Limnol. Oceanogr.

, 55(6), 2010, 22752284

2010, by the American Society of Limnology and Oceanography, Inc.
doi:10.4319/lo.2010.55.6.2275

On the role of natural water level fluctuation in structuring littoral benthic

macroinvertebrate community composition in lakes
Michael S. White,a,1,* Marguerite A. Xenopoulos,b Robert A. Metcalfe,c and Keith M. Somersd
a Watershed

Ecosystems Graduate Program, Trent University, Peterborough, Ontario

of Biology, Trent University, Peterborough, Ontario
c Renewable Energy Section, Ministry of Natural Resources, Trent University, Peterborough, Ontario
d Dorset Environmental Science Centre, Ontario Ministry of the Environment, Dorset, Ontario
b Department

Abstract
We used traditional hydrologic endpoints and extracted novel water-level fluctuation (WLF) characteristics
through principal components analysis (PCA), to determine the relationship between WLF and rocky littoral
benthic macroinvertebrate (BMI) richness in 16 boreal lakes. Yearly WLF amplitude (maximum minus minimum
water levels) ranged from 35.9 cm to 157.5 cm. Using PCA we derived a new variable D80-D210 (31 March minus
01 August) as a surrogate for change in mean water level and potential habitat squeeze (loss). Analyses of BMI
richness with several physicochemical variables, including water quality (8), habitat variability (4), lake and basin
morphology (6), land classification (28), water temperature (8), hydrology (9), and PCA axes (10) resulted in only
three significant relationships. We found a classic speciesarea relationship, as BMI richness increases with
increasing lake area (r2 5 0.38linear, r2 5 0.69unimodal). Similarly, as littoral slope increases macroinvertebrate
richness decreases (r2 5 0.32linear). Most importantly, lower water levels, quantified using D80-D210, have higher
macroinvertebrate richness (r2 5 0.38linear). Together these results suggest that a habitat squeeze in littoral areas is
the direct result of lower mean water levels and that relatively small changes in natural WLF can be associated
with changes in BMI communities.

The response of organisms to changes in their environment

(disturbance ecology) is a well-studied phenomenon in both
terrestrial and aquatic biomes (Mackey and Currie 2001). In
order to better understand the effects of anthropogenic
alterations and disturbance on biotic communities, researchers must first recognize the role of natural variability. This is
important so that unnatural disturbances can be assessed
against naturally occurring levels of variability. Lack of a
thorough understanding of natural conditions limits the
strength of conclusions drawn from studies of effects induced
by humans (Bowman and Somers 2005).
One area in freshwater aquatic ecology where there is
limited understanding of the natural variability and ecological
effects is the role of natural water-level fluctuation (WLF),
particularly, in lake ecosystems (Wantzen et al. 2008; White et
al. 2008). Considerable research has focused on studying
unnatural or manmade changes in water levels through
studies of systems regulated by humans (e.g., hydroelectric
reservoirs (Hill et al. 1998; Aroviita and Hamalainen 2008);
however, we have a very poor understanding of natural WLF
and its affect on aquatic communities.
In reservoirs and other manmade aquatic systems, WLF
decreases macrophyte diversity (Hill et al. 1998), macroinvertebrate diversity (Aroviita and Hamalainen 2008), fish
hl 2005), and waterfowl recruitrecruitment (Fisher and O
ment (Sayler and Willms 1997). Regulated WLF in
reservoirs often results in altered timing of flood events
* Corresponding author: mwhite@laurentian.ca
1 Present

and extremely low winter water levels (Aroviita and

Hamalainen 2008) compared with natural systems. Therefore, the usefulness of this WLF information from
reservoirs for making predictions on how changes in
natural water levels will affect biological communities is
questionable. In addition, most reservoir studies focus on
only one WLF measure: amplitude (maximum minus
minimum yearly water level). Although this basic metric
often produces significant results, effects are only found at
levels that exceed those of natural lakes (White et al. 2008).
While studies concerning the effects of WLF on aquatic
biota in hydroelectric reservoirs help us better understand
the effect of water regulation, these studies have a limited
capacity to explain the role of natural WLF in structuring
aquatic communities. Furthermore, climate scenarios
predict more subtle changes in natural lake WLF (Magnuson et al. 1997) than fluctuations observed in reservoirs. As
a result, to understand the potential effects of climate
change, a thorough understanding of natural WLF and its
ecological relationships is necessary. The objective of this
study was to evaluate natural WLF character, to identify its
predictors, and to identify components of natural WLF
that correlate with nearshore stony littoral benthic macroinvertebrate (BMI) communities. We compared annual
WLF and BMI communities in 16 relatively large Boreal
Shield lakes. We hypothesized that landscape and climatic
conditions influence WLF character which, in turn,
structures littoral BMI communities.

Methods

address: Cooperative Freshwater Ecology Unit,

Laurentian University, Department of Biology, Sudbury, Ontario,
Canada.

Study siteSixteen lakes located in the Boreal Shield

ecoregion of Ontario, Canada were used in this study

2275

2276

White et al.

