARPOLE, W S. ............................. 140
HARPOLE, W.S............................... 150
HARRELL, WADE C........................ 140
HARRINGTON, ROBIN A. ................ 186
HARRIS, PEGGY ............................ 147
HARRIS, REID N. ........................... 141
HARRIS, WYLIE N. ........................ 141
HARRISON, AUTUMN-LYNN .............30
HARRISON, PHILIP M. .................... 141
HARRISON, SANDY P. .................... 141
HARRISON, SUSAN ..........................81
HARRISS, ROBERT......................... 162
HART, MIRANDA M........................ 141
HART, STEPHEN C. ..................251, 304
HART, ZACHARY H. ....................... 142
HARTE, JOHN ............................... 195
HARTLE, R. TODD ......................... 142
HARTLEY, ANNE E......................... 142
HARTLEY, LAUREL M. ................... 142
HARTMAN, WYATT H..................... 142
HARTNETT, DAVID C...................... 192
HARTSHORN, ANTHONY S.............. 366
HARTVIGSEN, GREGG.................... 143
HARTWAY, CYNTHIA ..................... 143
HARTWELL, ANNA L. ......................89
HARVEY, CHAD T. ......................... 143
HASSAN, HASSAN K. ..................... 340
HASSELQUIST, NILES J. .................. 143
HASSETT, BROOKE A. .................... 249
HASTINGS, ALAN M. ..................... 332
HASTINGS, STEVE J. ........................63
HATCH, KENT A. .....................138, 143
HATTON, ELIZABETH S. ................. 143
HAWKES, CHRISTINE V. ................. 144
HAWLEY, GARY J. ...........................38
HAY, MARK E..................... 51, 180, 210
HAYES, JACK................................ 144
HAYWARD, APRIL J. ...................... 144
HE, FANGLIANG............................ 144
HE, JIN-SHENG.............................. 364
HE, XINHUA ................................. 144
HEARD, STEPHEN B....................... 145
HEATH, LINDA S. ............................64
HECKATHORN, SCOTT A. ............... 137
HECKMAN, KATHERINE A. ............. 145
HEDIN, LARS O. ......... 145, 155, 228, 344
HEDRICK, VICKI J. ........................ 145
HEIKOOP, JEFFREY M..................... 178
HEIMAN, KIMBERLY W. .... 145, 175, 235

HEIMPEL, GEORGE ........................ 157

HEIN, CATHERINE L....................... 146
HEINSCH, FAITH ANN .................... 174
HEISLER, JANA L. ......................... 146
HEITZMAN, ERIC.............................43
HELBING, CAREN.......................... 133
HELFMAN, GENE........................... 118
HELLQUIST, C. E............................ 146
HELMS, BRIAN S. .......................... 146
HELMS, J. H. ................................. 240
HELMS, SANDRA E. ....................... 147
HELMUS, MATTHEW R. .................. 147
HELMUTH, BRIAN ......................... 147
HEMBRE, LEIF K. .......................... 147
HEMMER, MICHAEL J. ................... 147
HEMMER, REBECCA L. .................. 147
HENDRICK, RONALD L................... 366
HENDRICK, RONALD ..................... 274
HENDRICKS, JOSEPH J. .................. 134
HENDRIX, PAUL F. ......................... 361
HENDRIXSON, HEATHER ................ 321
HENEBRY, GEOFFREY M. .........148, 340
HENNE, PAUL D. ........................... 148
HENRY, DAVID................................29
HENRY, HUGH A. L. ....................... 148
HENSON, SHANDELLE M. ............... 182
HEPPELL, SELINA.......................... 206
HERBERS, JOAN M. .........................18
HERENDEEN, ROBERT A. ............... 148
HERMS, CATHERINE P. ................... 316
HERMS, DANIEL A......................... 195
HERNANDEZ, DANIEL..............176, 253
NDEZ, LAURA ......................31
HERNA
HERRERA, CARLOS M......................10
HERRICK, JEFFREY E. .................... 148
HERRICK, JEFFREY........................ 238
HERRON, PATRICK M. .................... 149
HERSHEY, ANNE E...........................34
HESLER, STEVE ............................ 253
HESS, GEORGE R........................... 343
HESTER, MARK............................. 359
HETTINGER, ANNALIESE..................50
HEWITT, JENNIFER E...................... 319
HIAASEN, BARBARA A. ................. 149
HIBINO, TADASHI....................189, 330
HICKEY, LEO J. ............................. 129
HIERS, J. K. .................................. 342
HIGGINS, PAUL A. ......................... 149
HIGUCHI, NIRO ...............................60
HIGUCHI, TAKAYA ........................ 175
HILDEBRAND, T. ........................... 245
HILDREW, ALAN ........................... 149
HILL, DEVETTA............................. 285
HILL, GEOFF................................. 340
HILL, JUDSON P............................. 149
HILL, RONALD L. .......................... 150
HILL, WALTER R. .......................... 148
HILLE RIS LAMBERS, J................... 150
HILLE RIS LAMBERS, JANNEKE ...... 104
HILLHOUSE, HEIDI L...................... 150
HINCKLEY, THOMAS M. ................. 182
HINES, JESSICA E. ......................... 150
HINKLE, C. R. ............................... 206

HINKLE, C. ROSS.............................43
HINKLE, ROSS .............................. 271
HINTON, JULIANA G. ..................... 234
HITOSHI, MIYASAKA ..................... 150
HLADIK, CHRISTINE ...................... 151
HO, CHUAN-KAI............................ 151
HO, MOON-HO .............................. 241
HOBBIE, SARAH E. ...... 88, 176, 179, 230,
253, 297, 355
HOBBS, N. THOMPSON.............. 33, 151
HOCH, GREG ..................................58
HOCHACHKA, WESLEY M. ............. 155
HOCHSTRASSER, TAMARA ............. 151
HODGSON, DOMINIC A. ................. 200
HODGSON, JAMES R. ..................... 340
HOFFMAN, KATHERINE E. .............. 151
HOFMOCKEL, KIRSTEN S. .............. 151
HOFSTETTER, RICHARD W......... 18, 152
HOGSETT, WILLIAM....................... 229
HOHMAN, DOUGLAS S. .................. 152
HOLBEN, WILLIAM..........................55
HOLBROOK, NOEL M. .................... 317
HOLDO, RICARDO M...................... 152
HOLDSWORTH, ANDREW R. ........... 152
HOLLAND, J. N.............................. 250
HOLLAND, MARJORIE M. ............... 152
HOLLENHORST, TOM ..................... 251
HOLLINGER, DAVID....................... 237
HOLM, PATRICIA.............................39
HOLMES, BILL .............................. 192
HOLMES, WILLIAM E. .................... 371
HOLMQUIST, JEFF G....................... 128
HOLMQUIST, KARSTEN G. .............. 239
HOLSOMBACK, TYLA .................... 229
HOLT, CARRIE.................................16
HOLT, ROBERT D. ............. 153, 182, 187
HOLTHAM, ANITA ......................... 265
HOLZMUELLER, ERIC J. ................. 153
HONDORP, DARRYL W.................... 153
HONG, BONGGHI........................... 153
HONORIO, NILDIMAR A. ................ 173
HONU, YOHANES A.K. ................... 121
HOOD, JAMES M. .......................... 321
HOOK, JAMES E. ........................... 153
HOOK, JAMES .................................36
HOOKER, BETH A.......................... 154
HOOKER, TOBY D. ........................ 154
HOOPER, DAVID U......................... 154
HOOPER-BUI, LINDA M. ................. 305
HOPE, DIANE ..........................130, 326
HOPKINSON, CHARLES S................ 258
HOPPER, WILLIAM E................. 62, 154
HORNER-DEVINE, M. CLAIRE .... 99, 154
HORSLEY, STEPHEN B..............155, 210
HORTON, JONATHAN L. ...................67
HORVITZ, CAROL C. ...................... 155
HORWATH, WILLIAM R. ...........144, 361
HOSAKA, NAOMI .......................... 155
HOSSEINI, PARVIEZ R. ................... 155
HOST, GEORGE ............................. 251
HOTCHKISS, SARA ..................213, 344
HOU, LUCY .................................. 373
HOUCK, MARILYN A...................... 229

HOUDE, EDWARD D....................... 258

HOULTON, BENJAMIN Z. ................ 155
HOUSEMAN, GREGORY R. .............. 156
HOUSER, JEFFREY N...................... 156
HOUSER, LETISE T. ........................ 266
HOVERMAN, JASON T. .............156, 217
HOWARD, JEROME J. ................ 47, 327
HOWARD, TIMOTHY G. ............134, 328
HOWE, HENRY F. ........................... 156
HOWE, KATHERINE M.................... 157
HOWE, ROBERT W. ........................ 157
HOWE, ROBERT ............................ 139
HOWELL, EVELYN...........................24
HOWELL, JONATHAN P................... 157
HOYE, KATHERINE ........................ 157
HOYER, ERIK................................ 121
HRABIK, THOMAS R. ..................... 288
HSU, CYNTHIA L. .......................... 157
HU, FENG SHENG .. 12, 119, 148, 213, 249
HU, SHAOMIN............................... 372
HU, SHUIJIN ...................... 63, 158, 372
HUANG, QINGYU .......................... 373
HUBBARD, JAKARA....................... 158
HUBBELL, STEPHEN P. ........ 70, 158, 195
HUBERTY, ANDREA F. .................... 158
HUDSON, GREG ............................ 233
HUDSON, PETER J. ........................ 158
HUEBNER, CYNTHIA D................... 158
HUEMMRICH, KARL F. .............117, 187
HUENNEKE, LAURA F..................... 159
HUFBAUER, RUTH A. ...............214, 218
HUFFMAN, JEAN M........................ 159
HUFFORD, KRISTINA M.................. 159
HUGHES, KELLY ........................... 288
HUGHES, R. FLINT.................... 28, 159
HUGHES, TERRENCE P. ....................92
HUGHIE, HOYT H. ......................... 127
HUI, DAFENG................... 160, 327, 367
HUIJBREGTS, BAS ......................... 348
HULL-SANDERS, HELEN M. ............ 160
HULTINE, KEVIN R. ....................... 314
HUMPHREY, LEWIS D..................... 160
HUMSTON, ROBERT ...................... 160
HUNGATE, BRUCE A. ........ 160, 258, 363
HUNT, JOHN ................................. 336
HUNT, LUKE J.H. ........................... 140
HUNTER, MARK D. .... 115, 135, 147, 215,
280
HUNTZINGER, MIKAELA ................ 161
HUPP, CLIFF R. .............................. 161
HURTON, LENKA V. ....................... 161
HURTT, GEORGE C...........................92
HUTCHINSON, MARK N.................. 316
HUTCHINSON, TODD F. .................. 161
HUXEL, GARY R............................ 162
HUXMAN, TRAVIS E. .... 53, 97, 100, 137,
162, 271, 314
HUXMAN, TRAVIS ......................... 248
HWANG, BERNICE ......................... 172
HYATT, LAURA A. ...................115, 134
HYATT, LAURA ............................. 162
HYNES, MEAGAN .......................... 229

I
N
EZ, INES ............................... 162
IBA
IBARRA-NUNEZ, GUILLERMO ......... 267
IDE, CHARLES F. ........................... 167
IDIATA MAMBOUNGA, DANIEL....... 348
IDOL, TRAVIS W. ........................... 144
IELMINI, MICHAEL ........................ 309
IGNACE, DANIELLE D. .............100, 162
IGOR, KHANAYEV V. ..................... 150
IGUCHI, TAISEN ............................ 133
IHUE-UMIRE, YENNY ..................... 123
IMAI, TSUYOSHI .....................175, 350
IMHOFF, MARC L........................... 162
IMM, DONALD W. ....................163, 305
IMM, DONALD .............................. 215
INCHAUSTI, PABLO .........................56
INGRAM, ELLA L.....................163, 202
INNIS, ANNE F............................... 163
INOUYE, BRIAN D. ................... 81, 163
INOUYE, DAVID W. ........................ 163
INOUYE, DAVID ............................ 195
IRVINE, ROBYN L. ...................164, 196
ITLE, CAROLYN ............................ 204
ITO, AKIHIKO ............................... 164
IVANS, CAROLYN Y. ................265, 293
IVERSON, LOUIS R. .................. 63, 164
IVES, ANTHONY R. ........ 14, 56, 109, 111
IVES, TONY R. .............................. 358
IVEY, CHRISTOPHER T.................... 164
IYER, MEERA................................ 165
J
JABLONSKI, LEANNE M. ................ 165
JACKSON, AARON ......................... 214
JACKSON, GEORGE A. ................... 349
JACKSON, RHETT .......................... 235
JACKSON, ROB B........................... 314
JACKSON, ROBERT B. .. 99, 122, 157, 165,
225, 226, 244, 268, 360
JACKSON, STEPHEN T. ..38, 165, 307, 373
JACOBS, ALYSSA B........................ 165
JACOBS, LEE ANNE ....................... 355
JACOBSEN, ANNA L.................166, 272
JACOBY, GORDON ......................... 264
JAEGER, ROBERT G. ...................... 330
JAFFE, PETER R...............................97
JALICS RAUSCHERT, EMILY S. ........ 166
JAMES, JEREMY J. ......................... 166
JAROSZ, ANDREW M. ......... 80, 166, 220
JASTROW, JULIE D.................... 10, 166
JASTROW, JULIE............................ 311
JAWDY, SARA S............................. 252
JAYACHANDRAN, KRISH..........167, 172
JEFFERIES, ROBERT L. ................... 251
JELASO, ANNA M. ......................... 167
JENKINS, DAVID G.............. 61, 167, 224
JENKINS, MICHAEL A. .............153, 167
JENNINGS, CECIL .......................... 235
JEONG, KWANG-SEUK ................... 167
JETZ, WALTER .............................. 168
JHEE, EDWARD M.......................... 168
JHEE, EDWARD ...............................42

Author Index

383

JIANG, LIN ................................... 168

JIMENEZ-LOBATO, VANIA .............. 168
JIN, VIRGINIA L. ........................... 169
JOBBAGY, ESTEBAN G. ............. 99, 268
JOBBAGY, ESTEBAN ...................... 225
JOBE, R T...................................... 114
JOERN, ANTHONY ......................... 169
JOHANSEN, JEFFREY R. ................. 266
JOHANSEN, MATHEW P. ................. 169
JOHN, SCHADE ............................. 176
JOHNSON, BRETT M. ..................... 190
JOHNSON, D.A............................... 255
JOHNSON, DALE W. ............ 16, 169, 252
JOHNSON, DANIEL M. .................... 169
JOHNSON, DAN ............................. 121
JOHNSON, DEREK M. ..................... 170
JOHNSON, DOUGLAS E................... 129
JOHNSON, EDWARD A.......... 61, 64, 235
JOHNSON, HYRUM B. .................... 122
JOHNSON, JERALD B. .................... 170
JOHNSON, KELLY S. ................. 78, 170
JOHNSON, LORETTA C. ....... 70, 311, 315
JOHNSON, LORETTA ................. 67, 185
JOHNSON, LUCINDA B. .................. 251
JOHNSON, LUCINDA ...................... 273
JOHNSON, MARC T.J. ..................... 170
JOHNSON, MICHELE A. .................. 170
JOHNSON, PIETER TJ ..................... 171
JOHNSON, SARAH ......................... 279
JOHNSON, SCOTT N. ...................... 171
JOHNSON-BAWE, MIREILLE ..............14
JOHNSTON, CAROL A. ................... 171
JOHNSTON, JILL A. ........................ 171
JOHNSTON, TAMMY ...................... 171
JOHNSTONE, JILL ............................30
JONES, ANDY ............................... 172
JONES, CLAYTON A. ...................... 310
JONES, CLIVE G. .....................115, 298
JONES, CRYSTAL A........................ 172
JONES, CYNTHIA S. ....................... 202
JONES, DAVID T. .....................172, 294
JONES, JEANNE............................. 367
JONES, JULIA A............................. 242
JONES, LAURA.............................. 370
JONES, MALCOLM T.........................79
JONES, ROBERT H. ...... 36, 172, 249, 264,
322
JONES, RONALD D......................... 334
JONES, THOMAS G. ....................... 308
JONSEN, IAN D.............................. 173
JONSSON, COLLEEN ...................... 229
JOO, GEA-JAE ............................... 167
JORDAN, NICHOLAS R. ....................36
JOSE, SHIBU ...........................153, 167
JOST, JENNIFER............................. 147
JOYCE, LINDA A. ............................24
JUANG, JEHN-YIH.......................... 325
JUDD, KRISTIN E. .......................... 173
JUDD, KRISTI................................ 185
JUICE, STEPHANIE M. .................... 173
JULIANO, STEVEN A. .... 14, 73, 133, 173,
180, 321, 368
JUNG, DAWOON ............................ 173

