Ecological Indicators 6 (2006) 543553

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Utility and limitations of species richness metrics for

conservation planning
Erica Fleishman a,*, Reed F. Noss b, Barry R. Noon c
a

Center for Conservation Biology, Department of Biological Sciences, Stanford University, Stanford, CA 94305-5020, USA
b
University of Central Florida, Department of Biology, 4000 Central Florida Blvd., Orlando, FL 32816-2368, USA
c
Department of Fishery and Wildlife Biology, Colorado State University, Fort Collins, CO 80523-1474, USA
Received 22 April 2005; received in revised form 23 July 2005; accepted 25 July 2005

Abstract
The appropriateness of species richness as an ecological indicator or decision variable for setting conservation and management
priorities depends on many assumptions. Most critical is that current levels of species richness allow prediction of future
contributions of various locations to biodiversity conservation and ecological function. Also important is the assumption that
estimates of species richness can be compared among locations. Challenges arise because estimates of species richness are affected
by area, scale and intensity of sampling, taxonomic grouping, estimation methods, and the dynamic nature of species richness.
Nonetheless, species richness can contribute to prioritizing locations for biodiversity conservation provided it is not used in
isolationadditional metrics, such as species composition, endemism, functional significance, and the severity of threats, are also
required. The spatial domain of measurement also must be documented and justified. A multicriteria decision process is more likely
to realize comprehensive conservation goals than prioritization of locations based on species richness alone.
# 2005 Elsevier Ltd. All rights reserved.
Keywords: Biodiversity; Conservation planning; Hotspots; Scale dependence; Species richness

1. Introduction
Based on the number of publications reporting
estimates of species richness, it is clear that biologists
have a penchant for counting species. Counts are made
to understand why locations differ in the number of
species, what controls the number of species, and
especially why some locations are more species-rich
* Corresponding author. Tel.: +1 650 725 9914;
fax: +1 650 723 5920.
E-mail address: efleish@stanford.edu (E. Fleishman).

than others (Hutchinson, 1959). Biologists also hope to

understand how number of species (hereafter species
richness) is related to underlying environmental
variables such as latitude, elevation, and land cover,
and how species richness affects fundamental ecological processes such as primary productivity (e.g.,
MacArthur, 1972; Waide et al., 1999; Gaston, 2000;
Lyons and Willig, 2002; Pimm and Brown, 2004).
Among current principles used to guide conservation
and management is that protection of locations with
high species richness is an efficient way to conserve overall biodiversity and sustain key ecological

1470-160X/$ see front matter # 2005 Elsevier Ltd. All rights reserved.
doi:10.1016/j.ecolind.2005.07.005

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E. Fleishman et al. / Ecological Indicators 6 (2006) 543553

functions (Scott et al., 1987; Myers et al., 2000). In other

words, species richness is assumed to be an indicator of
conservation value (e.g., Meir et al., 2004).
Because of inherent interest in patterns and
processes associated with species richness, and because
maintenance of species-rich locations is believed to
protect multiple levels of biodiversity, ecologists have
put considerable effort into documenting species
richness and developing methods to identify potential
predictors of species richness. However, at any one
location, species richness is variable across time and
depends on dynamic processes. Therefore, protection of
locations that currently support a large number of
species may not be a reliable long-term conservation
strategy. Establishment of conservation reserves that
support high levels of biodiversity and associated
ecological processes over the long-term requires not
only information on species richness but also complementary measures of ecological integrity (Karr,
1981; Smogor and Angermeier, 2001; Davis and
Slobodkin, 2004) and resilience (Gunderson, 2000;
Allison, 2004).
In this paper, we first examine the utility and limitations of species richness metrics from a conceptual
perspective. We consider the extent to which protection of locations with a large number of species has
been achieved in practice, and whether this strategy
successfully has conserved biodiversity and ecological
function. Next, we provide an overview of existing
tools to make establishment of conservation priorities
based on species richness a more exact science. In
particular, we discuss methods for computing unbiased estimates of species richness to facilitate
reliable comparisons among locations and for factoring temporal variability in species richness into landuse decisions. Finally, we address several integrative
approaches for conservation assessment that build on
species richness to increase the probability of achieving multiple-criteria conservation objectives.