Fig. 1. Map of Ontario depicting locations of the 16

study lakes.

(Fig. 1). This geographic area covers approximately

300,000 km2, ensuring that study lakes experience a range
of WLF due to climatic variability and geographic location.
The dominant forest cover is black spruce (Picea mariana),
white spruce (Picea glauca), white pine (Pinus strobes), jack
pine (Pinus banksiana), poplar (Populus spp.), and birch
(Betula spp.). Further, forests near the Great Lakes often
include hardwoods such as maple (Acer spp.) and ash
(Fraxinus spp.). In general, this region has a temperate
climate, receiving 600800 mm annual precipitation in the
northwest (mean annual low temperature between 245uC
and 240uC, mean annual high temperature between 26uC
and 38uC) and 8001200 mm annual precipitation in the
northeast (mean annual low temperature between 240uC
and 234uC, mean annual high temperature between 26uC
and 38uC) (Natural Resources Canada 2004). The study
lakes are part of the Ontario Water Level Lake Series
(OWLLS) project; primarily funded by the Ontario
Ministry of Natural Resources (OMNR). The focus of
OWLLS is to monitor and study water levels in lakes across
the province. Lake selection was based on the following
criteria: surface area . 10 km2, mean depth , 10 m, no
dam control structure upstream or immediately downstream, accessible by road, and geographically dispersed
across the province.
Sampling protocolAll 16 lakes were outfitted with
water-level loggers in October 2005. Both SolinstH model
2001 and HOBOH model U20-001-01 (Onset) loggers were
employed, each logger was surveyed to a reference pin and
set to record temperature and water level at hourly intervals
365 d/yr. Biotic and abiotic sampling was conducted in late
summer of 2006 during a 4-week period (14 August to 11
September), which limits temporal variation (Reid et al.
1995). Each lake was sampled at one location only. At each

site the following water-quality measurements were taken

using a YSI 600QS SondeH: conductivity, pH, oxidation
reduction potential (ORP), water temperature, and dissolved oxygen (DO). A single 1-liter water sample was
collected for total phosphorus (TP), total dissolved
nitrogen (TDN), and dissolved organic carbon (DOC). A
500-ml subsample was filtered using a polycarbonate,
0.2 mm-pore, 47-mm filter (GE Water and Process
Technologies) within 4 h of sample collection for TDN
and DOC. Immediate processing of the water samples was
not feasible; therefore, all water samples were stored at 4uC
in the dark for 4 weeks until analysis in the lab was
possible.
Rocky littoral benthic macroinvertebrates were sampled
in triplicate at each sampling location following the Ontario
Ministry of the Environments (OMOE) lake benthos
sampling protocol (Jones et al. 2007). Each of three transects
were perpendicular to the shore and were kick sampled
starting at 1 m depth and proceeding back to shore until the
waters edge was reached. Kick sampling was standardized
by sampling for 1 min using a 500-mm mesh kick net. Total
transect length was recorded along with GPS coordinates
and multiple habitat descriptors (% canopy coverage, three
cobble size classes 6.525 cm, % woody debris, riparian
vegetation type, and three attached algae classes) (Jones et
al. 2007). Each kick-and-sweep transect was located 5 m
apart. To reduce habitat variability, sampling locations were
constrained to the following criteria: stony littoral substrate
of 6.525 cm in length, west-facing shoreline, with fetch
. 1 km. In situ measurements were taken for the abovementioned parameters to account for any unexpected
influence that may be attributed to the site. Benthic
macroinvertebrate samples were preserved using 200 mL of
Kahles solution (Martin 1977) and stored in 500-mL
polyethylene jars. In the lab, the BMI samples were sieved
to 500 mm and all macroinvertebrates were removed using
standard quality assurance and quality control (Jones et al.
2007) procedures at 10X magnification. Due to the large
geographic extent of this study, heterogeneity between sites
was reduced by identifying BMI to the family level of
taxonomic resolution (Bailey et al. 2004). Water mites were
the only taxon not identified to family, and all mites were
identified collectively as the Hydracarina. All BMI were
identified using either the work of Merritt et al. (2008) or of
Peckarski et al. (1990).
Landscape analysisLand classification (28 land classifications) and delineation was determined using Ontarios
Natural Resources and Values Information System
(NRVIS) database (OMNR 2002) and the Water Resources
and Information Project (WRIP) (OMNR 2006). These
data were analyzed using ArcMapH (ESRI) to determine
lake and watershed boundaries, watershed slope, and land
use. Lake morphology was extracted from the Ontario
Ministry of Natural Resources (OMNR) lake database for
the following three variables: maximum depth, mean depth,
and shoreline development (shape).
Hydrological data analysisHydrologic parameters were
calculated from water-level readings spanning the period