384

Author Index

JURENA, P. N................................. 174

JURGENSEN-ARMSTRONG,
MELISSA
J. ................................................. 174
JURGENSEN-ARMSTRONG, MELISSA ............................................... 151
K
KACHI, NAOKI .............................. 155
KALIM, SAHIR ................................55
KALKHAN, MOHAMMED A. ............ 174
KAMMERDIENER, SUSAN A. ........... 174
KAMPF, KATY M............................ 220
KANG, SINKYU ............................. 174
KAPLAN, IAN................................ 175
KAPLAN, JESSICA A. ..................... 114
KAPPEL, CARRIE V. .................175, 235
KARBAN, RICHARD ....................... 175
KARBERG, NOAH J. ....................... 211
KAREL, IRENE S............................ 175
KARIM, REZAUL ........................... 175
KARIMPOUR, M.H.......................... 121
KARLEN, DOUGLAS L. .....................13
KARNOSKY, DAVID ....................... 192
KARPANTY, SARAH M.................... 176
KARR, JAMES R. ........................... 157
KARRON, JEFFREY D. .................... 239
KARTESZ, JOHN ............................ 324
KASHIAN, DANIEL M. .................... 300
KATHOLI, CHARLES ...................... 340
KATS, LEE B. ..................................35
KATUL, GABRIEL G. ...................... 325
KAUFMANN, MERRILL R. ............... 136
KAVANAUGH, MARIA ....................... 9
KAY, ADAM ............................176, 253
KAYE, THOMAS N. ........................ 176
KEARSLEY, MICHAEL J.C. .............. 176
KEDDY, PAUL ............................... 288
KEELEY, JON E................. 227, 341, 362
KEELIN, CHERYL .......................... 176
KEELING, MATT J. ......................... 177
KEESING, FELICIA...................177, 257
KEFAUVER, SHAWN C.................... 324
KEFERL, EUGENE P. ....................... 328
KEIFER, MARYBETH...................... 341
KEIFFER, CAROLYN H............... 17, 223
KEIM, PAUL S. .............................. 360
KEITH, DAVID A............................ 278
KELLER, BILL .................................16
KELLER, EMILY A. ..................177, 310
KELLER, TROY A. ......................... 177
KELLEY, ALEXIA M. ...................... 177
KELLOGG, CHEV H. ....................... 178
KELLY, CHARLENE N. .................... 178
KELLY, DAVE................................ 298
KELLY, STEPHANIE J...................... 178
KELT, DOUGLAS A. ........................ 256
KEMP, W. MICHAEL ....................... 267
KEMPES, CHRISTOPHER P............... 178
KENDALL, BRUCE E. ...............111, 178
KENDALL, WILLIAM L. .................. 286
KENNAMER, ROBERT A. ................ 179
KENNARD, DEBORAH K. ................ 179
KENNEDY, THEODORE A. ............... 179

KENNEY, AMANDA M. ................... 132

KEOUGH, CINDY .............................84
KERBY, JACOB.............................. 179
KERBY, JAY .................................. 179
KERKHOFF, ANDREW J......... 21, 96, 180
KERN, KRISTINE A. ....................... 166
KERN, RUTH ANN .................... 44, 180
KERSHNER, MARK W. .................... 287
KESAVARAJU, BANUGOPAN ...... 73, 180
KESSLER, ANDRE .......................... 180
KESTER, KAREN M. ....................... 203
KICKLIGHTER, CYNTHIA E. ............ 180
KICKLIGHTER, DAVID W. ............... 372
KICKLIGHTER, DAVID....... 209, 335, 371
KIEHL, KATHRYN.......................... 167
KIESECKER, JOSEPH M. ...................35
KIKUZAWA, KIHACHIRO ................ 184
KIKVIDZE, ZAAL........................... 211
KILHAM, SUSAN S......................... 181
KILLILEA, MARY E. ....................... 181
KILLINGBECK, KEITH T.................. 181
KILPATRICK, A. MARM .................. 181
KILPATRICK, MARMADUKE............ 111
KIM, CHANG KYUN ....................... 182
KIM, CHONG-KYU ......................... 189
KIM, DAN..................................... 324
KIM, DONGGIL.............................. 181
KIM, HYUN-SEOK.......................... 182
KIM, IN SUNG .........................182, 201
KIM, JOON..............................174, 181
KIM, KE CHUNG..............................53
KIM, MYOUNG-CHUL..................... 167
KIM, RAE-HYUN............................ 182
KIMBALL, JOHN S. ........................ 174
KIMBRELL, TRISTAN S................... 182
KIMMEL, DAVID ........................... 258
KIMMERER, ROBIN W. ................... 117
KINDSCHER, KELLY ........................16
KING, AARON A. ........................... 182
KING, ELIZABETH G. ..................... 182
KING, JENNIFER Y. ........................ 266
KING, JOHN S. .............................. 211
KING, JOHN .................................. 192
KING, RACHEL T. .......................... 183
KING, RYAN S............................... 281
KING, SAMMY L............................ 161
KINKEAD, KAREN E....................... 183
KINTER, CECILIA LYNN ................. 183
KINZIG, ANN P. .......................183, 349
KIRCH, PATRICK ........................... 344
KIRKMAN, KATHERINE ............. 68, 183
KIRKMAN, L. KATHERINE ................. 8
KIRWAN, MATTHEW L. .................. 184
KITAJIMA, KAORU ..................184, 270
KITCHENS, WILEY M. ......... 94, 210, 356
KITZBERGER, THOMAS.................. 222
KLAPER, REBECCA........................ 184
KLAUSMEIER, CHRISTOPHER A. ..... 184
KLAUSMEIER, CHRISTOPHER ......... 208
KLEIN, ADAM ............................... 112
KLEMOW, KENNETH M. ................. 184
KLEPEIS, DEBRA M. ...................... 268
KLEPPEL, G. S. ................................89

KLEPPEL, GARY S. ........................ 172

KLEPZIG, KIER D.............................18
KLEPZIG, KIER.............................. 152
KLIMAS, CHRISTIE ........................ 185
KLINE, KERRY A. .......................... 185
KLING, GEORGE W. .......... 173, 185, 315
KLIPS, ROBERT A. ......................... 165
KLIRONOMOS, JOHN N................... 185
KLOEPPEL, B. D. ........................... 368
KLOEPPEL, BRIAN D. ...............185, 321
KLOPATEK, JEFFREY M. ............ 22, 188
KLUEPFEL, DANIEL A. ................... 331
KLUG, MIKE J. .............................. 261
KLUGER, EMILY C...................186, 252
KLUGH, KATRINA R.................186, 355
KNAPP, ALAN K. ................. 58, 70, 252
KNAPP, ALAN .................................67
KNAPP, LIZA B. ............................. 186
KNAPP, SHANNON M...................... 135
KNEITEL, JAMIE............................ 186
KNEPP, RACHEL G. ........................ 187
KNIGHT, TIFFANY M. ................ 61, 187
KNOEBL, IRIS ............................... 147
KNOEPP, JENNIFER D. ......................98
KNOPS, JOHANNES M H ................. 187
KNOPS, JOHANNES M.H. ........... 43, 150
KNOPS, JOHANNES MH .................. 187
KNORR, MELISSA A....................... 113
KNOWLTON, FREDERICK F. ..............22
KNOX, ROBERT G.......................... 187
KOBE, RICHARD K. .................165, 224
KOBE, RICHARD .....................193, 214
KOCHSIEK, AMY E. ....................... 187
KOECHY, MARTIN ......................... 188
KOEL, TODD M. ..............................64
KOELLE, KATIA V. ......................... 188
KOELLIKER, JAMES K. .....................70
KOELLIKER, JAMES.........................67
KOEPFLER, ERIC ........................... 329
KOERNER, BRENDA A.................... 188
KOERNER, CHRISTIAN ................... 243
KOFORD, ROLF R. ......................... 109
KOIKE, FUMITO ............................ 188
KOLAR, CINDY S........................... 188
KOLASA, JUREK...........33, 144, 189, 286
KOLB, THOMAS E............................. 5
KOMAI, KATSUAKI ........................ 189
KONG, WOO-SEOK ........................ 189
KOO, KYUNG-AH .......................... 189
KOPTUR, SUZANNE ....................... 189
KOSKI, MARCI L. .......................... 190
KOSLOW, JENNIFER M. .................. 190
KOTA, NATHAN L. ......................... 190
KOVACIC, DAVID A. ................190, 192
KOY, KEVIN....................................40
KRAL, ANDREA M. ........................ 113
KRAMER, KAREN A. ...................... 190
KRANKINA, OLGA N. ..................... 254
KRASNY, MARIANNE E. ............ 37, 306
KRATZ, TIMOTHY K....................... 328
KRAUS, JOHANNA M. .................... 191
KRAUSE, ANN............................... 191
KREEGER, DANIELLE A....................52

KREMEN, CLAIRE....................252, 313

KRENZ, CHRISTOPHER................... 191
KREPS, TIMOTHY A. ...................... 191
KRESS, W. JOHN............................ 195
KROES, DANIEL E. ........................ 368
KRUEGER, DAMON........................ 288
KRUGER, ERIC L. ............... 91, 101, 165
KRZYSIK, ANTHONY J. ..... 127, 190, 192
KUBISKE, MARK E. ....................... 192
KUDEYAROV, V N. ........................ 192
KULA, ABIGAIL R.......................... 192
KULAKOWSKI, DOMINIK................ 192
KULKARNI, MADHURA V. .............. 193
KULMATISKI, ANDREW.................. 193
KUMMEL, MIROSLAV .................... 193
KUNKLE, JUSTIN M. ...................... 193
KUO, HUANG-CHI............................85
KUPPINGER, DANE ..................342, 355
KURSAR, THOMAS A. ............... 69, 193
KWIT, CHARLES ............................ 194
KYLE, SEAN C. ............................. 194
KYSER, KURT T. ............................ 253
L
LABRAM, JILL A. .......................... 194
LACERTE, VALERIE ....................... 197
LACEY, EMERSON S....................... 194
LADD, DAVID G. ........................... 194
LADEAU, SHANNON L..............162, 194
LADEAU, SHANNON ...................... 364
LADOFOGED, THEGN..................... 344
LAFFERTY, KEVIN D. ..................... 223
LAFOREST, JOSEPH H..................... 195
LAHM, SALLY A............................ 348
LAHTI, MEGAN ............................. 195
LAIRD, DAVID A............................ 292
LAKE, FRANK K. ........................... 195
LAKE, JEFFREY K.......................... 195
LAMAR, WILLIAM R. ..................... 195
LAMBERT-JACK, TRACY ................ 277
LAMBRECHT, SUSAN ..................... 195
LAMBRINOS, JOHN G. .................... 339
LAMBRINOS, JOHN........................ 326
LAMONTAGNE, JALENE M.............. 196
LAMP, WILLIAM O. ..........................19
LAMPHERE, BRADLEY A. ............... 196
LANCASTER, JILL.......................... 196
LANDENBERGER, RICK E. ........190, 196
LANDIS, FRANK ............................ 197
LANDIS, MATT.............................. 248
LANE, DIANA ............................... 156
LANGDON, CHRIS ......................... 197
LANGLEY, SUSAN K....................... 197
LANZA, JANET.............................. 197
LAPOINTE, DENNIS .......................... 6
LARA, RUBEN............................... 302
LARKIN, PATRICK ......................... 147
LAROCQUE, GUY R. ...................... 197
LARRIMER, AUDREY K. ................. 198
LARSON, KATHERINE C. ................ 198
LASSEN, KARI E............................ 198
LATTY, ERIKA F.......................198, 259
LAU, JENNIFER ............................. 198

LAUENROTH, WILLIAM K........ 6, 43, 51

LAUGHLIN, KAREN D. ................... 199
LAURENCE, JOHN ......................... 229
LAURENCO-DE-OLIVEIRA, RICARDO .............................................. 173
LAURENT, EDWARD J...............199, 201
LAWLER, SHARON P....................... 199
LAWRENCE, DEBORAH ... 70, 88, 95, 158,
365
LAWRENCE, ERIN M. ..................... 308
LAWRENCE, WILLIAM T. ................ 162
LAWSON, DAWN M. ....................... 199
LAWSON, MICHAEL C. ................... 272
LAYBOURN-PARRY, JOHANNA ........ 229
LAYMAN, CRAIG A. ....................... 199
LAYNE, DESMOND ........................ 354
LAZORCHAK, JAMES M.................. 131
LEACH, ANDREA D........................ 200
LEAKEY, ANDREW ........................ 200
LEAR, JENNIFER A. ....................... 200
LEASE, HILARY M. ........................ 244
LEAVITT, PETER R. ........................ 200
LEBOUTON, JOSEPH P. .............199, 201
LECHLER, PAUL ............................ 216
LECHOWICZ, MARTIN J.................. 122
LEDFORD, JOANNE........................ 252
LEE, DOWON ................................ 181
LEE, EUN JU ...........................182, 201
LEE, G-J ....................................... 255
LEE, GILJAE ................................. 256
LEE, R. W................................146, 259
LEE, RICHARD F. ........................... 366
LEE, SARAH C............................... 201
LEE, WILLIAM G. ............................75
LEFEVRE, KARA L......................... 201
LEFF, LAURA G. ....................... 19, 255
LEFFLER, A JOSHUA...................... 293
LEFFLER, A. JOSHUA ..................... 201
LEFFLER, JOSH ............................. 265
LEHMAN, ELIZABETH.................... 202
LEIBOLD, MATHEW A. .............202, 321
LEICHT, STACEY A. ....................... 202
LEININGER, TED ........................... 324
LEISHMAN, MICHELLE .................. 202
LEMASSON, BERTRAND H. ............. 202
LEMOINE, NICOLE...........................53
LENHARD, GERALD J..................... 316
LENNON, JAY T. ............................ 203
LENSING, JANET R. ....................... 203
LENTILE, LEIGH B. ..................203, 311
LENTZ, AMANDA J. ....................... 203
LEONARD, NORMAN E. .................. 203
LEOPOLD, DONALD J. ......................28
LERDAU, MANUEL T. ..................... 115
LERDAU, MANUEL ..........................16
LESLIE, HEATHER ......................... 203
LETOURNEAU, DEBORAH K............ 204
LEUNG, BRIAN.............................. 204
LEVEY, DOUGLAS J. ........... 38, 129, 194
LEVIN, LISA ................................. 132
LEVINE, JONATHAN M. .................. 204
LEVINE, MIA T. ............................. 204
LEVINSON, BARBARA.................... 225