2. Origin and application of the species

richness paradigm
2.1. Motivation
For practical reasons, counts of species often are
emphasized throughout the process of ecological

assessment and monitoring. For many taxa of

conservation concern, species are discrete entities
that can be easily tallied. In addition, inexperienced
observers may not have sufficient skills to collect
unbiased data on abundance (Link and Sauer, 1998),
especially for taxa with considerable variation in
abundance within or between years. Using simple
models based on land cover to predict presence/
absence patterns (e.g., Scott et al., 1993, 2002; Caicco
et al., 1995) also is more feasible logistically than
collecting the detailed data on resource use or
demography necessary to model habitat quality or
viability. In addition, if sampling effort is standardized
among locations, it may be possible to make
inferences from count data about relative species
richness even if it is not possible to infer absolute
species richness (Link and Sauer, 1998).
A long history of counting species in ecosystems
worldwide has yielded a large amount of comparative
information. For example, available data clearly
illustrate global (chiefly latitudinal) gradients in
species richness, although the factors responsible
are still strongly debated (Gaston, 2000; Pimm and
Brown, 2004). Although some conclusions about
coarse-grained patterns of species richnesse.g.,
tropical latitudes harbor more species than polar
latitudesmay seem almost trivial, those conclusions
have motivated searches for mechanistic explanations
of species richness as a function of major gradients
including elevation (Janzen et al., 1976), primary
productivity or surrogates thereof (Hawkins et al.,
2003; Hurlbert and Haskell, 2003), and urbanization
(Pickett et al., 2001). The functional form of
relationships between species richness and environmental gradients often depends on the resolution and
extent of observation and on variation in disturbance
regimes (Pickett and White, 1985).
Existing information on species richness has
supported a range of high-profile conservation efforts,
particularly when data on rarity or endemism were
also available. For example, several major nongovernmental organizations have developed global maps of
the richness of endemic species in conjunction with
data on land use and land cover change. Areas with
high concentrations of endemics and rapid or
extensive land cover change commonly are known
as hotspots (Myers et al., 2000), whereas areas with
low species richness of endemics and more stable land

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cover patterns have been dubbed coldspots (Kareiva

and Marvier, 2003). Similarly, hotspots at subglobal
levels have been identified through application of rarityweighted indices of species richness (e.g., Stein et al.,
2000). The location of most hotspots based on these
criteria is not surprising to biologists. Nevertheless,
syntheses of comprehensive information on patterns of
endemism, imperilment, and projected land-use change
in repeatable ways have encouraged strategic conservation action based on species richness values.
2.2. Effectiveness
Hotspots of native and endemic species richness are
of obvious conservation value, but protection of
hotspots alone will not sustain global ecological
integrity. Intact ecosystems, such as much of the
boreal forest of Canada and Russia (Bryant et al.,
1997), are critical carbon sinks and reservoirs of littleknown biodiversity, yet they are not considered
hotspots and have received little conservation attention. For example, only 5.3% of the global land area
between 508 and 608 north latitude is protected,
compared to 9.4% of the tropics (UNEP-WCMC,
2002). Furthermore, we suggest that hotspots as they
usually are mapped often are too large (e.g., the
Caribbean; Myers et al., 2000) to guide land
acquisition or reserve design at realistic scales. Much
if not most land protection is driven by local
opportunities and politics rather than by a priori
assessment of ecological value (Prendergast et al.,
1999)indeed, the best-protected lands in many
regions (e.g., high-elevation areas) often have relatively low species richness (Harris, 1984; Noss and
Cooperrider, 1994; Scott et al., 2001). Specifying
large areas for conservation priority based on high
species richness ignores many relevant social factors.
Among these are patterns of land ownership and
whether economic incentives exist to encourage
private landowners to contribute to conservation
goals. Moreover, establishing a protected area on
paper does not imply the area will be managed in a
way that sustains biological diversity and ecological
processes (McCarthy et al., 2001; Pressey et al., 2003).
Whether protection of locations with high species
richness at the time of their designation has achieved
broad conservation goals remains an open question;
little data exists to assess whether this approach has

545

proven an efficient way to minimize extinctions.