Water level and benthos

from 01 March 2006 until 30 November 2006. However, of
the calculated hydrologic parameters, it is expected that
only those occurring before BMI collection (14 August
2006 to 11 September 2006) will demonstrate meaningful
relationships with BMI community structure. Standardization to a common WLF was accomplished by adjusting the
water level for each lake to identical starting heights at 01
March 2006. This standardization date was selected due to
the fact that this date occurred at least a week before any of
the 16 study lakes had started to exhibit a spring melt
hydrologic signature. Variation in hourly logger readings
due to instrument sensitivity required that the hourly data
be smoothed by taking a 12-h running average. Mean
daily values were then calculated and used in calculating
hydrologic endpoints. The following metrics were calculated from the water temperature data: mean temperature,
minimum temperature, maximum temperature, coefficient
of variation, and day of year at which a temperature of 4uC,
10uC, 15uC, and 20uC was first attained. The following
hydrologic endpoints were calculated: water-level coefficient of variation, amplitude (maximum minus minimum
water level), melt day (first day when water level exceeds
water level during winter months), fastest 15-d flood rate
(rate of water-level rise during spring), flood rate (melt day
to day of maximum water level), recessional limb rate (day
of maximum water level to 01 September), day of year
when maximum water level was attained (MWLD), waterlevel rise duration (WLRD, calculated as the number of
days between melt day and MWLD), and mean annual
runoff. Mean annual runoff for each watershed was
estimated using Canada Gridded Climate Data 19611990
(Hopkinson 2000) with ArcMapH by subtracting raster
grids of total annual potential evapotranspiration from
total annual precipitation, assuming no change in storage.
Further, a novel hydrologic endpoint (D80-D210) was
calculated by subtracting the water level at day 80 from the
water level at day 210. Water level at day 80 is a surrogate
for the winter water level, and water level at day 210 is
shown in the results to be predominantly important in the
PCA extraction (see below) and precedes BMI sampling
(day 226254). Thus, D80-D210 is a measure of the change
in water level from ice-covered winter months to midsummer, representing an index of the change in habitat
availability.
Statistical analysisA principal components analysis
(PCA) was used to summarize the water level of the 16
lakes using Personal Computer Ordination (PC-ORD)
multivariate analysis of ecological data (McCune and
Mefford 1999). The resulting ordination was then overlaid
with a second matrix containing landscape, lake morphology, water quality, BMI richness, and commonly measured
hydrologic endpoints to determine correlations with the
extracted axes.
We used regression analysis (SPSS 2000) to evaluate the
relationships among extracted principal components, commonly measured hydrological variables, water temperature,
water quality, lake morphology, basin morphology, habitat
characteristics, and land classification with BMI richness.
Where necessary, data was transformed to satisfy normality

2277

Table 1. Limnological properties of the 16 study lakes. Water

quality was sampled from 14 August to 11 September 2006.
Lake properties (n516)
(km2)

Mean 6 SD

Lake area
31.52620.85
507.986571.71
Basin area (km2)
Basin : lake area ratio
17.29614.29
Elevation (m)
364.19657.15
Max depth (m)
33.7467.33
Mean depth (m)
8.5361.17
Shoreline development (shape)
7.3662.69
Littoral slope (u)
11.5864.16
Fetch (m)
2667611146
DOC (mg L21)
7.3961.18
TDN (mg L21)
0.2160.04
TP (mg L21)
14.7565.20
DO (mg L21)
8.6860.40
Conductivity (mS cm21)
69.75646.34
pH
7.6460.36
ORP
85.15631.02

Max

Min

67.85
2355.15
56.27
437.00
48.80
10.30
12.19
21.80
5360
9.56
0.32
23.71
9.30
167.00
8.47
141.10

8.14
87.51
3.87
252.00
16.20
6.10
3.05
6.30
971
4.90
0.16
4.72
7.92
23.00
7.18
17.10

assumptions. We also conducted stepwise multiple regressions using the same variables employed in regression
analyses to determine which combination of variables best
explained BMI richness (SAS 2001). Where necessary, a
test of collinearity was conducted as described in Xenopoulos et al. (2003) using SAS (SAS Institute 2001). The
collinearities that resulted were all low and did not affect
model outputs.
To identify taxa that are most affected by WLF, an
indicator taxa analysis was conducted following the
method of Dufrene and Legendre (1997) using PC-ORD
(McCune and Mefford 1999). This method combines
relative abundance and relative frequency of occurrence
to identify indicator taxa.

Fig. 2. Hydrograph showing mean water level (solid line) of

the 16 study lakes 6 1 standard deviation (dashed line). Water
levels were recorded at hourly intervals and were smoothed using
a 12-h running average. All analyses were conducting using
average daily values.

2278

White et al.