Author Index

385

LEVRI, EDWARD P. ........................ 204

LEWIS, DANNY ............................. 205
LEWIS, DAVID B............................ 205
LEWIS, GREGORY ......................... 205
LEWIS, J D................. 205, 219, 291, 339
LEWIS, JAMES D. ....................220, 305
LEWIS, KRISTIN C. ........................ 206
LEWIS, MARK A. ................ 67, 212, 364
LEWISON, REBECCA...................... 206
LEYSHON, BRADLEY D. ................. 361
LI, BAI-LIAN ............................ 63, 206
LI, HARBIN.............................206, 315
LI, HUA..........................................76
LI, J. -H. ....................................... 206
LI, PEIGANG ................................. 334
LICHTER, JOHN............22, 108, 110, 207
LICITRA, JEFF ............................... 339
LIEBHERR, JAME K........................ 215
LIEBHOLD, ANDREW M.................. 207
LIGHTFOOT, DAVID C.......................81
LIGHTFOOT, DAVID ....................... 176
LILJA, PAUL R............................... 210
LILL, JOHN T...........................110, 207
LIM, JONG-HWAN.......................... 182
LIMMER, JACOB..............................55
LIN, GUANGHUI ......................207, 367
LIN, YICHING ............................... 207
LINCOLN, DAVID E. ....................... 208
LIND, JIM .......................................79
LINDAHL, AMY E. ...........................97
LINDER, ERIC T. ....................... 71, 208
LINDER, ERIC ............................... 324
LINDROTH, RICHARD L....... 91, 208, 322
LINKE, R. B. ................................. 206
LIPS, KAREN R.............................. 181
LIPS, KAREN................................. 246
LIPSON, DAVID ........................ 78, 361
LISA, BELDEN K..............................35
LISCHKA, STACY .......................... 288
LITCHMAN, ELENA .................184, 208
LIU, HONG .............................189, 208
LIU, JIANGUO .................. 199, 201, 209
LIU, JIYUAN ...........................259, 335
LIU, MINGLIANG.............. 209, 259, 335
LIU, XIAO-CHUAN ...........................78
LLOYD, ANDREA H........................ 209
LOARIE, SCOTT R.......................... 291
LOARIE, SCOTT............................. 304
LOCKABY, B. GRAEME.............209, 301
LOCKABY, GRAEME ........... 59, 179, 299
LODGE, DAVID M. ............ 191, 204, 287
LOEW, SABINE S..............................12
LOEWENSTEIN, NANCY J. .............. 209
LOFTIN, CYNTHIA S....................... 210
LOFTUS, WILLIAM F. ................ 94, 337
LOGIUDICE, KATHLEEN ................. 257
LOHSE, DAVE ............................... 275
LOIK, MICHAEL ............................ 195
LONG, JEREMY D. ......................... 210
LONG, NOAH ..................................22
LONG, ROBERT P. ............. 155, 210, 277
LONG, STEPHANIE ........................ 237
LONG, STEPHEN P............................86

386

Author Index

LONG, STEVE ............................... 200

LONGSTRETH, DAVID J. ...................77
LONSDORF, ERIC V. ....................... 210
LOPEZ, RYAN P. ............................ 180
LORD, CYNTHIA C......................... 210
LOREAU, MICHEL ......................... 104
LORENZ, JEROME J........................ 337
LORTIE, CHRISTOPHER J. ............... 211
LOTT, MICHAEL A. ........................ 211
LOTT, MICHAEL J. ......................... 345
LOUCKS, COLBY ........................... 162
LOUDA, SVATA M.......................... 370
LOUGHEED, VANESSA L. ............... 211
LOUNIBOS, L P. .......................131, 368
LOUNIBOS, L. PHILIP ..................... 173
LOVE, ALAN C. ............................. 202
LOVELOCK, CATHERINE E......... 62, 102
LOVETT, GARY M. ......................... 108
LOVEYS, BETH R........................... 211
VGREN, JOHAN ......................... 213
LO
LOWE, KATHRYN E. ...................... 109
LOYA, WENDY M. ......................... 211
LUBCHENCO, JANE .........9, 60, 212, 233
LUCASH, MELISSA S...................... 212
LUCK, GARY W. ............................ 209
LUDWIG, FULCO ........................... 287
LUDWIG, PATRICE M. .................... 141
LUGO, ARIEL ................................ 291
LUNCH, CLAIRE K. ........................ 212
LUNDE, KEVIN B. .......................... 171
LUNNEN, SHANE........................... 204
LUO, HONGYANG.......................... 309
LUO, YIQI...... 16, 160, 169, 212, 306, 327,
347, 367
LUTGEN, EMILY R. ........................ 282
LUTSCHER, FRITHJOF .................... 212
LUTZ, KIM.................................... 235
LUX, HEIDI B. .................................77
LYNCH, ANN M. ............................ 213
LYNCH, ELIZABETH A.................... 213
LYNCH, JASON A........................... 213
LYNCH, JONATHAN P. .................... 318
LYON, BRUCE E. .............................59
LYONS, KELLY G........................... 213
LYONS, WILLIAM B. ........................21
M
MAAS, MARTHA M. ....................... 214
MACEK, PETR ............................... 279
MACFARLANE, DAVID W. ............... 214
MACISAAC, HUGH J.........................68
MACK, RICHARD N..... 110, 183, 214, 237
MACKES, KURT H............................24
MACKINNON, DANIEL ................... 214
MACMAHON, JAMES A. ................. 250
MADDEN, KATHRYN................215, 305
MADRITCH, MICHAEL D................. 215
MADRITSCH, A. ............................ 206
MADSON, STEPHANIE L. ................ 215
MAERZ, JOHN C. ........................... 215
MAGEE, TERESA ........................... 282
MAGILL, ALISON .......................... 237
MAGNSUON, JOHN J. ..................... 288

MAGNUSON, JOHN J. ..................... 146

MAHALL, BRUCE E........................ 268
MAHER, ELIZA L. .......................... 215
MAHERALI, HAFIZ ........................ 360
MAIERSPERGER, THOMAS K........... 254
MAKIN, JESSICA L......................... 143
MALLARINO, ANTONIO P................ 292
MALLISON, SHANNON M................ 216
MALMSTROM, CAROLYN M....... 52, 216
MALONEY, KELLY O. ........ 216, 236, 239
MALONEY, KELLY......................... 156
MALONEY, PATRICIA E. ...................81
MANISH, CHINTAMANI S................ 216
MANOS, PAUL S. .....................230, 360
MARC, ELLA AKOU ....................... 348
MARCH, JOHN .............................. 347
MARCHINKO, KERRY B....................10
MARCO, ADOLFO ............................35
MARIANO, NESTOR A. ................... 216
MARITAN, AMOS........................... 158
MARKEWITZ, DANIEL.................... 217
MARKO, ADAM C. ...................156, 217
MARKO, MICHELLE D. ................... 217
MARKOW, T A.................................58
MARKS, DAVID ............................. 139
MARKS, PETER L........................... 343
MARQUARDT, PAULA E.................. 217
MARQUARDT, PAULA .................... 192
MARQUET, PABLO A. ..................... 357
MARQUIS, ROBERT J................110, 207
MARR, DEBORAH L. ................217, 256
MARRA, PETER P. .......................... 253
MARRS, ROBIN A. ......................... 218
MARSCHALL, ELIZABETH A. .......... 218
MARSE, TODD J. ........................... 218
MARSH-MATTHEWS, EDIE ....47, 83, 218,
221
MARSHALL, JOHN D. ...... 22, 60, 95, 130,
218, 338
MARTENS, SCOTT N. ..................... 124
MARTIN, ANDREW ..................205, 219
MARTIN, ANDY............................. 291
MARTIN, CHRIS A....................130, 205
MARTIN, CHRIS ............................ 349
MARTIN, JENNIFER L. .................... 219
MARTIN, TIMOTHY A..................... 319
MARTINELLI, LUIZ A. ......................91
MARTINEZ, JEANNETTE C. ............. 219
MARTINEZ, M. LUISA .................... 219
MARTINEZ, MARIA........................ 177
MARTINEZ-YRIZAR, ANGELINA ...... 125
MARTINS, HELEN R. ........................34
MARTINS, LILIANE T........................60
MARTINSEN, GREG D..................... 360
MARTINSON, SHARON J. ..................18
MARTYNIUK, CHRIS ...................... 338
MARUCA, SUSAN ....................219, 232
MARUSSICH, WENDY A. ................ 220
MARX, LAURA.............................. 220
MASIELLO, CAROLINE A. ............... 337
MASON, DORAN............................ 191
MASON, ERIN E............................. 220
MASUOKA, PENNY ........................ 279

MATAMALA, ROSER ........... 10, 311, 331

MATES-MUCHIN, JONATHAN .......... 220
MATHIAS, ANDREA ....................... 220
MATHIASEN, ROBERT L. ................ 229
MATHIS, MARILYN J. ..................... 121
MATSON, PAMELA A................. 57, 344
MATSON, PAMELA ........................ 344
MATSUMOTO, HIDEO..................... 330
MATTHEWS, WILLIAM J. ................ 221
MATTICE, JENNIFER A. .................. 221
MATTSON, WILLIAM........................51
MATZNER, STEVEN L.......................86
MAUN, ........................................ 221
MAY, DAVE Z. ............................... 221
MAYER, AUDREY L. ................221, 263
MAYFIELD, MARGARET M.............. 222
MAZER, SUSAN............................. 159
MAZIA, NOEMI C...................... 61, 222
MAZZARELLI, LISA ....................... 185
MAZZARI, DARREL ....................... 334
MAZZOTTI, FRANK J...................... 263
MCAULIFFE, JOSEPH R................... 222
MCCAIN, CHRISTY M..................... 222
MCCALL, ANDREW C..................... 222
MCCALLUM, HAMISH I. ................. 223
MCCAMENT, CORINNE L. ............... 223
MCCARLIE, V. W............................ 335
MCCARRON, JAMES K. .................. 223
MCCARTHY, BRIAN C. .. 8, 198, 223, 226,
259, 280, 282
MCCARTHY, HEATHER ................... 223
MCCARTHY, MEGAN C................... 224
MCCARTHY, SARAH E.................... 224
MCCARTHY, THOMAS M. ............... 285
MCCARTY, GREGORY W. ................ 282
MCCARTY, JOHN P. ........................ 194
MCCAULEY, LISA .......................... 224
MCCLAIN, MICHAEL E. .................. 297
MCCLOSKEY, JON ......................... 210
MCCOWN, JAMES W. ..................... 244
MCCOWN, WALTER .........................89
MCCOY, MICHAEL W...................... 224
MCCULLEY, REBECCA L................. 225
MCCULLOH, KATHERINE A. ........... 225
MCDANIEL, JOSH M....................... 225
MCDILL, MARC E. ......................... 105
MCDONALD, EVAN........................ 192
MCDONALD, MICHAEL E................ 225
MCDONALD, ROBERT I. ................. 225
MCDONNELL, MARK J. .................. 359
MCDOWELL, WILLIAM H................ 128
MCELRONE, ANDREW J. ... 157, 226, 244
MCEUEN, AMY B.....................226, 320
MCEWAN, RYAN W. ....................... 226
MCFARLIN, CAROLINE R. ............... 226
MCGHEE, JAY D. ........................... 227
MCGINNIS, NATHAN ...................... 276
MCGINNIS, THOMAS W. ...........227, 362
MCGONIGLE, TERENCE P................ 143
MCGRAW, JAMES B.... 116, 190, 195, 196,
227, 295
MCGRAW, JODI M. ......................... 227
MCGREGOR, RACHELLE L. ............. 228

MCGRODDY, MEGAN E. ................. 228

MCGUIRE, DAVID A. ...................... 372
MCGUIRE, MARY ANNE .... 109, 228, 333
MCINNIS, MICHAEL L. .....................95
MCINTOSH, MOLLIE D. .................. 211
MCINTYRE, AIMEE P. ..................... 228
MCINTYRE, BLODWYN M............... 228
MCINTYRE, NANCY E. ................... 229
MCINTYRE, PETER B. .................... 229
MCIVER, JAMES ............................ 347
MCKANE, ROBERT ........................ 229
MCKEE, KAREN ............................ 359
MCKELVEY, STEPHEN R. ................ 229
MCKENNA, KERRY C. .................... 229
MCKERNAN, PAMELA.................... 229
MCKNIGHT, MEGHAN ..............114, 357
MCLACHLAN, JASON S. ................. 230
MCLACHLAN, JASON.......................36
MCLANE, TRACY ............................95
MCLAUCHLAN, KENDRA K............. 230
MCLENNAN, DONALD ................... 270
MCLEOD, KEN W. .......................... 163
MCMANUS, JOHN .......................... 269
MCMILLAN, BRETT A. ................... 230
MCMILLIAN, MIKE ..........................21
MCMURTREY, JAMES E. ...... 73, 100, 236
MCMURTREY, JIM E.........................52
MCNEELY, CAMILLE...................... 230
MCNULTY, STEVEN G. ................... 260
MCNULTY, STEVE.......................... 274
MCPHEARSON, P. TIMON................ 231
MCPHERSON, GUY R. .................... 119
MCQUATTIE, CAROLYN J. .............. 231
MCSHEA, WILLIAM J. ......................40
MEADOR, ANDREW J. .................... 231
MEENTEMEYER, ROSS K. ............... 241
MEGONIGAL, J P............................ 296
MEGONIGAL, JAMES P. .................. 150
MEGONIGAL, PATRICK .....123, 231, 250,
363
MEIER, ALBERT J. ...................231, 319
MEIER, OUIDA W. .......................... 231
MEINERS, SCOTT J. .................232, 370
MEINI, MONICA ............................ 317
MEIR, ELI ...............................219, 232
MEISTER, M. H. ............................. 206
MELACK, JOHN M. ........................ 236
MELBOURNE, BRETT A. ...................63
MELBOURNE, BRETT ..................... 232
MELILLO, JERRY M. ................335, 372
MELILLO, JERRY........ 209, 232, 259, 371
MELLARD, JARAD P. ...................... 232
MENA-ALI, JORGE I. ...................... 321
MENDELSSOHN, IRVING A. ............ 232
MENDELSSOHN, IRV...................... 359
MENGE, BRUCE A. ................ 9, 60, 233
MENGE, DUNCAN L. ...................... 233
MENGES, ERIC S. .......................... 274
MENGES, ERIC .............................. 208
MENNINGER, HOLLY L................... 233
MENTZER, JESSICA L..................... 233
MERCADO-SILVA, NORMAN ........... 288
MERCHANT, AMETHYST G. ............ 234