Conservation organizations that purchase land with
the purported aim of maintaining or enhancing
biodiversity traditionally have measured success in
terms of tangible variables such as dollars donated or
hectares protected. That is, assessment of success has
not been measured in terms of the original objectives.
Increasingly, however, conservation organizations are
making an effort to measure success in terms of the
condition and trend of populations, species, and
ecosystems (Salafsky et al., 2002). The African
Wildlife Foundation, Conservation International,
The Nature Conservancy, the Wildlife Conservation
Society, and the World Wide Fund for Nature/World
Wildlife Fund recently launched an initiative, the
Conservation Measures Partnership, to develop a
common framework for evaluating conservation
success and the effectiveness of diverse conservation
strategies.
2.3. Information content
Biologists have emphasized measures of species
richness as a way to establish conservation priorities,
but they also have begun to acknowledge limitations
of those measures and to develop complementary
metrics. For example, methods have been formulated
to prioritize hotspots by incorporating explicit
information on the vulnerability of particular locations
to biodiversity loss, our ability to manage threats, and
whether locations will accommodate ecological and
evolutionary processes necessary to maintain species
richness over time (Pressey et al., 1996, 2003; Stein
et al., 2000; Noss et al., 2002). Although such methods
require additional information, they require little
supplementary information on species biology.
Proponents of the use of species richness to set
conservation priorities cite not only the comparative
ease of counting species or predicting species
occurrence, but also the conviction that information
on species richness provides insights to the state of
other ecological processes. However, the evidence that
species richness by itself provides insights to
ecosystem properties is equivocal. Inferences to
ecosystem properties require information on species
functional effects on ecological properties and their
functional responses to disturbance (Hooper et al.,
2005). Species richness per se does not distinguish

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between native and non-native species and conveys no

information on endemism, rarity, function, or sensitivity to changes in land use. In addition, species
richness provides no information on density or
demography and thus provides no insights to likelihoods of species persistence. Despite its shortcomings as an indicator of ecological condition, we
believe species richness will remain a popular metric
for comparing conservation alternatives if for no other
reason than its wide availability. Given this reality, it
is important to ask how species richness can be
estimated more accurately so as to allow more reliable
comparisons among candidate conservation areas.

3. Estimation of species richness

Observed differences in species richness among
locations may reflect true differences in species
richness as well as differences in sampling effort or
dissimilarity in the underlying distributions of
species abundance (Wintle et al., 2004). As
ornithologists have long recognized (e.g., Emlen,
1971), counts must be adjusted for variation in
probabilities of detection among species (Boulinier
et al., 1998; Gotelli and Colwell, 2001; Cam et al.,
2002; Royle and Nichols, 2003). Counts of species
are challenging for many taxonomic groups, geographic locations, and ecosystems. Microbes, invertebrates, and many marine and tropical assemblages
are among the more difficult to evaluate accurately
and efficiently (Walters, 1986). Field measurements
tend to underestimate species richness (Magurran,
1988; Soberon and Llorente, 1993; Hayek and
Buzas, 1997), and, as we address in detail later in
this paper, short-term snapshots of species richness
and composition may not accurately reflect longer
term patterns in response to deterministic or
stochastic environmental changes (Conroy and
Noon, 1996; Tyre et al., 2001).
Standardizing for survey effort is essential before
comparisons are made. However, it is not a trivial
exercise because the two ways of standardizing
species richness estimatesaccording to the area or
number of individuals sampledmay lead to different
estimates of species richness and, in turn, different
priority rankings (Gotelli and Colwell, 2001). Legitimate arguments can be made for either method of