Table 2. Eigenvalues for the first 10 axes extracted through

PCA of WLF data from 01 March 2006 until 30 November 2006.
Ordination of axis 1 and axis 2 presented in Fig. 3a.
Axis

Eigenvalue

% variance explained

1
2
3
4
5
6
7
8
9
10

102.66
85.01
36.10
11.86
5.27
2.96
1.58
1.48
1.13
0.52

41.06
34.00
14.44
4.75
2.11
1.18
0.63
0.59
0.45
0.21

For the indicator-taxa analysis, lakes were categorized

into three groups based on the magnitude of change in
water level: the group with the highest change had D80D210 values between +8.42 and 23.30 cm, n 5 5; the group
with mid-range change had D80-D210 values between
23.30 cm and 28.90 cm, n 5 5; and the group with the
lowest change had D80-D210 values between 28.90 cm and
224.20 cm, n 5 6. Significance of a taxon belonging to
either the high, mid, or low group was determined using
Monte Carlo randomization procedure using 1000 permutations (Dufrene and Legendre 1997).

Results
Lake morphology and water quality varied among lakes
(Table 1). Similarly, amplitude ranged between 157.50 cm
and 35.97 cm, with a mean and standard deviation of
73.29 cm and 34.90, respectively (Fig. 2). Melt day varied
from 11 March to 30 March 2006. The fastest 15-d flood
rate ranged from 0.90 cm d21 to 7.83 cm d21, with mean 6
standard deviation of 2.79 cm d21 6 2.05, respectively.
Generally, lakes in Northeastern Ontario had larger
amplitudes and greater flood rates.
We used PCA to extract empirical hydrologic endpoints
associated with WLF. A total of 10 axes were extracted; the
first three axes explained 89.9% of the cumulative variance
(Table 2). Based on day of year, axis 1 separated lakes
along day 210; axis 2 separated lakes between day 80 and
days 126147; and axis 3 separated lakes along day 300
(Fig. 3a; axis 3 not shown). BMI richness was most
strongly correlated with axis 1 (r 5 20.51). D80-D210
values ranged between +8.42 cm and 224.17 cm, with a
mean 6 standard deviation of 27.39 cm 6 7.97,
respectively. A set of hydrologic endpoints was correlated
with the WLF-derived principal components. We found
R
Fig. 3. Ordination of PCA showing (a) vector response of WLF
with axes 1 and 2, (b) vector response of measured hydrologic and
temperature variables with axes 1 and 2, and (c) vector response of
water quality, landscape and lake morphology variables with axes
1 and 2. Although 73 environmental variables were included in
analyses only variables with r values greater than 0.45 with either
axis 1, 2, or 3 are displayed. Vector length and direction is

proportional to its relationship with each axes. Open circles

represent lakes in northeastern Ontario, solid circles represent
lakes in northwestern Ontario, and solid triangles represent lakes
in north-central Ontario (see Fig. 1).

Water level and benthos

that axis 1 had the highest correlation with D80-D210 (r 5
0.83); axis 2 was correlated with both amplitude (r 5 0.87)
and flood rate (r 5 0.77), and recessional limb rate (r 5
20.84); and axis 3 was correlated with MWLD (r 5 20.74)
and WLRD (r 5 20.58) (Fig. 3b; Table 3). All watertemperature data were most strongly correlated with axis 1.
This could be due to the complementary effect of changes
in mean water level adding increased variability to watertemperature readings: as the data logger records temperature at a fixed position. Only three water-quality variables
demonstrated strong correlations with axis 1, axis 2, or axis
3: conductivity correlated with axis 1 (r 5 20.71); DO
correlated with axis 2 (r 5 0.70); and ORP between axis 1
and axis 2 correlated with r values of 0.44 and 20.47,
respectively. Interestingly, no lake morphometry or landclass variables had strong correlations with axis 1. The four
parameters: basin area, basin : lake area ratio, deep water,
and open fen showed the strongest correlations with axis 2,
with r values of 0.68, 0.71, 20.61, and 0.52, respectively.
Mean depth, lake area, and littoral slope had similar
strength correlations with both axis 1 and axis 3 (Table 3).
Linear regression of: principal components (n 5 10),
hydrologic variables (n 5 9), water temperature (n 5 8),
habitat characteristics and lake morphology (n 5 10), water
quality (n 5 8), and land classification (n 5 28) with BMI
richness revealed four significant (p , 0.05) relationships:
BMI richness decreased as axis 1 scores increased (r2 5
0.25); BMI richness decreased with increased D80-D210
values (r2 5 0.38); BMI richness decreased with increased
littoral slope (r2 5 0.32); and BMI richness increased with
lake area (r2 5 0.38) (Fig. 4; axis 1 relationship not

2279

displayed). A better fit (higher r2) of lake area with BMI

richness was found using a unimodal Gaussian (threeparameter) equation: f 5 a?exp(20.5?((x 2 x0)/b)2), p 5
0.001, r2 5 0.69 (Fig. 4b). Forward, stepwise, multipleregression analysis resulted in lake area and littoral slope
explaining BMI richness y 5 1.1764e27(lake area) 2
0.5069(littoral slope) + 26.0972 (p 5 0.03). Predicted versus
actual BMI richness resulted in an r2 of 0.58. The inclusion
of any predictor with D80-D210 did not yield significant
results. Analysis of collinearity confirmed that D80-D210,
lake area, and littoral slope are not collinear with respect to
BMI species richness because the variance inflation number
was , 10 and condition index was , 30.
A total of 49 taxa were identified, and mean BMI richness
values ranged from 16 to 33, with mean 6 standard
deviation of 23.94 6 4.71, respectively. Indicator species
analysis identified four taxa with significant D80-D210
group designation (Table 4). These taxa were the Gomphidae, Hydracarina, Leptoceridae, and Macromiidae and were
all determined to be indicative of mid D80-D210 water levels
(23.30 cm to 28.90 cm) and had p values of 0.01, 0.01, 0.02,
and 0.03, respectively. Most taxa were designated to the mid
and low D80-D210 groups but had p values . 0.05.