MERCHANT, JAMES W. ................... 340

MERRITT, RICHARD W............... 27, 211
MESONES A., YTALO ..................... 107
METHOT, GINETTE ........................ 202
METTLER, PAIGE........................... 312
METTS, BRIAN S. .......................... 263
METZGER, KRISTINE L................... 234
MEYER, CLIF W............................. 278
MEYER, CLIFTON W.................124, 234
MEYER, EUGENE........................... 234
MEYER, HARRY A. ........................ 234
MEYER, J. L. ................................. 368
MEYER, JUDY L. ........................... 235
MEYER, JUDY ............................... 118
MEYERSON, LAURA A. .................. 235
MIAO, SHILI ................................. 235
MICHAELSEN, JOEL....................... 333
MICHALET, RICHARD .................... 211
MICHALETZ, SEAN T...................... 235
MICHELI, FIORENZA......... 145, 175, 235
MICHELS, GERALD J...................... 237
MICKLER, ROBERT A. ......................64
MIDDENDORF, GEORGE ...................30
MIDDLEMAN, ROSALI......................55
MIDDLETON, BETH A..................... 121
MIDDLETON, ELIZABETH M. ...... 52, 73,
100, 236
MIHAIL, JEANNE D. .........................48
MIHUC, JANET A. ............................47
MIKAMI, KOICHI ........................... 188
MILCHUNAS, DANIEL G. ................... 6
MILLAR, BONNIE .......................... 223
MILLAR, JOCELYN G. ..............123, 194
MILLER, AMY E. ........................... 236
MILLER, DEBORAH L. .................... 367
MILLER, JERRY............................. 216
MILLER, MICHAEL W. .................... 151
MILLER, MIKE ................................10
MILLER, RICHARD M. .................... 311
MILLER, STEPHANIE A.............236, 239
MILLER, THOMAS E....................... 236
MILLS, APRIL V. ............................ 237
MILLS, APRIL ............................... 358
MINCHIN, PETER R. ....................... 237
MINER, BENJAMIN G. .................... 237
MINER, BENJAMIN ........................ 224
MINGUS, PATRICK...........................90
MINK, KIM .....................................65
MINKKINEN, KARI ........................ 206
MINOCHA, RAKESH....................... 237
MINTON, MARK S.......................... 237
MIRIK, MUSTAFA .......................... 237
MIRITI, MARIA N........................... 238
MISURELLI, DARCY....................... 177
MITAL, STEPHEN........................... 238
MITCHELL, CHARLES E............238, 271
MITCHELL, KATHERINE A. ............. 238
MITCHELL, RANDALL J.................. 239
MITCHELL, RICHARD M. ..........216, 239
MITCHELL, RICHARD .................... 236
MITCHELL, ROBERT J. ......109, 134, 264,
342
MITCHELL, ROBERT ...................... 183

Author Index

387

MLADENOFF, DAVID J. ................... 198

MO, WENHONG............................. 256
MOELLER, DAVID A....................... 239
MOFFETT, J. MINCY....................... 239
MOHAN, JACQUELINE E. ................ 239
MOHAN, SANDHYA ....................... 240
MOKOS, JENNIFER T. ..................... 240
MOLL, JASON D. ........................... 240
MOLLOY, THOMAS ........................ 139
MOMEN, B.................................... 240
MONACO, THOMAS A. ................... 241
MONGER, CURTIS ......................... 238
MONSON, RUSSELL K. ................... 303
MONTZ, GLEN .............................. 288
MOODY, AARON ........................... 241
MOODY-WEIS, JENNIFER M. ........... 241
MOON, DANIEL C. ...................135, 288
MOON, DANIEL............................. 323
MOONEY, HAROLD A. ..............241, 267
MOONEY, KAILEN A. ..................... 241
MOONEY, SARAH E........................ 330
MOORCROFT, PAUL R. ...................... 7
MOORCROFT, PAUL .........................92
MOORE, DAVID J. .......................... 241
MOORE, DOUGLAS I. ..................... 356
MOORE, GEORGIANNE W. .............. 242
MOORE, JEFFREY E. ...................... 242
MOORE, JENNIFER ........................ 274
MOORE, JOHN C............................ 242
MOORE, JONATHAN W. .................. 242
MOORE, MARGARET M. ................. 231
MOORHEAD, DARYL L. ..... 229, 242, 351
MOORHEAD, KEVIN ...................... 350
MOPPER, SUSAN .....................119, 299
MORAES, JORGE ........................... 217
MORALES, MANUEL A. .................. 243
MORAN, MATTHEW D. .. 35, 42, 243, 319,
323
MORAN, ROBBIN........................... 350
MORENO-CASASOLA, PATRICIA ...... 243
MORGAN, JACK A. ................... 51, 243
MORGAN, JACK ............................ 262
MORGAN, JOHN W. ........................ 359
MORIN, PETER J. .............. 168, 231, 273
MORRIS, J T. ................................. 329
MORRIS, JAMES T............... 79, 243, 258
MORRIS, SHERRI J. ..........................45
MORRIS, THOMAS F. ...................... 154
MORRIS, TIFFANY ......................... 207
MORTENSEN, DAVID A................... 160
MORTON, JOSEPH B....................... 178
MOSER, JOHN ............................... 152
MOSES, MELANIE E. ...................... 244
MOSIER, ARVIN R. .. 51, 84, 244, 262, 266
MOTOMI, GENKAI-KATO ................ 150
MOTT, KEITH A. ..............................50
MOTZKIN, GLENN ......................... 345
MOU, PAUL P. ............... 34, 36, 264, 348
MOURA, CATARINA F..................... 244
MOUSSY, FRANCIS ..........................57
MOYER, MELISSA A. ..................... 244
MOYES, ANDREW B....................... 244
MU, SHOU GUO ............................ 181

388

Author Index

MULDER, C. P. H............................ 245

MULDER, CHRISTA P.H...... 283, 289, 362
MULDER, CHRISTA........................ 286
MULHOLLAND, PATRICK J....... 156, 216,
239
MULKEY, STEPHEN S. ....... 184, 234, 270
MULLAN, CAITLIN P. ..................... 245
MULLER, ROBERT N. ..................... 226
MULLER-NAVARRA, DOERTHE .........20
MULLIGAN, MARGARET R.............. 245
MUMME, RONALD L. ..................... 194
MUNGUIA, PABLO ......................... 245
OZ, ALEJANDRO A.................. 247
MUN
MURAKAMI, KAZUO...................... 330
MURAKAMI, MASASHI.....................25
MURDOCH, WILLIAM .................... 245
MURIE, JAN O. ................................90
MURPHY, C. A. .............................. 245
MURPHY, JOHN F. .......................... 101
MURPHY, KATHRYN A. .................. 246
MURPHY, PETER J. ........................ 246
MURPHY, SHANNON K. .................. 246
MURPHY-MEDLEY, DEENA ............. 287
MURRAY, MELISSA........................ 288
MURRAY, TANYA .......................... 246
MURRELL, KATHREN E. ................. 247
MURTHY, RAMESH ...22, 36, 78, 130, 185,
200, 218, 265, 338, 348, 363, 365
MURTHY, R .................................. 192
MUSS, JORDAN D. ...................247, 345
MUTCHLER, TROY ........................ 247
MUTH, CHRISTINE C...................... 247
MYER POLACEK, KELLY ................ 202
MYERS, JONATHAN A. ................... 343
MYERS, ORRIN B........................... 234
MYERS, WAYNE L. ........................ 349
N
N.A., N.A. ..................................... 152
NADELHOFFER, KNUTE J. .............. 315
NADELHOFFER, KNUTE ................. 185
NAEEM, SHAHID .....................187, 365
NAGEL, JENNIFER ......................... 248
NAGELKERKE, KEES ..................... 248
NAKATANI, NAOKI ........................ 248
NANNINI, MICHAEL A.................... 248
NASHAGNIK, JOSHUA.................... 109
NASON, JOHN D. ........................... 145
NASSAR, JAFET...............................23
NATALI, SUSAN M. ........................ 248
NATALIA, NELNIK G. ..................... 150
NATE, NANCY A............................ 317
NCEAS N WORKING GROUP, .......... 212
NDONG OBIANG, SOSTHE`NE .......... 348
NEATOUR, MATTHEW A. ................ 172
NEATROUR, MATTHEW A. .............. 249
NEFF, JASON ................................. 278
NEIRA, CARLOS ............................ 132
NELISSEN, HANS J.M. ......................20
NELSON, DAVID H. ........................ 249
NELSON, DAVID M. ....................... 249
NELSON, JENNIFER A..................... 249
NELSON, KAREN ...................... 71, 249

NELSON, S.................................... 146

NELSON, STEVEN G. ...................... 197
NENZEN, HEDVIG.......................... 249
NESBIT, APRIL D. .......................... 341
NESS, JOSHUA H. .......................... 250
NEUBAUER, SCOTT C. .............123, 250
NEUBAUER, SCOTT ....................... 231
NEUFELD, HOWARD L......................61
NEVAI, ANDREW L. .................250, 341
NEWBOLD, T. A. SCOTT ................. 250
NEWELL, STEVEN Y....................... 226
NEWINGHAM, BETH A. .................. 251
NEWMAN, GREGORY S. ................. 251
NEWMAN, RAYMOND M................. 217
NEWTON, PAUL ............................ 243
NGAI, JACKIE ............................... 251
NGO, HAIVAN V. .............................26
NGUGI, MOFFATT K. ................. 70, 251
NICHOLLS, MELVILLE ................... 273
NICHOLS, JEFFREY V. .................... 313
NIEDZWIECKI, JOHN H................... 285
NIELSEN, KARINA J. ...................... 233
NIELSEN, KARINA ...........................60
NIEMI, GERALD J. ......................... 251
NIEMI, GERALD ..............................79
NIESENBAUM, RICHARD A. ......186, 252
NIKLAS, KARL J. ........................... 224
NIKLAUS, PASCAL A...................... 243
NILES, JOHN O. ............................. 252
NILES, KATHRYN J. ....................... 325
NILON, CHARLES H. ........... 30, 347, 349
NILSEN, ERIK T. ..............................67
NIPPERT, JESSE B. .................... 95, 252
NISBET, ROGER M. ..........................11
NISBETT, RICHARD A..................... 229
NITTA, KEIJI ...................................99
NOORDELOOS, MARCO.................. 284
NORBY, RICHARD J..................252, 371
NORBY, RICHARD ......................... 348
NORLAND, JACK E. ....................... 237
NORRIS, MARK D. ......................... 253
NORRIS, RYAN D........................... 253
NORTH, GRETCHEN B. ................... 335
NORTON, ANDREW P...................... 253
NORTON, ANDREW..................190, 214
NORTON, SUSAN B. ....................... 353
NORTON, URSZULA ....................... 154
NOVAK, DANIEL B. ....................... 109
NOVOTNY, AMY M. ....................... 253
NOVOTNY, AMY............................ 176
NOWAK, JOHN T. .............................75
NOWAK, ROBERT S................... 14, 243
NOWAK, ROBERT ............................. 9
NUFIO, CESAR R. .......................... 253
NUTTER, DOUGLAS A. ................... 254
NUTTLE, TIM ................................ 254
NUZZO, VICTORIA......................... 215
NYBERG, DENNIS W.........................11
NYDICK, KOREN ........................... 254
O
OBRIEN, SEAN ............................. 176
OCONNELL, LISA ......................... 254

OCONNOR, ROBERT E. .................. 307

OMALLEY, RACHEL ...................... 246
ONEILL, ELIZABETH ..................... 348
OBERBAUER, STEVEN F. ..................66
OBERBAUER, STEVEN ................... 172
ODDY, DONNA M........................... 327
OECHEL, WALTER C. .......... 63, 117, 309
OESTERHELD, MARTIN .................. 268
OETTER, DOUG R. ......................... 254
OGDAHL, MEGAN ...................176, 253
OGLE, KIONA ............................... 255
OGLE, STEPHEN M. ....................... 255
OIKAWA, TAKEHISA ...................... 256
OJIMA, DENNIS S. .............. 84, 255, 262
OJIMA, DENNIS ............................. 255
OKUNO, TAKI ............................... 248
OKUYAMA, TOSHINORI ...................38
OLAPADE, OLA A. ......................... 255
OLEKSYN, J ....................................96
OLFF, HAN......................................20
OLI, MADAN K................. 244, 255, 258
OLI, MADAN...................................89
OLIVER, TAYLOR .......................... 306
OLLINGER, SCOTT...........................75
OLSEN, LISA M. ............................ 216
OLSON, BRET E............................. 149
OLSON, JERRY S. .......................... 256
OMETTO, JEAN P. ............................91
ONESIMUS, OETINO....................... 276
OREN, RAM ..................... 182, 223, 325
ORIANS, COLIN............................. 256
ORJUELA, MARIA A. ...................... 285
ORLAND, MARY C......................... 256
ORR, STUART B. ........................... 256
ORR, TERI J. ...........................130, 256
ORROCK, JOHN L. ......................... 257
ORTIZ-BARNEY, ELENA.................. 257
OSBORNE, MEGAN J. .......................27
OSENBERG, CRAIG W..................... 345
OSENBERG, CRAIG ........................ 224
OSIO, GIACOMO C. ........................ 175
OSTERTAG, REBECCA.................... 257
OSTFELD, RICHARD S. ...... 177, 257, 298
OTHERS, 17 .................................. 139
OTIS, DAVID L. ............................. 183
OTSUKA, KOJI .............................. 248
OTT, JAMES R. ................................97
OTTMAR, ROGER .......................... 342
OTZ, INES.......................................74
OUELLETTE, JOHN ........................ 112
OUTCALT, KENNETH W. ...................46
OUTCALT, KENNETH...................... 179
OVADIA, OFER.............................. 257
OVERBY, STEVEN T. ...................... 251
OWEN, DIANNE V. ...................247, 308
OWEN, DIANNE............................. 345
OZGUL, ARPAT ............................. 258
P
PACALA, STEPHEN W. ......................19
PACE, MICHAEL L. ............. 18, 283, 340
PACKER, ALISSA A. ....................... 258
PACKER, ALISSA ...................... 67, 279

PADDOCK, WILLIAM A. ................. 166

PAERL, HANS W. ........................... 258
PAGEL, ALISHA L. ......................... 258
PAHL, JAMES W............................. 258
PAI, ASWINI.................................. 259
PAIGE, KEN N. ......................... 12, 204
PAIGE, TERENCE E. .........................22
PAIGE, TERENCE ........................... 365
PAINE, ROBERT T. ......................... 140
PAINE, TIMOTHY D................... 96, 365
PAL BAIS, HARSH ......................... 218
PALIK, BRIAN J. ............................ 264
PALMER, BRENT D. ....................... 285
PALMER, MARGARET A. ..........233, 249
PALMER, MARGARET ............... 31, 329
PALMER, MATHEW I. ..................... 231
PALOPOLI, MICHAEL F. .................. 110
PALTA, MONICA............................ 235
PALUMBI, STEPHEN....................... 316
PAMPLIN, JACQUE......................... 233
PAN, SHUFEN ............ 209, 259, 335, 371
PANKEY, J. R................................. 259
PAPAIK, MICHAEL J. ...................... 259
PAPAJ, DANIEL R........................... 253
PARADISE, CHRIS J........................ 260
PARDO, LINDA H........................... 260
PARe, DAVID................................. 197
PARHAM, JAMES E. .................260, 320
PARK, RICHARD A...................260, 353
PARKER, CHRISTIAN A. ................. 211
PARKER, GEOFFREY G. ............. 30, 353
PARKER, INGRID M. ........................26
PARKER, JOHN D.............................51
PARKER, SOPHIE S. ....................... 260
PARKER, V.T.................................. 103
PARKS, CATHERINE G. .....................95
PARMENTER, BOB ...........................82
PARMENTER, ROBERT R.... 114, 260, 371
PARMENTER, ROBERT ................... 143
PARMESAN, CAMILLE.................... 261
PARRY HECHT, BROOKE ................ 261
PARRY, DYLAN ............................. 261
PARSHALL, TIM E.......................... 261
PARSONS, GEORGE R..................... 279
PARSONS, WILLIAM F.J................... 208
PARTON, WILLIAM J. ................ 84, 262
PARTOW, HASSAN ......................... 322
PARUELO, JOSE M. ................... 43, 268
PARYSOW, PABLO ...................229, 231
PASCARELLA, JOHN B. .................. 262
PASCUAL, MERCEDES.......188, 262, 280,
289
PASSAFARO, STEVE....................... 290
PASSMORE, HEATHER A................. 262
PATAKI, DIANE E.....................243, 262
PATIL, G. P. ................................... 349
PATTEN, BERNARD C. ..............105, 262
PATTEN, DUNCAN T. ...................... 300
PATTERSON, KATHRYN L. .............. 280
PATTERSON, RANDY L. ............. 22, 130
PATTERSON, RANDY ..................... 365
PAUL, ELDOR A. .............................45
PAUL, VALERIE J........................... 333