standardization. The area-based approach, which

accumulates area by the addition of successive sample
units, better accounts for heterogeneity in species
richness among samples, but is affected by differences
in the mean number of individuals per sample unit.
The individual approach may require measurements
from sample units of substantially different size.
Importantly, if locations being compared differ in the
mean number of individuals per sample, then their
rank order of species richness may depend on how
estimates of species richness are standardized (Gotelli
and Colwell, 2001). Differences can be explored by
comparing area versus individual-based rarefaction
curves separately for each location. Rarefaction
curves are generated by repeatedly resampling from
the pool of sample units or individuals and plotting
species richness as a function of sample size
(Simberloff, 1972). Use of rarefaction to standardize
survey data (whether by total area or number of
individuals) prior to comparison among locations,
however, must be done carefully because the method
assumes, among other things, that distributions of
species abundance in the locations being compared are
similar (Gotelli and Graves, 1996).
For conservation decision-making, it may be more
appropriate to standardize species richness estimates
by area sampled than by number of individuals
because protected areas have fixed dimensions and
specific costs per unit area. Nevertheless, if data are
standardized to a common area, it is possible to infer
that a relatively species-poor sample has higher
species richness than if the same data were
standardized to number of individuals. When
estimates are standardized by area, the statistic
estimated is more properly called species density
than species richness because it reflects the number of
species within a specified area (Gotelli and Colwell,
2001).
Equally relevant to the standardization of richness
estimates prior to making comparisons among
locations is the adequacy of the sample. If the area
being surveyed contains a large number of rare
species, then it is likely that many species will not be
included in the sample. Thus, methods used to
estimate species richness based on interpolation from
the sample data (i.e., species accumulation or
rarefaction curves) will be biased low (Boulinier
et al., 1998). In general, estimates of species richness

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when detection probabilities are both <1.0 and

heterogeneous among species will be negatively
biased (Boulinier et al., 1998; Cam et al., 2002).
Sampling rarely continues until all species have
been enumerated. As a result, extrapolation methods
are needed to estimate the number of undetected
species. Recently, many methods have been developed that allow extrapolation from survey data to
estimate the true number of species (e.g., Keating
et al., 1998; Nichols et al., 1998). Nonparametric
estimators based on mark-recapture data seem most
reliable because they estimate species richness by
accounting for the variation in species detection
probabilities that exists in real surveys. These
methods are applicable to any taxonomic group,
and do not require organisms to be marked. Instead,
they are based on records of species occurrence
(presence or absence) obtained during repeated visits
to sample units. These nonparametric estimates of
species richness are unbiased and independent of the
sampling effort devoted to a location of a given size
(Cam et al., 2002).
It is well known that measures of species richness
and diversity, and inferences drawn from these
patterns, are dependent on the spatial and temporal
scale of measurement (Noss, 1983; Wilson and
Shmida, 1984; Conroy and Noon, 1996). Sometimes
the effects of scale are subtle and go unnoticed. For
example, extent can have two different interpretations:
(1) the total area sampled based on the cumulative area
pooled across sample units (i.e., from species
accumulation curves or rarefaction curves) or (2)
the area spanned by the sample units, which increases
as the distance between sample units increases. Using
sample-based accumulation methods to estimate
species richness fails to distinguish between these
two interpretations. Even if controlled for area
sampled, two locations that have been sampled may
differ in spatial extent, and thus species richness,
because one set of samples spans a greater area
(Whittaker et al., 2001). In addition, for management
purposes it is necessary to recognize that manipulations of land cover or ecological processes that
increase species richness at the local level, for
example through increased heterogeneity of vegetation, edge effects, and changes in disturbance regimes,
may reduce species richness at larger spatial levels if
opportunistic species prosper at the expense of

547

sensitive species (Noss, 1983; Shea et al., 2002;

Pressey et al., 2003).

4. The dynamic nature of species richness

Using comparative values of species richness as a
criterion for establishing land-use priorities is prone to
error unless the changing nature of species richness
and the dynamic processes that give rise to it are taken
into account (Conroy and Noon, 1996; Tyre et al.,
2001; Bestelmeyer et al., 2003). Island biogeography
theory (MacArthur and Wilson, 1967) is the historical
cornerstone of many strategies intended to maximize
conservation of biodiversity. Often forgotten, however, is that the MacArthurWilson model is explicitly
dynamic in both space and time. At any given location,
the current value of species richness reflects species
richness in a previous time period, augmented by the
number of species that have colonized the location
since then and diminished by the number of species
that have gone locally extinct. Thus, a change in
species richness over time is a function of the
difference between local colonization and extinction
rates, both of which are functions of the current value
of species richness. Colonization rate decreases and
extinction rate increases as the number of species
approaches that available in the regional species pool.
Many conservation and management strategies
implicitly assume that species richness is constant
i.e., that the rates of colonization and extinction are
constant. However, constant rates of colonization and
extinction do not imply the absence of colonization
and extinctioneven with constant rates, ecological
processes are not static and there is turnover in species
composition. Further, temporal variability in colonization and extinction rates introduces an additional
source of variability both in the value of species
richness and in species composition.
In the absence of reliable estimates of colonization
and extinction rates, general knowledge about a system
can be used to factor dynamic attributes of species
richness into conceptual models for ecological assessment and management (Conroy and Noon, 1996). For
example, are the species most likely to colonize or be
extirpated a random subset of the species in the regional
pool, or is the order of colonization and extinction
predictable based on differences in their ecology and