Discussion
We found that natural WLF in large boreal lakes varied
little (35.9157.5 cm) compared with reservoirs created for
power production, which vary several fold more (e.g., 1
7 m) (Aroviita and Hamalainen 2008). Although the WLF
were small, we still found significant effects on the BMI

Table 3. Pearson correlation coefficients (r) and p values for axis 1, axis 2, and axis 3 with environmental variables having r values
. 0.45 with one of the first three axes. All p values , 0.05 are identified in bold.
Axis 1

Axis 2

Axis 3

Environmental variable

D80-D210
Amplitude
Flood rate (15 d)
Flood rate
Recessional limb rate
Melt day
MWLD
WLRD
CV water level
CV water temperature
Mean temperature
Temperature to 4uC
Elevation
DO
Conductivity
ORP
Mean Depth
Basin area
Lake area
Basin : lake area ratio
Littoral slope
Open water
Open fen
Treed bog
BMI richness

0.83
20.34
20.35
20.23
0.45
20.05
20.33
20.30
20.30
20.54
0.59
20.76
0.72
20.37
20.71
0.44
20.43
20.26
20.47
20.12
0.45
0.09
20.33
0.16
20.51

0.0001
0.1930
0.1863
0.3892
0.0804
0.8449
0.2199
0.2646
0.2643
0.0326
0.0171
0.0006
0.0018
0.1601
0.0022
0.0855
0.3845
0.3317
0.0659
0.6644
0.0792
0.7510
0.2129
0.5489
0.0463

20.37
0.87
0.76
0.77
20.84
0.64
20.11
20.42
0.85
20.25
20.03
0.15
20.28
0.70
0.33
20.47
20.30
0.68
20.05
0.71
20.10
20.61
0.52
0.70
0.15

0.1582
0.0001
0.0006
0.0005
0.0001
0.0071
0.6904
0.1083
0.0001
0.3535
0.9187
0.5875
0.2922
0.0024
0.2152
0.0654
0.5322
0.0038
0.8637
0.0021
0.7121
0.0117
0.0398
0.0025
0.5701

0.31
0.26
0.46
0.54
20.20
20.33
20.74
20.58
0.41
20.10
20.16
0.22
20.11
0.10
20.01
0.03
0.48
20.15
20.53
0.35
0.44
20.39
20.07
0.04
20.45

0.2371
0.3281
0.0714
0.0326
0.4524
0.2140
0.0010
0.0185
0.1123
0.7029
0.5443
0.4245
0.6757
0.7186
0.9587
0.9186
0.6635
0.5900
0.0363
0.1802
0.0907
0.1403
0.8011
0.8919
0.0821

2280

White et al.

Fig. 4. Regression analysis of (a) littoral slope with macroinvertebrate richness (slope 5 20.64, p 5 0.02linear), (b) lake area with
macroinvertebrate richness (slope 5 0.14, p 5 0.01linear; p 5
0.0005unimodal), and (c) D80-D210 with macroinvertebrate richness
(slope 5 20.36, p 5 0.01linear). Solid lines represent linear regression
and dashed line represents nonlinear regression employing a
unimodal Gaussian (3 parameter) equation: f 5 a?exp(2.5?((x 2
x0)/b)2). Open circles represent lakes in northeastern Ontario, solid
circles represent lakes in northwestern Ontario, and solid triangles
represent lakes in north-central Ontario (see Fig. 1).