PAULK, ERIC ..................................65

PAVAO-ZUCKERMAN, MITCHELL
A. ................................................ 263
PAWLOWSKI, CHRISTOPHER W. ..... 221,
263
PAYTAN, ADINA............................ 270
PEARLSTINE, LEONARD G. ............. 263
PEARS, SARAH L. .......................... 264
PEARSON, SCOTT F. ....................... 194
PEARSON, SCOTT M. ..................... 113
PEART, DAVID R. ........................... 103
PECHMANN, JOSEPH H. K............... 263
PECK, AMANDA E. ........................ 264
PECOT, STEPHEN D. ....................... 264
PEDERSEN, AMY B. ....................... 264
PEDERSEN, BRIAN S. ..................... 264
PEDERSON, NEIL........................... 264
PEEK, MICHAEL S. ........... 201, 265, 293
PEET, ROBERT K........ 114, 127, 265, 342,
355, 366
PEFFLEY, ELLEN B......................... 341
PEGG, MARK A. ..............................64
PEGORARO, EMILIANO .................. 265
PELLANT, MIKE ............................ 148
PELLATT, MARLOW G. ................... 265
PELTZER, DUANE A. ...................... 130
PENDALL, ELISE ........................... 266
PENDLETON, BURTON K. ............... 266
PENDLETON, ROSEMARY L. ........... 266
PENG, CHANGHUI ......................... 266
PENG, TSUNG-HUNG...................... 197
PENNINGS, STEVEN C. ...... 151, 226, 266
PENNINGTON, DEANA D. ............... 371
PERFECTO, IVETTE........................ 267
PERKINS, ALISON E. H. ............. 46, 267
PERKINS, DANA L. ..........................95
PERRONE, MARC........................... 334
PERRY, MICHEAL E.L. ......................18
PERSSON, LENNART ...................... 213
PETERJOHN, WILLIAM T................. 216
PETERS, DEBRA P.C. ...................... 151
PETERS, DEBRA P. ......................... 238
PETERS, HALTON A. ...................... 267
PETERSEN, JOHN E. ....................... 267
PETERSON, CHRIS J. ...................... 200
PETERSON, CHRIS ......................... 135
PETERSON, GARRY D.......................28
PETERSON, GARRY .......................... 9
PETERSON, GARY A....................... 244
PETERSON, LAURA A..................... 147
PETES, LAURA ................................50
PEZESHKI, S REZA ..........................66
PFAFF, LIZA E. .............................. 203
PHILLIPS, NATHAN ........................ 120
PHILPOT, WILLIAM ........................ 181
PHILPOTT, STACY M. ..................... 267
PICKETT, STEWARD T A................. 232
PICKETT, STEWARD T.A....... 54, 285, 302
PIELKE, ROGER A............................51
PIENITZ, REINHARD ...................... 200
PIERCE, DANIELLE ........................ 207
PINCKNEY, JAMES L. ..................... 258
PINDER, JOHN E. ........................... 169

Author Index

389

PINEL-ALLOUL, BERNADETTE ........ 202

EIRO, GERVASIO ...................... 268
PIN
PINGALI, PRABHU ...........................28
PIPPEN, JEFFREY S. ....................... 268
PIPPEN, JEFFREY........................... 241
PIPPEN, WAYNE E.......................... 211
PITHER, JASON ............................. 268
PLATT, BILL............................. 26, 268
PLATT, ROBERT............................. 204
PLATT, WILLIAM J. .... 116, 122, 159, 262,
369
PLAUT, JENNIFER .......................... 219
PLOG, LAURA B. ........................... 268
POCKMAN, WILL T. ....................... 226
POCKMAN, WILL........................... 225
POFF, LEROY ................................ 353
POHLAD, BOB R. ........................... 333
POLING, TARA C. .......................... 165
POLLEY, H. W................................ 243
POLLEY, H. WAYNE ....................... 122
POLLEY, WAYNE ........................... 269
POLLMANN, WILLIAM ................... 269
POLLOCK, KENNETH H. ...................19
POLSENBERG, JOHANNA F.............. 269
PONOMARENKO, ELENA ................ 269
PONOMARENKO, SERGUEI .......269, 270
PONTIUS, JEFFREY S...................... 283
POORTER, LOURENS...................... 270
POPE, KEVIN ................................ 279
POPLE, ANTHONY ......................... 333
PORDER, STEPHEN ........................ 270
PORTER, JAMES W. ........................ 280
PORTER, WARREN P. ...................... 294
POSADA, JUAN M. ......................... 270
POSSINGHAM, HUGH P. .................. 333
POST, DAVID M. ............................ 270
POST, ERIC S................................. 118
POST, ERIC .....................................98
POST, WILFRED M. ........................ 164
POTEET, MARY F. .......................... 271
POTOSNAK, MARK ........................ 185
POTTS, DANIEL L. .................... 62, 271
POULTER, BENJAMIN..................... 271
POUYAT, RICHARD V........................30
POVAK, NICHOLAS A..................... 115
POWELL, THOMAS ................... 43, 271
POWER, ALISON G............ 199, 238, 271
POWER, MARY E. .......................... 230
PRASAD, ANANTHA M. ............. 63, 164
PRATER, MARGARET R. ................. 271
PRATT, MARNEY C. ....................... 272
PRATT, PAUL D.............................. 113
PRATT, R BRANDON ...................... 272
PRATT, R. BRANDON................. 35, 102
PREGITZER, KURT S. ................ 84, 211
PREGITZER, KURT ...................192, 348
PREISSER, EVAN L.................... 24, 272
PRENGER, JOSEPH......................... 272
PRENTICE, COLIN I. ....................... 141
PRICE, CHARLES A. ....................... 273
PRICE, JENNIFER E. ....................... 273
PRICE, K. P.................................... 245
PRICE, KEVIN ............................... 175

390

Author Index

PRICE, STEVE ............................... 139

PRIHODKO, LARA ......................... 273
PRINGLE, CATHERINE M. .........128, 181
PRINGLE, CATHERINE.................... 273
PRINGLE, CATHY .......................... 118
PRINN, RONALD G......................... 372
PROSTKO, ERIC P.............................51
PRYSBY, MICHELLE D.................... 273
PSUTY, NORBERT P. ....................... 274
PUBANZ, DAN M. .......................... 217
PUGNAIRE, FRANCISCO I. .............. 211
PULLIAM, H. R. ............................. 121
PULLIAM, H. RONALD .....................88
PUTZ, BRIAN C. ............................ 340
PUTZ, FRANCIS E. ......................... 342
PYKE, CHRISTOPHER R. ................. 274
PYKE, DAVID A. ............................ 148
PYKE, DAVID................................ 176
PYLE, ERIC J................................. 355
PYPKER, THOMAS ......................... 120
Q
QI, SHI ......................................... 274
QI, YE .......................................... 331
QIAN, SONG S. .............................. 281
QIAN, YONGHAI..............................63
QUEREJETA, JOSE IGNACIO..............97
QUINTANA-ASCENCIO, PEDRO F...... 274
R
RACHEL, GARY............................. 274
RACZ, GABOR R............................ 256
RAFERT, GREG ................................55
RAGUSO, ROBERT A. .......................48
RAHMAN, FAIZ ............................. 117
RAICH, JAMES W. ..................... 91, 275
RAIMONDI, PETER T. ..................... 275
RAINS, KAI C. ............................... 275
RAJANIEMI, TARA K. ..................... 275
RAM, SETHA DR............................ 337
RAMSPOTT, M. E. .......................... 245
RAMSPOTT, MATTHEW .................. 175
RANDALL, DAVID ......................... 276
RANDALL, JESSE A........................ 275
RANDALL, LORI......................229, 357
RANDERSON, JAMES T. ....................76
RANDOLPH, J.C. ............................ 353
RANDOLPH, JC.............................. 350
RANDRIAMAHEFA, ALEXANDRINE.. 276
RANVESTEL, ANTHONY W.............. 181
RASCHER, UWE ............................ 276
RASHLEIGH, BRENDA.................... 276
RASMUSSEN, JOSEPH B....................48
RASTETTER, EDWARD B. ............... 315
RASTETTER, EDWARD ................... 185
RATCLIFFE, LAURENE M. ............... 253
RAVE, DAVID P................................80
RAVIT, ELIZABETH ........................ 273
RAY, CHRIS .................................. 276
READ, BRIDGETT ............................95
REAGEL, PETER F. ......................... 276
REAGON, MICHAEL ....................... 171

REAL, LESLIE A. .....................277, 292

REALE, JOSEPH A............................35
REASER, JAMIE K.......................... 235
REASONER, MEL ........................... 200
REBBECK, JOANNE..................231, 277
REBEL, KARIN T. ........................... 277
REDDY, SRINIVAS ......................... 352
REDMAN, CHARLES L.................... 183
REED, DAVID L. ............... 277, 294, 325
REED, SETH.................................. 320
REES, MARK................................. 204
REEVES, GORDIE H. ........................29
REGAL, RONALD R. ....................... 251
REGAN, HELEN M. ........................ 278
REHEIS, MARITH C. ....................... 278
REICE, SETH R. ............................. 278
REICH, CHRISTINE ........................ 219
REICH, PAUL ................................ 196
REICH, PETER B. ..... 75, 88, 152, 253, 355
REICH, P ........................................96
REICHERT, PETER............................39
REICHMAN, JIM ............................ 294
REICHMAN, O. J. ........................... 350
REID, CHANTAL D. ........................ 157
REID, MARY L. ............................. 309
REIGHARD, GREGORY L................. 331
REILEY, DAVID K. ...................199, 278
REILLY, JOHN ............................... 209
REILLY, MATT............................... 268
REINERS, WILLIAM A. ................... 107
REINHARDT, CARRIE H. ................. 278
REINHART, KURT O. ...................... 279
REINHART, KURT .......................... 134
REINIKAINEN, MARKO .................. 213
REITER, MICHAEL A. ..................... 279
NKOVA
, ELISKA.................. 279
REJMA
RELYEA, RICK A. .. 17, 156, 217, 279, 300
RENNE, IAN J................................ 279
RENWICK, WILLIAM H. ....13, 41, 87, 342
RESETARITS, WILLIAM J. .................33
RETZLAFF, WILLIAM ..................... 343
REYNOLDS, BARBARA C. ............... 280
REYNOLDS, HEATHER L................. 142
REYNOLDS, JAMES F................255, 296
REYNOLDS, RICHARD.................... 278
RHAM, JRENE ............................... 242
RHEIS, HAROLD F. ......................... 280
RHOADS, ANNE G. ........................ 280
RHODES, B. .................................. 146
RICCARDI, CYNTHIA L................... 280
RICHARDS, CHRISTINA L. .............. 280
RICHARDS, JAMES H. ............... 44, 166
RICHARDS, JEANNINE .............205, 220
RICHARDS, RENEE ..........................14
RICHARDS, SHANE A. .................... 280
RICHARDSON, BARBARA ............... 318
RICHARDSON, C J.......................... 329
RICHARDSON, CURTIS J. .... 49, 142, 258,
281
RICHARDSON, ELIZABETH ............. 235
RICHARDSON, LAURIE L. ............... 346
RICHARDSON, MAXWELL J. ........... 296
RICHARDSON, SARA J.................... 130

RICHARDSON, SARAH J. ................ 281

RICHTER, BRIAN ........................... 235
RICKETTS, TAYLOR H. .............162, 281
RIEDER, JULIE P. ........................... 281
RIGGS, MICHAEL R................... 80, 281
RIHA, SUSAN J. ............................. 277
RIHA, SUSAN ................................ 181
RILEY, MELISSA B. ........................ 331
RILEY, WILLIAM J. ..........................71
RILLIG, MATTHIAS C. .................... 282
RILOV, GIL ................................... 282
RINGELBERG, CAROL S. ..................64
RINGOLD, PAUL L. ........................ 282
RINKES, ZACHARY L. .................... 282
RIOS, BIANCA G............................ 279
RITCHIE, JERRY C. ........................ 282
RITCHIE, MARK E.......................... 194
RITLAND, DAVID B........................ 283
RITLAND, KERMIT ........................ 254
RITTER, AMY F.............................. 283
RIVERS, JAMES W.......................... 283
RIXEN, CHRISTIAN ........................ 283
RIZZO, DAVID M. ..................... 81, 246
ROBERT, JONES H.......................... 348
ROBERTS, BRIAN J. ....................... 283
ROBERTS, EMILY .......................... 229
ROBERTS, KELLY A. ...................... 283
ROBERTSON, G. P. ......................... 261
ROBERTSON, G. PHILIP .................. 128
ROBERTSON, KEVIN M................... 284
ROBERTSON, PHILIP A. ....................60
ROBERTSON, TRAESHA R............... 284
ROBINS, JAMES H.......................... 124
ROBINSON, BRADFORD A. ............. 371
ROBINSON, JULIE A. ................102, 284
ROBINSON, TODD M.P. ................... 284
ROBINSON, W. DOUGLAS ............... 326
ROBISON, LAURA A....................... 285
ROBITAILLE, GILLES ..................... 197
RODD, F. HELEN............................ 201
RODRIGUEZ, SUSANA.................... 285
RODRIGUEZ-ITURBE, IGNACIO .........59
RODRIGUEZ-SAONA, CESAR R. ....... 285
ROGERS, ALISTAIR..................137, 200
ROGERS, KEVIN H. ........................ 285
ROGSTAD, STEVEN H..................... 223
ROHANI, PEJMAN....................285, 351
ROHR, JASON R............................. 285
ROHRS, JENNIFER K. ..................... 286
ROLAND, JENS................................67
ROLLINS, SCOTT L. .........................27
ROMANEK, CHRISTOPHER S. .......... 169
ROMANUK, TAMARA N.................. 286
ROMME, WILLIAM H. .................... 234
ROONEY, THOMAS P. .......................11
ROOSENBURG, WILLEM M. ............ 286
ROOT, KAREN V. ........................... 286
ROSE, JOSHUA S. .......................... 286
ROSENBERG, ANDREW .................. 212
ROSENFELD, KRISTEN M................ 287
ROSENTHAL, DAVID M. ................. 287
ROSENTHAL, SADIE K.................... 287
ROSENZWEIG, MICHAEL L. ............ 273