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life histories? If the order of colonization and extinction

is predictable, are species likely to colonize or go
extinct those of greatest conservation concern? How
does the colonization process affect the extinction
processthat is, do the species most likely to colonize
disproportionately affect the persistence of the species
most in need of conservation? Based on answers to
these questions, is there a predictable rate and trajectory
of change in species richness (and species identity) that
would undermine the utility of species richness as an
ecological indicator or conservation criterion? Ultimately, active and flexible management plans that
address colonization and extinction processes are more
likely to succeed than static land-use delineations
(Bestelmeyer et al., 2003; Shea et al., 2002; Pyke,
2004).

5. Complementary approaches
The value of estimates of species richness for
setting conservation and management priorities is
enhanced when those data are linked with more
revealing statistics associated with species composition, species functional roles, or the ability of a
location to sustain ecological processes associated
with species richness and viability (Burgman et al.,
2001). Among the species-based approaches to
complement or substitute for direct measures of
species richness are conservation of species with
particular attributes based on their area requirements,
trophic position, functional uniqueness, or irreplaceability. Surrogate-based approaches include coarsefilter methods based on vegetation composition and
seral stage. These approaches are popular with
conservation biologists and land managers, but
inadequate attention to their conceptual basis and
appropriate use sometimes has led to inconsistent and
questionable applications. We do not provide an
exhaustive review of complementary approaches. For
example, we do not explore estimation of population
viability, which can play a useful role in conservation
assessment and land-use planning (for recent reviews,
see Sjogren-Gulve and Ebenhard, 2000; Beissinger
and McCullough, 2002). Nor do we explore fully
emerging methods to incorporate ecological processes
into identification of conservation and management
targets (e.g., Gerber et al., 2003; Pressey et al., 2003).

5.1. Species status

The potential success of conservation planning can
be improved dramatically when data on species
richness can be coupled explicitly with information
on species composition and knowledge of the basic
life history of taxonomic groups. Diverse assessment
approaches have been designed based on the argument
that species composition reflects land use and
associated land cover change. Examples include the
Floristic Quality Index (Nichols, 1999; Rooney and
Rogers, 2002), indices of biotic integrity (Karr, 1981;
Fausch et al., 1984; Kerans and Karr, 1994; OConnell
et al., 1998; Hill et al., 2000; Smogor and Angermeier,
2001), and ratios of edge species to interior species.
Three especially useful metrics that can be derived
from species lists and rudimentary life history
information are proportion of native species, proportion of species characteristic of undisturbed locations
(as opposed to ubiquitous or weedy species
characteristic of biotic homogenisation; Noss, 1990;
Lockwood and McKinney, 2001), and proportion of
species with narrow geographic distributions (Noss,
1983). Further analysis may be used to classify species
as avoiders, adapters, or exploiters of specific natural
and human disturbances (McKinney, 2002).
Studies of the biological effects of urbanization
illustrate the value of combining estimates of species
richness with information on species composition, life
history, and geographic distribution. Although urbanization is one of the primary threats to species
viability (Czech et al., 2000), species richness often
peaks in an intermediate location along an urbanization gradient. For example, golf courses may have
more species of birds and butterflies than either nature
preserves or business districts (Blair, 1999). However,
a high proportion of the species found in locations
with intermediate levels of urbanization are relatively
ubiquitous. By contrast, many native species are
restricted to locations with comparatively little urban
development despite the fact that species richness of
those locations is relatively low (McKinney, 2002).
Information on ecology and life history also
facilitates identification of species that play key roles
in sustaining ecological processes and community
structure (Kinzig et al., 2001). Asking what species do
in ecosystems (Lawton, 1994) is the relevant question
for setting conservation priorities. Species with high