community. As such, it appears that BMI are influenced by

natural WLF and our hypothesis was supported.
The two most prominent hydrologic features of lake
WLF, were amplitude and change in mean water level
measured as D80-D210. Of the many water-level parameters measured, only the D80-D210 measure yielded
significant results with BMI richness. With the exception
of one lake, our study lakes experienced a decrease in mean
water levels over the 20062007 period. This reduction in
mean water level, quantified as D80-D210, ranged between
+8.42 cm and 224.17 cm. We found that lakes with the
greatest decrease in water levels (at time of sampling)
relative to winter water level had elevated BMI richness in
stony littoral habitats (Fig. 4c). The causality of this
relationship is unclear. It is highly plausible that decreased
water level reduces available littoral habitats resulting in a
habitat squeeze effect.
In addition to D80-D210, lake area and littoral slope were
significantly correlated with BMI richness and, although not
collinear, all three environmental descriptors had high
correlations. Although lakes were chosen based on a
minimum size (. 10 km2), not having more stringent lakesize and littoral-slope criteria limit the strength of the study
results: as lake area revealed a classic speciesarea curve
relationship (Fig. 4b) and species richness declined with
increased littoral slope (Fig. 4a). The best multivariate
model predicting BMI richness included lake area and
littoral slope. This indicates that although D80-D210
explains some variance in BMI richness it may not be as
important as lake area in predicting richness. The negative
influence of increased littoral slope on BMI diversity has
been shown in previous studies (Scheifhacken et al. 2007);
however, its univariate relationship with BMI richness in our
study proved weaker than D80-D210 change in water level.
The linear relationships of D80-D210 and lake area with
BMI richness had identical r2 values (0.38), but we also
found that lake area exhibited a strong unimodal relationship with BMI richness (r2 5 0.69). The lake-area
relationship with BMI diversity is not surprising (Heino
2008) and has been documented for many aquatic species
(Dodson 1992; Griffiths 1997). However, evidence that
changes in mean water level alter BMI richness regardless
of lake area or littoral slope is supported by White et al.
(2008), who analyzed a long-term data set (20 yr) and
found similar results to this study: a long-term unimodal
pattern between mean water level and BMI richness in one
lake. We show here that the pattern holds across multiple
lakes. Although our study focuses on large lakes, similar
results are expected in smaller lakes; however, it is
important to recognize that smaller lakes have different
environmental controls that may further confound an
assessment of the effects of WLF. For example, water
transparency is a good predictor of epilimnion depth in
small lakes (,5 km2), but no such pattern was found in
large lakes (Fee et al. 1996).
We also found that the WLFBMI relationship is
dictated by the region in which the lake is found. There is
clearly a regional relationship among western, central, and
eastern regions of Ontario (Fig. 4c). The water level of a
lake is indeed driven by regional, and to a lesser degree

Water level and benthos

2281

Table 4. Indicator taxa analysis of 49 identified BMI families with three D80D210 WLF groups. Relative abundance is the average
abundance of a given taxa in a given group of sites over the average abundance of that taxa in all sites, expressed as percentage. Relative
frequency is the percent of sites in a given group where the given taxon is present. Indicator value is the relative abundance multiplied by
the percent relative frequency. Monte Carlo significant tests of group designations were run using 1000 permutations. The high water
level group had D80-D210 values between +8.42 and 23.30 cm, n 5 5; the middle water level group had D80-D210 values between 23.30
and 28.90 cm, n 5 5; the low water level group had D80-D210 values between 28.90 and 224.20 cm, n 5 6. Taxa with significant p value
(, 0.05) are in bold.
Relative abundance

Relative frequency

Monte Carlo
significance test

Indicator value

Taxa

High

Mid

Low

High

Mid

Low

High

Mid

Low

Group

Gomphidae
Hydracarina
Leptoceridae
Macromiidae
Ephemeridae
Corixidae
Chironomidae
Lumbriculidae
Chaoboridae
Dyticidae
Helicopsychidae
Empididae
Hydroptilidae
Lymnaeidae
Tabanidae
Erpobdellidae
Tubificidae
Polycentropodidae
Physidae
Planorbidae
Hydrobiidae
Psephenidae
Elmidae
Sphaeridae
Caenidae
Tipulidae
Ancylidae
Hyalellidae
Lepidostomatidae
Ceratopogonidae
Prostoma
Coenagrionidae
Aeshnidae
Baetiscidae
Perlidae
Glossiphonidae
Valvatidae
Cambridae
Phygranidae
Baetidae
Naididae
Heptageniidae
Ephemerellidae
Leptophlebiidae
Capniidae
Chloroperlidae
Glossosomatidae
Psychomyiidae
Sisyridae

0
18
16
0
6
0
11
14
0
0
3
21
25
0
0
23
3
30
32
2
2
52
19
11
6
78
0
14
35
16
100
100
0
0
39
21
0
79
55
19
36
40
27
35
22
0
0
0
0

93
60
56
100
24
0
64
28
77
91
41
62
62
0
0
13
32
46
61
75
66
28
31
41
56
0
64
42
45
50
0
0
100
100
37
44
52
12
0
35
36
27
35
32
0
0
0
0
0

7
23
28
0
69
100
25
58
23
9
56
17
13
100
100
64
65
24
6
23
33
20
50
48
39
22
36
44
19
34
0
0
0
0
25
35
48
9
45
46
28
33
38
33
78
100
100
100
100

0
100
100
0
20
0
100
80
0
0
20
40
20
0
0
20
20
100
60
40
20
80
80
100
40
20
0
100
40
60
20
20
0
0
40
20
0
20
20
60
100
100
40
100
20
0
0
0
0

80
100
100
60
40
0
100
100
40
40
60
100
60
0
0
20
80
100
60
80
60
80
80
100
60
0
20
80
60
80
0
0
20
20
60
40
20
20
0
80
100
100
60
100
0
0
0
0
0