ROSIER, CARL L............................ 282

ROSNEBERG, DANIEL K. ..................59
ROSS, KATIE................................. 195
ROSS, MICHAEL S. .... 117, 172, 189, 287,
294, 325, 370
ROSS, MIKE .................................. 167
ROSSELL, IRENE ........................... 350
ROSSI, ANTHONY M. ...............288, 330
ROSSON, JAMES F.......................... 288
ROSSON, JESSICA L. ...................... 288
ROTH, AGATHA-MARIE .................. 288
ROTH, BRIAN M. .....................146, 288
ROTH, JAMES D. ........................... 268
ROTH, SUSAN ............................... 127
ROTHSTEIN, DAVID E. ................... 369
ROUGHGARDEN, JOAN E...... 15, 50, 289
ROULSTON, TAI H. .................. 57, 289
ROY, ALLISON H. .......................... 354
ROY, B. A. .................................... 245
ROY, BITTY A. .............................. 289
ROY, MANOJIT .............................. 289
ROYCE, GITANE L. ........................ 289
ROYCE, GITANE ............................ 169
ROYO, ALEJANDRO A. ................... 290
RUBBO, MICHAEL J. ...................... 166
RUBECK-SCHURTZ, NICHOLE ...........86
RUBINO, LUCY M. ......................... 290
RUBINO, LUCY ............................. 219
RUBIO, MARIA G........................... 290
RUDGERS, JENNIFER A. ...........190, 290
RUEDAS, MARCELA ...................... 290
RUIZ, PABLO L................. 287, 294, 325
RUIZ, PABLO ................................ 117
RUIZ-BERNARD, IVELISSE ........135, 291
RUIZ-GUTIERREZ, VIVIANA......291, 304
RUMBAITIS-DEL RIO, CRISTINA M. 291
RUNNING, STEVEN W..................... 174
RUOFF, RACHEL H. ........................ 291
RUSS, ANDREW L.......................... 292
RUSSELL, ANN E. .......................... 292
RUSSELL, COLIN A. ....................... 292
RUSSELL, JOHN ............................ 254
RUSSELL, ROBIN E. ....................... 292
RUSSO, SABRINA E........................ 292
RUTHIG, GREGORY R..................... 293
RYAN, MICHAEL G. ....................... 293
RYAN-SCHNEIDER, SUSAN ............. 167
RYEL, RON J. ................................ 265
RYEL, RONALD J. ....................201, 293
RYGIEWICZ, PAUL......................... 229
RYKIEL, EDWARD J. ...................... 293
S
S.K., RHEE .................................... 347
SAAH, DAVID S. ............................ 294
SABO, JOHN L............................... 294
SACCHI, CHRISTOPHER F................ 294
SACK, LAWREN ............................ 256
SADLER, KIM C............................. 294
SADOF, CLIFFORD S.........................28
SAGARIN, RAFAEL ........................ 175
SAH, JAY P.................................... 294
SAH, JAY...................................... 167

SAH, SEEMA ................................. 167

SAIT, STEVEN M. ........................... 351
SALA, OSVALDO E. .......................... 6
SALANT, STEPHEN W. .................... 193
SALAZAR-BRAVO, JORGE............... 363
SALGADO, CRISTIANO S. ............... 151
SALIENDRA, N. ............................. 255
SALK, CARL F. ................................17
SALKELD, DAN ............................. 295
SALO, LUCINDA F.......................... 295
SALTONSTALL, KRISTIN................. 295
SAMBROTTO, RAYMOND ............... 367
SAMUEL, MICHAEL .......................... 6
SAMUELS, IVAN A......................... 295
SAMUELSON, LISA J. ........ 105, 209, 324
NCHEZ-PIN
ERO, FRANCISCO.. 90, 162
SA
SANDERS, NATHAN J. .................... 295
SANDERS, SUZANNE M. ................. 295
SANDERSON, MATT A. ................6, 126
SANDERSON, MATT.........................70
SANDQUIST, DARREN .................... 354
SANFORD, JR, ROBERT L................ 336
SANFORD, ROBERT L........227, 278, 296,
362
SANFORD, ROBERT ....................... 255
SANKARAN, MAHESH..............138, 296
SANTIAGO, LOUIS S.................296, 308
SARGENT, KENNETH...................... 205
SARGENT, SARAH .................... 38, 194
SARNELLE, ORLANDO ................... 296
SAUER, SUZANNE L....................... 296
SAUNDERS, COLIN J. ................ 36, 296
SAUNDERS, THOMAS J................... 297
SAVAGE, HARRY M. ...................... 340
SAVAGE, VAN M. .......................... 297
SAX, DOV F................................... 297
SCARDAMALIA-NELSON, CYNTHIA
B. ................................................ 249
SCATENA, FREDERICK N. .................25
SCHAAFSMA, HOSKI...................... 297
SCHABERG, PAUL G. .......................38
SCHACHT, WALTER W.H. ..................18
SCHADE, JOHN D. ...................253, 297
SCHAEFFER, SEAN M..................... 298
SCHALLES, JOHN .......................... 151
SCHAMP, BRANDON S. ................... 302
SCHAUB, MICHAEL ....................... 299
SCHAUBER, ERIC M....................... 298
SCHAUER, ANDREW ...................... 120
SCHEESSELE, ERIN A. .................... 298
SCHEINER, SAMUEL M................... 298
SCHEINER, SAMUEL ...................... 323
SCHEMSKE, DOUGLAS W................ 155
SCHERSON, ROSA ......................... 365
SCHEUERELL, MARK D. ................. 242
SCHIEL, DAVID R. ...................298, 332
SCHILE, LISA M. ........................... 299
SCHILLING, ERIK B. ...................... 299
SCHIMEL, DAVID S. .........................16
SCHIMEL, JOSHUA P. ........ 236, 260, 352
SCHINDLER, DANIEL E...... 112, 138, 242
SCHLESINGER, WILLIAM H......... 22, 83,
108, 151, 207, 239, 268, 299, 331, 347, 371

Author Index

391

SCHLIEMANN, SARAH A. ............... 303

SCHMALZER, PAUL A. .............111, 299
SCHMID, BERNHARD ..................... 362
SCHMID, HANS PETER ................... 350
SCHMIDT, BENEDIKT ..................... 299
SCHMITZ, DENINE M. .................... 300
SCHMITZ, OSWALD ....................... 257
SCHMITZ, RICHARD A. .................. 228
SCHNABEL, ANDREW F. ................. 256
SCHNEIDER, DAVID C. ................... 300
SCHNURR, JACLYN L. .................... 300
SCHOCH, CARL ................................ 9
SCHOENNAGEL, TANIA.................. 300
SCHOEPPNER, NANCY M................ 300
SCHOETTLE, ANNA W. .............. 38, 301
SCHOFIELD, OSCAR....................... 208
SCHOLBERG, JOHANNES M. ........... 212
SCHOOLMASTER, DONALD R.......... 301
SCHOONOVER, JON E. ................... 301
SCHORRAN, DAVID .................. 16, 343
SCHROMEN-WAWRIN, LINDSEY ...... 267
SCHRUM, KIM............................... 307
SCHULER, AZENETH...................... 217
SCHULTZ, PEGGY A. ................. 32, 142
SCHULZ, KURT E........................... 301
SCHULZ, KURT ...............................51
SCHUPP, EUGENE W. ................160, 301
SCHURER, MAAYKE ...................... 302
SCHUSTER, WILLIAM ....... 220, 336, 339
SCHUUR, EDWARD A G.................. 338
SCHUUR, EDWARD AG................... 308
SCHWARTZ, MARK W..................... 164
SCHWARTZ, STEVE S. .................... 302
SCHWARZ, KIRSTEN ...................... 302
SCHWENDENMANN, LUITGARD ...... 302
SCHWILK, DYLAN W...................... 302
SCHWILK, DYLAN ............................ 5
SCINTO, LEONARD J. ..................... 334
SCOBELL, SUMMER A.................... 303
SCOFIELD, DOUGLAS G.................. 303
SCOTT, DAVID E. .....................263, 331
SCOTT, FINLAY A M ...................... 303
SCOTT, PETER E. .............................78
SCOTT, RUSSELL L. ....................... 314
SCOTT-DENTON, LAURA E.............. 303
SCOW, KATE M................................93
SEABLOOM, ERIC W. ..................... 303
SEABLOOM, ERIC..................... 39, 359
SEAMAN, JOHN............................. 277
SEARS, ANNA L. W. ....................... 304
SEARS, ROBIN R............................ 304
SEAVY, NATHANIEL.........................38
SEHTIYA, HARBANS L. .................. 308
SEIGLE, JOHN ............................... 109
SEKERCIOGLU, CAGAN H.........291, 304
SEKINE, MASAHIKO ................175, 350
SELIG, ELIZABETH R. .................... 266
SELMANTS, PAUL C....................... 304
SEMENOV, V M. ............................ 192
SENDEL, TANIA T. ......................... 304
SENDZIMIR, JAN .............................. 9
SENFT, AMANDA........................... 114
SENGUPTA-GOPLAN, CHAMPA..........69

392

Author Index

SENOCK, RANDY ............................34

SERGEY, SHUBENKOV G. ............... 150
SERMONS, SHANNON ......................51
SETHI, SURESH ............................. 175
SEYMOUR, MICHAEL A. ................. 305
SEZEN, ZEYNEP ............................ 305
SHAFER, SARAH ........................... 359
SHANDAS, VIVEK.......................... 358
SHAPIRO, JOSSLYN B..................... 305
SHARITZ, REBECCA R...92, 169, 301, 305
SHARITZ, REBECCA.................215, 235
SHARMA, BRIJ MOHAN DR............. 337
SHARSHIR, FARRG ........................ 330
SHAVER, PATRICK ......................... 148
SHAW, JOSEPH ................................68
SHAW, M. R.....................................94
SHAW, REBECCA ........................... 305
SHEA, KATHLEEN L. ........................23
SHEA, KATRIONA .......................... 305
SHEAR, THEODORE H. ................... 343
SHEIK, CODY................................ 306
SHELDON, JOAN ........................... 235
SHELTON, ANDREW O..............175, 235
SHEPHERD, BRIAN S...................... 285
SHEPPARD, MICHAEL E.................. 366
SHEPPARD, WALTER S. .................. 325
SHEPPERD, WAYNE D. ................24, 38
SHEPPERD, WAYNE .................203, 311
SHER, ANNA A.............................. 306
SHERICK, MICHAEL....................... 320
SHERRY, REBECCA A. ..............306, 347
SHERRY, THOMAS W...................... 253
SHIPLEY, BILL ........................306, 344
SHIRER, REBECCA R........................89
SHIRK, JENNIFER L........................ 306
SHOLTIS, JOHNNA D. ..................... 252
SHORTLE, JAMES S........................ 307
SHOWALTER, JULIA M. .................. 172
SHRESTHA, JUNO ............................97
SHUGART, HERMAN H. ........ 55, 59, 315
SHUMAN, BRYAN N....................... 307
SHUMAN, BRYAN............................55
SHURETTE, RYAN ......................... 307
SHURIN, JONATHAN ........................39
SICCAMA, THOMAS C. .....................25
SICKMAN, JAMES O....................... 236
SIEDSCHLAG, SARAH .................... 307
SIEVING, KATHRYN E. .............. 79, 307
SIGMAN, DANIEL M....................... 155
SIGURDARDOTTIR, RAGNHILDUR .....25
SIH, ANDREW ............................... 307
SIH, ANDY.................................... 285
SIKAROODI, MASOUMEH ............... 123
SILANDER, JOHN A........................ 202
SILIM, SALIM................................ 308
SILLIMAN, BRIAN R......31, 201, 245, 308
SILMAN, MILES R.......................... 366
SILVER, WHENDEE ........................ 332
SILVERA, KATIA ........................... 308
SIMMONS, JEFFREY A.................... 308
SIMMONS, STEVEN P................149, 308
SIMONS, THEODORE R. ....................19
SIMONSON, SARA E....................... 309

SIMPSON, COLLEEN M. .................. 309

SIMS, DANIEL A. ........................... 309
SIMS, DANIEL ............................... 117
SINGER, FRANCIS J..........................33
SINGER, MICHAEL S. ..................... 309
SINSABAUGH, ROBERT L. ........108, 173
SINSABAUGH, ROBERT .................. 242
SIQUEIRA, MARIO B.S. ................... 325
SKELLY, DAVID K.......................... 309
SKELLY, DAVID............................. 113
SKINNER, HOWARD.........................70
SLAYBACK, DANIEL ...................... 359
SLOCUM, MATTHEW G................... 232
SLONE, DANIEL H. ........................ 310
SLOTHOWER, ROGER .................... 155
SMALL, ERIC E. ............................ 314
SMALLWOOD, PETER..................... 320
SMEE, DELBERT L. ..................106, 310
SMILEY, PETER C. ......................... 310
SMITH, ANNEMARIE...................... 310
SMITH, BRUCE N.............. 177, 310, 335
SMITH, BRUCE.............................. 328
SMITH, CELIA M. ............................34
SMITH, CHARLES .......................... 139
SMITH, CURTIS A. ......................... 311
SMITH, DAVID L......................292, 311
SMITH, DENA M. ........................... 311
SMITH, DIXIE L. ............................ 311
SMITH, ELIZABETH A. ................... 311
SMITH, FREDERICK W. .............203, 311
SMITH, JAMES L............................ 312
SMITH, JAPHIA ............................. 312
SMITH, MARIAN............................ 312
SMITH, MELINDA D. ....... 58, 59, 99, 252,
283, 312
SMITH, RACHEL N......................... 312
SMITH, RICHARD G. ................133, 313
SMITH, RUTH M. .............................47
SMITH, SARAH A........................... 313
SMITH, SCOT E. ..............................34
SMITH, SHAWN E.L........................ 313
SMITH, SIGRID.............................. 256
SMITH, STANLEY D. ...................... 373
SMITH, STANLEY .......................... 248
SMITH, STEPHEN A........................ 289
SMITH, VAL H.......................... 95, 175
SMITH, WILLIAM J. ....................... 121
SMITH, WILLIAM K. .....47, 138, 143, 169
SMITH, WINSTON P. ....................... 313
SMITH-LONGOZO, VICKIE L..............14
SMITH-WALTERS, CINDI................. 294
SMOL, JOHN P. .............................. 200
SNAMAN, JENNIFER M. .................. 277
SNETSINGER, CAROL................ 46, 314
SNOVER, MELISSA L...................... 314
SNOW, ALLISON A. ..................171, 199
SNYDER, JAMES R......................... 189
SNYDER, KEIRITH A. ..................... 314
SNYDER, ROBIN E. ........................ 314
SOBALLE, DAVID M....................... 123
SOJA, AMBER J. ............................ 315
SOLOMESHCH, AYZIK I. ................. 315
SOLUK, DANIEL A. ........................ 342