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conservation value include those that contribute

disproportionately to the transfer of matter and energy
(sometimes called keystone species), structure the
environment and create opportunities for additional
species (ecological engineers), or exercise control
over competitive dominants, thereby promoting
increased biotic diversity (strong interactors) (Mills
et al., 1993; Soule et al., 2003, 2005). Knowledge of
the distribution patterns of species that play significant
functional roles, in conjunction with information on
species richness, encourages a balance between
species-centric and ecosystem-focused conservation
strategies and greatly improves the capacity for
conservation at multiple levels of biological organization (Kinzig et al., 2001).
5.2. Species-specific management
Data on the occurrence of species of concern
complement data on species richness, and are relevant
for delineation of conservation areas and other land
uses. Species of concern encompass taxa that are
targeted for either conservation or eradication. Socioeconomic as well as ecological criteria affect the value
attributed to individual species. Ecologists typically
assign high conservation priority to species that are
native, rare or threatened, phylogenetically distinct, or
play important roles in maintaining ecological
function (Noss and Cooperrider, 1994) and low
priority to cosmopolitan or introduced species. But,
as attitudes towards the conservation of top predators
demonstrate, ecological valuations of the complement
of species to conserve or eliminate often conflict with
other perspectives. Despite considerable scientific
evidence for the importance of predators in promoting
species richness and stabilizing food webs (Noss et al.,
1996; Soule et al., 2003, 2005; Ripple and Beschta,
2005), many sectors effectively cull native predators
to benefit game species (some of which are
introduced) or because of social values.
Caution is often necessary when using speciesbased approaches because methods for protecting or
controlling species of concern may not meet the needs
of the larger biotic community to which they belong.
For example, rarity can be driven by a complex array
of mechanisms, and rare species often do not occur in
locations with highest species richness (Cody, 1986).
Moreover, the distributions of different rare species

549

may not overlap (Freitag et al., 1997), making it

difficult to achieve comprehensive conservation goals
in a small number of reserves. As we learn more about
the functional traits of species some general rules may
be emerging. Conservation focused on highly interactive species (Soule et al., 2005)often those at the
topic of trophic pyramidsor species with large area
requirements (e.g., Roberge and Angelstam, 2004)
may provide some assurance that the resource
requirements of unmanaged species also may be
met. However, these rules should not be applied
uncritically.
In some situations, well intentioned species-based
approaches may conflict with maintenance of native
biodiversity. For instance, along riparian corridors in
arid ecosystems around the world, invasive species are
modifying ecosystem processes, species distributions,
and population dynamics of native species (Higgins
et al., 1999; Mooney and Hobbs, 2000; Rejmanek,
2000; Palumbi, 2001). Unless native riparian vegetation can be restored immediately following elimination of non-native species, however, rehabilitation
efforts inadvertently may threaten native biota
(Zavaleta et al., 2001). For example, non-native
willows (Salix spp.) once were planted extensively to
arrest erosion in streams and rivers in Australia.
Restoration ecologists have concluded that the
willows should not be eradicated without a concomitant plan to reestablish native riparian vegetation
(Cremer, 1999; Williams and West, 2000). Similarly,
efforts to eradicate introduced salt-cedar (Tamarix
ramosissima) along rivers in the Mojave Desert have
been confounded by the ability of many native
animals, including the endangered Southwestern
Willow Flycatcher (Empidonax traillii extimus), to
exploit areas that now are dominated by T. ramossissima (Cleverly et al., 1997; Sher et al., 2000).
5.3. Coarse filters
Coarse-filter strategies use metrics on the distribution of communities, ecosystems, or other environmental classes as surrogates for biodiversity. Although
the coarse-filter approach typically recognizes communities and ecosystems as worthy of protection in
their own right, it is often used to infer species richness
rather than directly enumerating or assessing the status
of individual species (Groves, 2003; Lombard et al.,