50
100
100
0
83
50
100
100
17
17
83
100
33
33
33
50
67
83
50
100
83
33
83
100
100
17
17
100
33
83
0
0
0
0
17
67
33
17
17
67
100
100
50
100
17
17
17
17
17

0
18
16
0
1
0
11
11
0
0
1
9
5
0
0
5
1
30
19
1
0
42
15
11
2
16
0
14
14
10
20
20
0
0
15
4
0
16
11
11
36
40
11
35
4
0
0
0
0

75
60
56
60
10
0
64
28
31
36
25
62
37
0
0
3
26
46
37
60
40
22
25
41
33
0
13
33
27
40
0
0
20
20
22
18
10
2
0
28
36
27
21
32
0
0
0
0
0

3
23
28
0
58
50
25
58
4
1
46
17
4
33
33
32
43
20
3
23
27
7
41
48
39
4
6
44
6
29
0
0
0
0
4
23
16
2
8
30
28
33
19
33
13
17
17
17
17

Mid
Mid
Mid
Mid
Low
Low
Mid
Low
Mid
Mid
Low
Mid
Mid
Low
Low
Low
Low
Mid
Mid
Mid
Mid
High
Low
Low
Low
High
Mid
Low
Mid
Mid
High
High
Mid
Mid
Mid
Low
Low
High
High
Low
High
High
Mid
High
Low
Low
Low
Low
Low

0.0097
0.0132
0.0238
0.0341
0.0666
0.0697
0.1075
0.1107
0.1432
0.1432
0.1918
0.2119
0.267
0.2884
0.2896
0.3165
0.3272
0.334
0.4399
0.4503
0.456
0.4722
0.4902
0.5225
0.5281
0.54
0.5455
0.5685
0.5764
0.6151
0.6229
0.6242
0.6267
0.6323
0.714
0.7194
0.7322
0.7797
0.799
0.8203
0.8975
0.8994
0.9288
0.9891
1
1
1
1
1

2282

White et al.

local, climatic influences (Bengtsson and Malm 1997).

Interestingly, inspection of D80-D210 regression with BMI
family richness (Fig. 4c) reveals that in both the northwest
and the northeast BMI family richness increases with
decreasing D80-D210; however, this pattern does not hold
when considering the north-central lakes separately. The
inconsistent pattern of north-central lakes is probably a
consequence of a low lake-sample size (three lakes) and that
all three lakes had similarly low D80-D210 water levels
(216 cm to 224 cm), with no lakes exhibiting water levels
similar to over-winter levels. Another regional effect may
result from possible geospatial patterns in fish community
structure, as top-down control of BMI community
structure through fish predation probably occurs in our
study lakes (Vander Zanden and Vadeboncoeur 2002).
Although we do not know the fish community structure of
our study lakes, it is expected that fish communities would
be similar across these large (. 10 km2) abiotically similar
northern temperate lakes (Jackson et al. 2001). As such,
slight differences in fish community structure would
probably only obscure detectability of BMIWLF relationships across our study lakes.
Only 4 of the 49 identified taxa were found to be
significant indicator taxa and all were designated to the mid
D80-D210 water-level group. The four taxa designated to
this group (Gomphidae, Macromiidae, Hydracarina, and
Leptoceridae) are relatively mobile species; however, there is
no experimental evidence in the literature to suggest that a
change of , 10 cm in mean water level would preferentially
benefit the survival of these taxa. One possibility is that mid
water levels concentrate detrital matter in rocky littoral
habitats, augmenting the proportionally dominant Chironomidae taxon (Table 3) and resulting in increased forage
efficiency, and thus survival, of predators such as Hydracarina (Proctor and Pritchard 1989) and Odonata (Merritt et al.
2008). However, at extremely low water levels it is possible
that disturbance is great enough that detrital matter is
severely agitated so that no taxon exhibits a preference
strong enough to be statistically designated to the low waterlevel group (Table 3). Further, there is increased taxa
richness in lakes designated to the mid and low D80-D210
taxa groups, supporting the occurrence of a habitat squeeze
(i.e., lakes with high water levels have lower taxa richness
and do not contain unique taxa that are not found in lakes
with mid and low water levels). Due to the qualitative nature
of our sampling regime, relative frequency may be a more
accurate measure of taxa response to changes in water level
than indicator value: which is a product of relative frequency
and relative abundance.
Our study shows that relatively small changes in mean
water level (33 cm) can affect BMI richness. The majority
of WLF research in temperate lake systems involves the
study of reservoirs with the conclusion that yearly
amplitude is a driver of biotic richness in littoral areas
(Hill et al. 1998; Aroviita and Hamalainen 2008). The
magnitude of yearly amplitudes (minimum minus maximum water level) reported in these studies varies several
fold but is almost always greater than 2 m. Our study shows
that a much smaller fluctuation (% 1 m) can be an
important factor structuring macroinvertebrate communi-