SOMERS, GREG L. ...........................61

SOMERSHOE, SCOTT G. ................. 339
SOMMERKORN, MARTIN ..........185, 315
SOMMERVILLE, GERICKE L. ........... 315
SON, YOWHAN ............................. 182
SONG, BO...............................315, 348
SORANNO, PATRICIA A. ................. 317
SORENSEN, MARY A...................... 329
SORRIE, BRUCE A. ........................ 316
SOSNOSKIE, LYNN M. .................... 316
SOTKA, ERIK ................................ 316
SOUTER, NICHOLAS J. ................... 316
SOUTHER, REBECCA F. .................. 316
SPENCE CHERUVELIL, KENDRA...... 317
SPENCER, DAVE............................ 306
SPERRY, JOHN S. ........................... 225
SPICER, RACHEL ........................... 317
SPILLER, DAVID A. ........................ 317
SPINELLI, GIANFRANCO ................ 317
SPIRA, TIMOTHY P...................332, 371
SPONSELLER, RYAN A. .................. 317
SPONSELLER, RYAN ...................... 297
SPRAGGINS, ALAN ........................ 284
SPRINGER, CLINT J..................318, 355
SPROTT, PATRICIA L. ..................... 318
SRIVASTAVA, DIANE S. .................. 318
ST. CLAIR, J. BRAD ........................ 177
ST. CLAIR, LARRY L. ..................... 266
ST.CLAIR, SAMUEL B. .................... 318
STABLER, BROOKE L. .................... 130
STABLER, LINDA B........................ 205
STACKHOUSE, PAUL W. ................. 315
STAFFORD, NATHANIEL B. ............. 318
STAL, DEREK................................ 219
STANLEY, EMILY H........................ 110
STANLEY, THOMAS R. ................... 318
STANOSZ, GLEN R. ........................ 101
STANTON, L.................................. 369
STANTON, MAUREEN L.................. 365
STANTURF, JOHN........................... 324
STARK, JOHN M. .....................154, 319
STARK, JOHN................................ 193
STARK, SCOTT C. .......................... 319
STARNES, JOHN H. ........................ 319
STARNES, SARAH M. ..................... 319
STARR, GREGORY .................... 34, 319
STEELE, KAREN E.H. .......................30
STEELE, MICHAEL A...................... 320
STEEN, HARALD .............................81
STEETS, JANETTE A....................... 320
STEFANOVA, IVANKA .................... 249
STEGEN, JAMES C. ........................ 320
STEINAKER, DIEGO F. .................... 320
STEINAUER, MICHELLE L............... 320
STEINBERG, ELEANOR................... 232
STEINER, CHRISTOPHER F. .......202, 321
STEINER, KIM C. ........................... 105
STEINER, S. .................................. 368
STEINER, SUSAN M. ...................... 321
STEMBERGER, RICHARD S. ............ 335
STEPHEN, MARJORIE ..................... 197
STEPHEN, PORTNOY ...................... 292
STEPHENS, LAURA S. .................... 115

STEPHENSON, ANDREW G. .......321, 361

STEPHENSON, NATHAN L. .............. 341
STEPHENSON, ROBERT E................ 321
STERNBERG, LEONEL .................... 129
STERNER, ROBERT W...............176, 321
STEUDLER, PAUL A. ...................... 372
STEVENS, GLEN N. ..................172, 322
STEVENS, MARTIN HENRY H. ......... 322
STEVENS, MICHAEL T. ................... 322
STEVENS, MICHELLE L. ................. 322
STEVENS, RICHARD D. .................. 322
STEVENSON, BLAKE K................... 323
STEVENSON, R. JAN ................211, 372
STEWART, JOY E. .......................... 345
STIEGLITZ, MARC ...........................37
STILES, ARTHUR ........................... 323
STILES, EDMUND W. ........................64
STILING, PETER ...... 22, 72, 135, 288, 323
STILLER, VOLKER ......................... 287
STILLWELL, CRAIG ....................... 323
STIMMLER, BRYAN .................307, 323
STIMSON, HUGH C. ....................... 324
STINSON, KRISTINA A.................... 324
STIREMAN, JOHN O. ...................... 145
STOCK, WILLIAM D. ...................... 330
STODOLA, KIRK W......................... 324
STOECKLI, VERONIKA ................... 362
STOHLGREN, THOMAS J.... 174, 309, 324
STOKER, JASON M......................... 136
STOKES, THOMAS A. ...............105, 324
STOMS, DAVID M. ......................... 303
STOMS, DAVID.............................. 359
STONE, PETER A. .......................... 325
STOUT, I. JACK ...............................66
STOVER, DANIEL B. ...................... 325
STOY, PAUL C. .............................. 325
STRAHM, SPRING ............................78
STRAND, ALLAN E. ....................... 325
STRANGE, JAMES P........................ 325
STRATFORD, JEFFREY A................. 326
STRECKER, ANGELA L. .................. 326
STRENGBOM, JOACHIM ................. 326
STROMBERG, JULIET C. ....... 25, 40, 257
STRONG, DON R..............................24
STRONG, DONALD R................272, 326
STUART, GLENN............................ 326
STUEFER, JOSEF F.......................... 155
STUREK, MICHAEL..........................46
STYRING, ALISON R. ..................... 327
STYRSKY, JOHN D. ........................ 327
SU, BO ...................................327, 351
SUAZO, ALEXIS A. ........................ 327
SUBERVIELLE, STACY ................... 327
SUBRAMANIAM, BANU....................32
SUDING, KATHARINE N.................. 328
SUELTER, TYLER.............................67
SUGDEN-NEWBERY, ANNA E. ......... 328
SUKHININ, ANATOLY..................... 315
SUKKESTAD, KATHRYN E. ............. 328
SULLIVAN, J. H. ............................ 240
SULLIVAN, MICHAEL J................... 247
SULTAN, SONIA E. ......................... 131
SUMMERS, HEIDI .......................... 328

SUN, GE ....................................... 274

SUNDARESHWAR, P V. ................... 329
SUNDARESHWAR, P.V. .................... 243
SUNDARESHWAR, PALLAOOR V. ..... 142
SUTHERLAND, ANDREW ................ 118
SUTHERLAND, ELAINE K. .............. 161
SUTTER, ROBERT D. ...................... 329
SUTTLE, BLAKE ............................ 334
SUTTLE, KENWYN B. ..................... 329
SVENSON, ELIZABETH L. .................36
SWAN, CHRIS................................ 329
SWANEY, DENNIS .......................... 153
SWANK, W. T................................. 368
SWANK, WAYNE T. ........................ 185
SWANSON, BROOK O. .................... 329
SWARBRICK, SUSAN...................... 245
SWART, CHARLES C....................... 330
SWEET, SARA ............................... 306
SWEMMER, TONY M...................... 330
SWENSON, JANICE E...................... 330
SWETNAM, THOMAS W. ................. 213
SWIFT, CHERYL ......... 195, 249, 307, 323
SWIHART, ROBERT K. ....... 124, 242, 292
SYMS, CRAIG .................................57
SZLAVECZ, KATALIN .......................30
T
TADA, KAZUFUMI ......................... 330
TADROS, MOHSEN 11351 ................ 330
TAILLIE, CHARLES ........................ 349
TAINTER, JOSEPH A....................... 331
TALLEY, DREW M.......................... 162
TANEVA, LINA.............................. 331
TANG, JIANWU ............................. 331
TANG, Y-H.................................... 255
TANNER, SHANN C. ....................... 331
TAPER, MARK L. ........................... 106
TAUB, DANIEL R. .......................... 176
TAUB, FRIEDA B. .......................... 331
TAVERNA, KRISTIN ....................... 355
TAYLOR, BARBARA E. .............. 84, 331
TAYLOR, CAZ M............................ 332
TAYLOR, CAZ ............................... 326
TAYLOR, DAVID I. ......................... 332
TAYLOR, JASON R. ........................ 151
TAYLOR, JASON ............... 128, 174, 332
TAYLOR, MILTON D. ...................... 332
TAYLOR, PHILLIP D. ...................... 338
TAYLOR, RONALD K. .......................64
TAYLOR, WILLIAM ........................ 191
TEMPLER, PAMELA H. ................... 332
TENHUMBERG, BRIGITTE.........333, 339
TERRY, MYLINDA L. ...................... 197
TESH, R B..................................... 368
TESKEY, ROBERT O. ......... 109, 228, 333
TESSIER, ALAN J. ............................53
TEWKSBURY, JOSHUA J. ..................38
THACKER, ROBERT W. ... 32, 55, 101, 333
THALER, JENNIFER S. ..............130, 285
THAXTON, JARROD .........................93
THELEN, GILES C. ...........................55
THOMAS, CAROLYN L. .................. 333
THOMAS, CASSONDRA R. .............. 262

Author Index

393

THOMAS, KATHRYN A. .................. 333

THOMAS, MICHAEL A. ................... 334
THOMAS, RICHARD B. ................... 318
THOMAS, SERGE ........................... 334
THOMPSON, ANDREW R................. 334
THOMPSON, IAN D. .........................13
THOMSEN, MEREDITH A. .......... 72, 334
THORPE, ANDREA S. ..................... 335
THRALL, PETER H. ........................... 8
THREAT, CALVIN J......................... 335
THROOP, HEATHER L. .................... 115
THUM, RYAN A. ............................ 335
THURSTON, ROBERT ..................... 107
THYGERSON, TONYA..................... 335
TIAN, HANQIN ........... 209, 259, 335, 371
TIAN, JIAN ................................... 249
TIELBOERGER, KATJA ................... 188
TILMAN, D. .................................. 150
TILMAN, DAVID ......................104, 123
TILMAN, D ................................... 140
TIMBERLAKE, JONATHAN .............. 152
TINGIRIS, PAM .............................. 111
TISSUE, DAVID T. ..42, 130, 205, 284, 291,
305, 341
TISSUE, DAVID ............................. 336
TITIZ, BEYHAN ............................. 336
TOLEDO, LIGIA C. ...........................60
TOMAS, ANDREA .......................... 336
TORN, MARGARET S................. 71, 337
TORREALBA, KARIMAR ...................23
TOWNSEND, ALAN R. .................... 336
TOWNSEND, PHILIP A. ................... 336
TOZER, MARK .............................. 278
TRACY, BENJAMIN F. ...............126, 279
TRAHAN, KATHLEEN ..................... 234
TRAN, KIM CHI ............................. 330
TRAUT, BIBIT H............................. 337
TRAVERS, STEVEN E................321, 361
TRESEDER, KATHLEEN K. .... 30, 76, 337
TRETTIN, CARL............................. 206
TREXLER, JOEL C....... 131, 313, 337, 358
TRIBUZY, EDGARD S........................60
TRIPATHI, GHANSHYAM DR............ 337
TROUT, ROGER ............................. 158
TROYER, MICHAEL ....................... 131
TRUDEAU, VANCE......................... 338
TRUEMAN, REBECCA J. ................. 338
TRUMBORE, SUSAN E. .....................60
TRUMBORE, SUSAN....................... 338
TRZCINSKI, KURTIS M. .................. 338
TU, CONG..................................... 372
TUBERVILLE, TRACEY................... 305
TUCKER, BRIAN C. ..........................11
TUCKFIELD, R. CARY TUCKFIELD... 338
TUHELA-REUNING, LAURA ..............86
TULJAPURKAR, SHRIPAD ............... 344
TURCHIN, PETER........................... 339
TURES, ELIZABETH ....................... 225
TURNBULL, MATTHEW H. ......... 42, 130
TURNBULL, MATTHEW .................. 336
TURNER, GREG D. ......................... 220
TURNER, GREG ................ 205, 291, 339

394

Author Index

UDOVIC, DANIEL ....................238, 340

UDOVIC, DAN .........................128, 332
UKITA, MASAO .......................175, 350
ULANOWICZ, ROBERT ................... 191
UNDERWOOD, NORA ..................6, 340
UNMACK, PETER J......................... 103
UNNASCH, THOMAS R. .................. 340
URBAN, DEAN L......................225, 366
USTIN, SUSAN L............................ 324

VIERLING, KERRI T........................ 343

VIERLING, LEE A.....................312, 344
VIGUE, LEANNE............................ 344
VILE, DENIS ................................. 344
VILE, MELANIE A. ......................... 344
VILGALYS, RYTAS......................... 142
VINTON, MARYANN ...................... 344
VIRGINIA, ROSS A. ..........................21
VITONE, NICHOLAS .........................82
VITOUSEK, PETER M. ......... 10, 270, 344
VIVANCO, JORGE M.................218, 251
VIVANCO, LUCIA ............................18
VOGEL, AMBER ............................ 247
VOGELSANG, KEITH M. ................. 345
VOGELSANG, KEITH........................32
VOGT, DANIEL J. .............................25
VOGT, KRISTIINA A. ................. 25, 261
VOLIN, JOHN C. ....94, 149, 165, 211, 247,
308, 345
VON HOLLE, BETSY ...................... 345
VONESH, JAMES R......................... 345
VONESH, JAMES ........................... 224
VOSE, JAMES M. .....................185, 345
VOSS, JOSHUA D. .......................... 346
VUONG, LINH ............................... 257

VACCARO, LYNN .......................... 171

VAITKUS, MILDA R. ...................... 340
VALDES, LEXIA M. ........................ 258
VALIENTE-BANUET, ALFONSO ..........32
VALVERDE, TERESA ........... 31, 168, 290
VAN AUKEN, O. W. ........................ 174
VAN BODEGOM, PETER M. ...............20
VAN DE BOGERT, MATTHEW C. ..... 283,
340
VAN DER MERWE, MARIUS ..............51
VAN DER PUTTEN, WIM H.............. 279
VAN GESTEL, NATASJA C............... 341
VAN HAREN, JOOST ...................... 207
VAN MANTGEM, PHILLIP J. ............ 341
VAN NIEL, KIMBERLY P. ................ 341
VAN SICKLE, JOHN ....................... 282
VAN ZANDT, PETER A.................... 341
VANCE, RICHARD R.................250, 341
VANCE-CHALCRAFT, HEATHER D. ... 342
VANDER ZANDEN, M JAKE ............ 146
VANDER ZANDEN, M. JAKE............ 360
VANDERMAST, DAVID B...... 99, 114, 342
VANHAREN, JOOST ....................... 265
VANNI, MICHAEL J. .........13, 41, 87, 342
VANO, JULIE A................................56
VARNER, J. M. .............................. 342
VAUGHN, KAREN ............................65
VEBLEN, THOMAS T. ..................... 192
VELINSKY, DAVID J. ........................52
VELLEND, MARK .......................... 343
VENTERIS, ERIK R......................... 282
VERBURG, PAUL S......................... 169
VERBURG, PAUL ...................... 16, 343
VERBYLA, DAVID L. ........................55
VERMILLION, BRIAN ..................... 343
VIDRA, REBECCA L. ...................... 343

WACHNICKA, ANNA ...................... 117

WACHTER, PAUWEL ...................... 348
WADDELL, PAUL A. ....................... 237
WAGNER, C. MICHAEL................... 346
WAGNER, DIANE........................... 373
WAGNER, RACHEL C. ......................36
WAHL, EUGENE R.......................... 346
WAHREN, CARL H. ........................ 347
WAIT, ALEXANDER................... 12, 145
WAIT, D. ALEXANDER......................21
WALBRIDGE, MARK R. ............. 93, 346
WALDE, SANDRA J. ....................... 338
WALDROP, THOMAS A. ............. 37, 347
WALKER, AMBER M. ..................... 347
WALKER, DONALD A. .................... 177
WALKER, ERIN M. ......................... 284
WALKER, JOAN L. ...................356, 357
WALKER, JOHN F.............................67
WALKER, MARILYN....................... 347
WALL, DIANA H. .............................21
WALLACE, LINDA L............ 16, 306, 347
WALLACE, LINDA ...................169, 351
WALLACE, MICHAEL P. ............190, 192
WALLENSTEIN, MATTHEW D. ...268, 347
WALLER, DEBORAH A. ............157, 347
WALLER, DONALD M.......................11
WALLER, LANCE A. ....................... 292
WALLER, LANCE........................... 277
WALLS, ELIZABETH A..............132, 161
WALSH, EMILY ............................. 147
WALSH, PETER D. .................... 19, 348
WALTER, ACHIM ........................... 348
WALTERS, JOSHUA ........................ 172
WALTERS, MICHAEL B. .....121, 165, 193,
199, 201, 275
WALTERS, MICHAEL...................... 220