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2003; Noon et al., 2003). Coarse-filter approaches

assume most species in a regional biota will be
conserved if representative land-cover types or
environmental domains (e.g., as determined by
climate and geology) are protected. Although a
coarse-filter approach alone rarely is adequate to
protect all native plant and animal species in the long
term, the method complements the so-called fine filter
of species-level conservation (Hunter et al., 1988;
Noss and Cooperrider, 1994; Noon and Dale, 2002).
The coarse-filter framework has been widely
applied by merging thematic layers of relatively
low resolution land-cover data, such as vegetation and
topography, in a GIS platform. Land-cover data are
linked with wildlife-habitat relationship models (e.g.,
Mayer and Laudenslayer, 1988) to infer the distribution of animal species across the landscape. The
number of species predicted to occur in a mapping unit
serves as an estimate of species richness at that
location. The broader purpose of these exercises is to
identify gaps in the distribution of land-cover types
and associated species in networks of protected areas
(Scott et al., 1993).
In principle, coarse-filter approaches are a costefficient, indirect method of assessing species
distributions and may facilitate interim protection of
taxa for which distributional data are limited. The
extent to which coarse-filter approaches are effective
in practice depends on the extent to which three
assumptions are valid: (1) attributes used to delineate
land-cover types are reliable surrogates for habitat
(and the occurrence of a species is strongly correlated
with the occurrence of its habitat), (2) management
actions at the ecosystem level will address the factors
currently limiting the distribution and abundance of
species, and (3) the spatial resolution of the coarse
filter matches the scale at which species respond to
environmental heterogeneity.
These three assumptions may hold for some
species, especially those that are small-bodied,
abundant, and tightly linked to a particular type of
land cover. However, these assumptions rarely are met
for all, or even most, species in an assemblage; coarsefilter assessments often overestimate the presence of a
species on the planning landscape. Thus, the success
of the coarse-filter approach has proven equivocal
(Boone and Krohn, 2000), albeit it is practically
impossible to test the coarse filter comprehensively,

given limited distributional data for many taxa. Errors

of commission (false positives) and omission (false
negatives) can be reduced simultaneously by increasing the resolution of the filter. The latter adjustment
reduces the area predicted to be suitable for each
species (Karl et al., 2000). In particular, the accuracy
of occurrence predictions for species that are rare,
have specialized resource requirements, or have large
home ranges usually can be increased by increasing
the number of mapped land cover types (Karl et al.,
2000).

6. Conclusions
Ecologists and conservation biologists have
focused on species richness, its geographic patterns,
and its environmental correlates for many decades.
Species richness has been a convenient indicator of
ecological condition or conservation value because it
has been measured in many places and over long
periods of time. Nevertheless, a preoccupation with
species richness may be more strongly related to data
availability and the desire to generate lists of species
than to the search for the ecological significance of the
statistic.
Certainly, reliable information on species richness,
when referenced to a specific location, is of value. Yet
species richness by itself is an insufficient measure to
characterize candidate conservation areas in terms of
their ability to sustain biodiversity and ecological
function over the long term. The primary reason is that
species richness, in isolation, provides no information
on species identity or the functional roles of individual
species as contributors to ecosystem processes and
resilience to stress. A simple estimate of species
richness provides few, if any, insights to the dynamic
processes of extinction and colonization that give rise
to species richness in the first place. In addition, most
measures of species richness are not true estimates in a
statistical sense because it is impossible to determine
their precision or the influence of possible biases. The
absence of standardized, statistically credible estimates of species richness makes suspect the use of this
statistic for assigning conservation priorities.
Nevertheless, we do not intend to dismiss a concept
that has some merit. Species richness remains a
valuable statisticwhen it is used in combination with

E. Fleishman et al. / Ecological Indicators 6 (2006) 543553

information on, for example, composition, function,

endemism, and the distribution of abundance. We also
should ask whether the complement of species implied
in any value of species richness represents one or more
entire food webs with resiliency to environmental
variation, or whether it lacks top predators or diverse
primary producers. Establishment of conservation strategies and priorities requires comprehensive suites of
complementary measures of ecological communities.
Using species richness as the principal criterion for
establishing priorities in the face of the biodiversity
crisis is convenient but not defensible. Given the value
of biodiversity to human welfare, conservation
planning must be scientifically rigorous and comprehensive. Continuing to rely exclusively on simple
metrics such as species richness, which are known to
be information-poor and subject to estimation errors,
is unacceptable. The tools exist to make conservation
planning a more exact science by implementing more
scrupulous standards for sampling ecological communities, estimating parameters, and using the
information derived to develop conservation priorities
that reflect the multiple values of natural ecosystems.

Acknowledgment
EF acknowledges the support of the Nevada
Biodiversity Research and Conservation Initiative
during preparation of this manuscript.

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