TURNER, MONICA G. .... 11, 56, 113, 234,

300, 328
TURNER, RAYMOND M. ...................44
TURNER, THOMAS F. .......................27
TURNER, WILL R........................... 273
TUTIN, CAROLINE E.G. ....................14
TWEDT, DANIEL J. ......................... 339
TWIGGS, JOHN P............................ 267
TYLER, A. C.................................. 132
TYLER, ANNA C. ........................... 339
TYRE, ANDREW J. ...................333, 339
TYREE, MELVIN T.......................... 193
TYSER, DEBORAH ...........................57
U

WAN, SHIQIANG............... 327, 347, 348

WANG, CHUAKUAN K. ................... 103
WANG, LIQIN..................................97
WANG, LIXIN................................ 348
WANG, WEI .................................. 112
WANG, XIANLI.............................. 348
WANG, XIANZHONG ...................... 348
WANG, YUNG-HO (OPHELIA) .......... 349
WANG, ZHENGQUAN ..................... 206
WARD, BESS B. ............................. 349
WARDROP, DENICE H. .................... 349
WARNER, TIMOTHY A.................... 196
WARREN, PAIGE S. ........................ 349
WARREN, ROBERT J....................... 350
WATANABE, MORIYOSHI ............... 350
WATKINS, JAMES E........................ 350
WATKINSON, JONATHAN................ 244
WATSON, MAXINE A...................... 163
WATTERS, GEORGE ....................... 314
WATTS, SEAN ............................... 350
WATTS, T .......................................58
WAUGAMAN, ROBIN D................... 222
WAYSON, CRAIG ........................... 350
WEAKLEY, ALAN S. ....................... 316
WEAR, DAVID N. ........................... 351
WEAR, DAVID............................... 118
WEARING, HELEN J. ...................... 351
WEATHERLY, HEATHER E. .............. 351
WEATHERLY, HEATHER.................. 289
WEATHERS, KATHLEEN C............... 108
WEAVER, S C. ............................... 368
WEBB, SARA L.............................. 290
WEBB, THOMPSON ........................ 307
WEBBER, MICHELE M.................... 319
WEBSTER, THEODORE M........... 51, 351
WEDIN, DAVID A........................... 253
WEDIN, DAVID.............................. 355
WEEKLEY, CARL W........................ 274
WEICHT, THOMAS R. ..................... 351
WEIHER, EVAN ........................ 46, 352
WEIN, GARY R. ............................. 352
WEINSTEIN, DAVID ....................... 153
WEINTRAUB, MICHAEL N............... 352
WEIR, ENRIQUE H. ........................ 352
WEISER, MICHAEL D. .................... 352
WEISHAMPEL, JOHN F. .............268, 353
WEISS, JOHANNA .......................... 231
WEISSBURG, MARC J. .................... 310
WELCH, NICOLE T. ........................ 353
WELLBORN, GARY A. .................... 353
WELLMAN, MARJORIE C. ............... 353
WELLNITZ, TODD.......................... 353
WELLS, CHRISTINA E...............331, 354
WELTER, JILL R............................. 317
WELTZIN, JAKE F................ 99, 271, 354
WELTZIN, JAKE...............................27
WENDEL, HEATHER E. ................... 354
WENDER, NAOMI J. ....................... 354
WENGER, SETH J........................... 354
WENNERBERG, SARAH B. .............. 355
WENTWORTH, THOMAS R. ...... 127, 128,
185, 197, 287, 355
WERNER, KIRWIN ......................... 171

WESSMAN, CAROL A. ..............124, 291

WEST, GEOFFREY B. ................. 45, 297
WEST, JASON B. ............................ 355
WESTBURY, HUGH M. .................... 355
WESTON, LESLIE A..........................21
WESTOVER, KRISTI M.................... 356
WESTRICK, LAWRENCE J. .............. 359
WETZEL, PAUL R........................... 356
WHEELER, BROOKE E. ................... 355
WHICKER, JEFFREY J..................... 169
WHIGHAM, DENNIS F. ..............155, 163
WHIPPLE, AMY ...............................89
WHIPPLE, STUART J....................... 262
WHITBECK, JULIE L.................218, 356
WHITE, CARLETON S. .................... 356
WHITE, DAVID L. .......................... 356
WHITE, ETHAN P. .......................... 357
WHITE, JENNIFER A....................... 357
WHITE, JOSEPH D....................229, 357
WHITE, LEE J.T.............................. 348
WHITE, LEE J. .................................14
WHITE, LUTHER............................ 367
WHITE, PETER S............... 167, 355, 357
WHITE, PETER .............................. 158
WHITE, SUSAN N........................... 357
WHITEHEAD, DAVID........... 42, 130, 336
WHITHAM, THOMAS G................... 360
WHITLOW, LINDSAY ...................... 358
WHITMIRE, STEFANIE L. ..................93
WHITTAKER, KARA ....................... 358
WHITTEN, W MARK....................... 308
WICKINGS, JEAN E. .........................14
WICKLAND, ALLISON C. ..................81
WIEDENHOEFT, JANE E. ...................11
WIEGERT, RICHARD G.................... 366
WILCOX, BRADFORD P................... 314
WILL, RODNEY E. ............................ 8
WILLARD, DEBRA A. ..................... 336
WILLIAMS, ALISSA J...................... 358
WILLIAMS, BOB J. ......................... 359
WILLIAMS, CHRISTOPHER K........... 358
WILLIAMS, CHRISTOPHER................98
WILLIAMS, DAVID G...................... 271
WILLIAMS, JACK W. ........................59
WILLIAMS, JOHN W. ...................... 359
WILLIAMS, KATHY S. .................... 359
WILLIAMS, KIMBERLYN ................ 359
WILLIAMS, LORI A. ....................... 135
WILLIAMS, NICHOLAS S G ............. 359
WILLIAMS, NORRIS H. ................... 308
WILLIG, MICHAEL R. .......................59
WILLIS, JONATHAN M.................... 359
WILLIS, THEODORE V. ................... 360
WILLSON, CYNTHIA J. ................... 360
WILSEY, BRIAN J. .......................... 269
WILSON, ALEXIS........................... 209
WILSON, GAIL W T........................ 192
WILSON, JACQUELINE A. ............... 360
WILSON, KAREN A. ....................... 360
WILSON, RICHARD ..........................78
WILSON, SCOTT D. ........................ 320
WILSON, WILLIAM G. ....... 249, 311, 372
WIMBERLY, MICHAEL C. ................ 360

WIMP, GINA M. ............................. 360

WINEMILLER, KIRK O. ................... 199
WINFREE, RACHAEL...................... 361
WINSOME, THAIS .......................... 361
WINSOR, JAMES A......................... 361
WINTER, KLAUS ........................... 308
WINTERS, JUSTIN.......................... 204
WINTERS, MONICA S. .................... 361
WIPF, SONJA ................................. 362
WISE, DAVID H. ............................ 203
WITHAM, THOMAS..........................89
WITHINGTON, CYNTHIA L.............. 362
WITMAN, JON D. .....................104, 362
WOFSY, STEVEN ........................... 160
WOJCIK, NATHAN J. ................296, 362
WOJDAK, JEREMY M. ............... 91, 362
WOLF, AMELIA A. ......................... 363
WOLF, AMY T................................ 157
WOLF, EVAN C. ............................. 363
WOLF, JOY J.................................. 363
WOLF, JULIE ................................. 363
WOLF, MOSHEH ............................ 363
WOLFE-BELLIN, KELLY S. ......... 25, 364
WOLOSIN, MICHAEL S. .................. 364
WONHAM, MARJOIRE J. ................. 364
WOOD, STANLEY R. ...................... 364
WOOD, TANA E. ............................ 365
WOOD, TANA .......................... 70, 158
WOOD, WAYNE R. ......................... 191
WOODBURY, PETER....................... 153
WOODS, JOEB ............................... 109
WOODS, KERRY D. ........................ 259
WOODSON, BROCK .........................51
WOODWARD, GUY ........................ 149
WOOLBRIGHT, SCOTT A. ................ 360
WOOLEY, STUART C. ..................... 365
WOOLFENDEN, GLEN .................... 178
WOOTEN, MICHAEL C.................... 239
WRAITH, JON M. ........................... 149
WRIGHT, ALLEN......................207, 365
WRIGHT, HERBERT E. .................... 249
WRIGHT, JESSICA W....................... 365
WRIGHT, JOSEPH........................... 184
WRIGHT, JUSTIN P.......................... 365
WRIGHT, MEREDITH S.................... 366
WRINN, KERRI .............................. 127
WRONA, AMANDA B. .................... 366
WU, JIANSHENG............................ 372
WURTSBAUGH, WAYNE ................. 254
WURZBURGER, NINA..................... 366
WYATT, JULIE L. ........................... 366
WYCKOFF, PETER H.........................41
WYDEVEN, ADRIAN P. .....................11
WYER, MARY B. ........................... 287
X
XI, WEIMIN .................................. 366
XIANG, BIXIA ............................... 334
XU, CHENGYUAN.......................... 367
XU, TAO.................................160, 367
Y
YAGER, LISA Y.............................. 367
YAKIMOVA, EVA.............................68

Author Index

395

YAKIR, DAN ............................ 41, 338

YAN, NORMAN D. ...........................16
YANAI, RUTH D. .....................212, 367
YANG, LOUIE H............................. 368
YANOSKY, THOMAS M. .................. 368
YANOVIAK, STEPHEN P. ................. 368
YARD, HELEN ............................... 176
YARNES, CHRISTOPHER T............... 368
YATES, TERRY L. ....................114, 290
YAUSSY, DANIEL A..................161, 277
YEAKLEY, J. A. ........................ 56, 368
YEATMAN, WILLIAM M.................. 100
YEE, DONALD A............................ 368
YEE, SUSAN M H .......................... 369
YELENA, DZYUBA V. ..................... 150
YERMAKOV, ZHANNA.................... 369
YLIOJA, TIINA ................................18
YODER, CARRIE L. ........................ 369
YOO, HO JUNG S. .......................... 369
YORK, ALAN C. ..............................51
YOSHIDA, LIDIA C...........................32
YOSHIDA, TAKEHITO..................... 370
YOUNG, LAUREN M....................... 370
YOUNG, WILLIAM P. ...................... 360

396

Author Index

YOUNGSTEADT, ELSA K................. 370

YU, FANGFANG............................. 335
YUND, PHIL O............................... 127
YURKONIS, KATHRYN A. ............... 370
Z
ZAFIRIS, ANGELIKIE A.............117, 370
ZAK, DON R.................................. 173
ZAK, DONALD R. ................ 65, 84, 371
ZAK, DON .................................... 192
ZAK, JOHN C. .................. 190, 192, 284
ZALIDIS, GEORGE ......................... 322
ZANGERL, ARTHUR R. ................... 137
ZASADA, JOHN ........................ 51, 301
ZASOSKI, ROBERT J....................... 144
ZEDLER, JOY ................................ 171
ZEDLER, PAUL H. ....................199, 278
ZEGLIN, LYDIA H. ......................... 371
ZEHRBACH, NANCY ................306, 347
ZELLMER, AMADA J. ..................... 371
ZELLMER, AMANDA J. ...................... 9
ZELMER, DEREK A. ....................... 171
ZERBACH, NANCY ........................ 169

ZETTLER, JENNIFER A. ............332, 371

ZHANG, CHI ................................. 371
ZHANG, DAOWEI .......................... 209
ZHANG, TAO ................................ 372
ZHANG, WEIJIAN .......................... 372
ZHANG, XINSHI ..............................63
ZHANG, YU .................................. 372
ZHAO, MAOSHENG........................ 174
ZHENG, LEI .................................. 372
ZHI, GUIYE.....................................63
ZHOU, JIZHONG ............................ 347
ZHOU, XIAOLU ............................. 266
ZHU, WEIXING.............................. 372
ZHUANG, QIANLAI ........................ 372
ZICUS, MICHAEL C. .........................80
ZIEGLER, SUSAN........................... 298
ZIMMERMANN, CRAIG R. ............... 373
ZIMPFER, JEFF F. ........................... 373
ZIPPERER, WAYNE C........................30
ZITZER, STEPHEN F........................ 373
ZOU, XIAOMING ........................... 373
ZU DOHNA, HEINRICH ................... 257
ZUREK, MONIKA.............................28
ZYGMUNT, JENNIFER R. ................. 373

USEFUL TELEPHONE NUMBERS and CONTACTS IN SAVANNAH

Savannah International Trade & Convention
Center

Savannah Area Convention and Visitors Bureau

(SAVCVB)

One International Drive, P.O. Box 248

Savannah, Georgia 31402-0248
Main Reception: (912) 447-4000; desk is staffed 9 AM to
5 PM weekdays

101 East Bay Street, Savannah, Georgia 31401

Main telephone number: (912) 644-6401
Email contacts for housing questions:
Diane Plunk at dplunk@savcvb.com or
Angie Cosenza at acosenza@savcvb.com

ESA/ISEM Annual Meeting Registration Desk

River Concourse, Savannah International Trade &
Convention Center
(912) 447-7300 (staff available during published hours)

Official Housing Locations:

Addresses, Telephone Numbers, and Shuttle Routes

Hotels
Hampton Inn in the Historic District
201 East Bay Street, Savannah

(912) 231-9700
Red Route

Savannah Marriott Riverfront

(912) 233-7722
100 General McIntosh Boulevard, Savannah
Red Route

In addition to serving as the Official Housing Bureau for

the ESA/ISEM Annual Meeting, the SAVCVB offers lots of
information about Savannah and wide variety of brochures
and maps. Offices are located less than a block from the
Hyatt Regency Savannah and Hampton Inn in the Historic
District.

Transportation Services
Airline Reservations and Car Rentals
Stellar Access/Conventions-in-America (Toll Free Number):
(800) 929-4242

Airport Services

The Mulberry Inn

601 East Bay Street, Savannah

(912) 238-1200
Red Route

B&B Shuttle Service: (912) 964-1411 or (912) 655-0441

Savannah International Airport: (912) 964-0541

Hyatt Regency Savannah

One West Bay Street, Savannah

(912) 238-1234
Blue Route

Local Taxi Services

Days Inn and Suites

201 West Bay Street, Savannah

(912) 236-1024
Blue Route

Radisson Hotel
411 West Bay Street, Savannah

(912) 790-7000
Green Route

Best Western Hotel

412 West Bay Street, Savannah

(912) 233-1011
Green Route

Comfort Inn and Suites

630 West Bay Street, Savannah

(912) 629-2001
Green Route

Savannah College of Art and Design (SCAD)

Dormitories
As individual rooms have no telephones in them, a general
SCAD contact number is provided.
Oglethorpe Residence Hall
201 West Oglethorpe Street, Savannah

(912) 525-7000
Orange Route

Turner House Residence Hall

302 West Boundary Street, Savannah

(912) 525-7000
Orange Route

Checker Yellow Cab: (912) 236-1133 or (912) 236-1951

Toucan Taxi Service: (912) 233-3700
Meeting Shuttle Service between official hotels and the
Savannah International Trade & Convention Center provided by Gray Line of Savannah (Cell phone contact): (912)
313-1501

Document Services
Kinko's Southside, 7929 Albercorn Extension, South
Savannah, Georgia, (912) 927-8119
Open 24 hours; 20% discount offered for attendees of the
ESA/ISEM Annual Meeting.

Child Care Services

Infant and Toddler Care and Youth Summer Camp
Parent and Child Development Services, Inc.
134 Houston Street, Savannah, Georgia 31401,
(912) 238-2778 or (912) 238-2777

Savannah Area Restaurants

Please refer to the information on Savannah restaurants
included with your meeting materials.