ties in natural lake ecosystems. This finding is important

because it highlights that even small incremental changes in
water quantity and water level associated with climate
change may alter benthic communities even without the
confounding effect of anticipated warmer water temperatures (Magnuson et al. 1997).
The sampling methodology of this study focused on
maximizing the number of lakes sampled at the cost of
reduced sampling precision of within lake variability (Bailey
et al. 2004). By increasing the number of stony littoral sites
sampled per lake, we would have increased the precision of
within lake variability (Reid et al. 1995), which would
further strengthen the relationship between BMI family
richness and the D80-D210 hydrologic measure. Nevertheless, our sole focus on stony littoral habitats reduced habitatrelated variability (Bailey et al. 2004) as well as our ability to
draw conclusions for other types of littoral habitats. Since
stony littoral habitats are structured and maintained by the
physical process of wave action (Jackson et al. 2001), it is
safe to assume that our results on the effects of WLF on BMI
are conservative. Stony littoral BMI communities are
comprised of taxa that are likely more tolerant of physical
water-level disturbance. It is possible that other habitats
(macrophyte, mud, woody debris) are composed of macroinvertebrate taxa that are more sensitive to WLF compared
to those in stony littoral areas.
In our study, many landscape variables can be used to
infer predictors typically extracted from a water balance
approach (Fig. 3). For example, an indicator of yearly
amplitude is basin area, or more specifically, as the basinto-lake-area ratio increases so does amplitude. This
relationship is well known (Bengtsson and Malm 1997),
but many other factors also contribute to the quantity of
water in lake systems: depth of snow pack (Nijssen et al.
2001), frost depth (Bayard et al. 2005), precipitation,
evaporation, preceding water content in surface stores
and soils (Metcalfe and Buttle 1999), and seepage at the
lake bottom (Sebestyen and Schneider 2001). Interestingly,
we found a positive correlation between amplitude and
percentage of treed bog and open fen in the catchment,
whereas amplitude was negatively correlated with open
water. This is surprising because it has been shown that
swamps seemingly do not influence or regulate seasonal
runoff in boreal areas (Devito et al. 1996); however, the
influence of the landscape to reduce runoff is dependant
upon the previous years hydrologic inputs (Metcalfe and
Buttle 1999). Boreal lake water levels in our study region
followed a 812-yr cycle that reached minimum levels in
2003 (White et al. 2008). Thus in our study lakes, which were
sampled in 2006, it is possible that open fen and treed bog
wetlands had recharged to maximum water levels, becoming
supersaturated and therefore increasing water runoff rather
than acting as a sink for spring melt. Conversely, open water
(lakes) had slower recharge responses to precipitation
compared to treed bog and open fens.
We found that large lakes (in both area and mean depth)
were at lower elevations and reached a temperature of 4uC
earlier in the year but nevertheless had lower mean
temperatures compared with smaller lakes found at higher
elevations. The larger low-elevation lakes had lower D80-

Water level and benthos

D210 water levels than smaller high-elevation lakes. The
earlier rise in water temperature of the lower larger lakes is
expected, as air temperatures are higher at lower elevations
and air temperature is the largest predictor of ice breakup
(Assel and Robertson 1995). Although ice breakup is
delayed in the higher elevation lakes, these lakes probably
reach a higher mean temperature due to their smaller water
volume and heat capacity. Direct linkages between
increased lake areas having low D80-D210 water levels
are unclear. Possible explanations include landscape
position (Webster et al. 1996), increased evaporation, or
increased seepage (Sebestyen and Schneider 2001).
Out of eight water-quality parameters that we examined,
only three had strong correlations with hydrologic character: DO, ORP, and conductivity. The separation of DO and
ORP along axis 2 and their relationship with amplitude
may be a result of increased water replenishment (Ambrosetti et al. 2003). During spring snowmelt lakes with
larger amplitudes receive larger inputs of melt water
relative to the lakes surface area, which may increase the
flushing and mixing of deoxygenated water (Ambrosetti et
al. 2003) accumulated over the winter months (Jarvinen et
al. 2002). The response of conductivity along axis 1 is a
direct effect of north-central Ontario lakes having elevated
conductance due to underlying glacial deposits.
We found evidence that a habitat squeeze may result from
small changes in water level (33 cm). The high productivity
of littoral areas warrants further investigation toward this
habitat squeeze hypothesis. Particularly, the responsiveness
of nutrient cycling processes to changes in natural water
level in boreal lacustrine environments needs to be better
understood. The biotic implications of a habitat squeeze may
lead to particularly insightful intra- and interspecific
competition research. A focused examination of trophic
dynamics under habitat squeeze may reveal significant
effects on higher trophic fauna that have large societal
influence such as birds and fish. Finally, while much research
has been devoted to understanding climatic, geomorphological, and landscape controls of WLF in lentic ecosystems,
what is needed is research that undertakes a long-term (10
20 yr), multidisciplinary approach that combines detailed
water balance study and aquatic community structure across
multiple lake and watershed basins.
Acknowledgments
We thank Ryan Stainton and Stephanie Lyons for their
assistance in the field and in the lab. We also thank Mark Hanson
and two anonymous reviewers whose suggestions improved this
manuscript. This work was supported by funding from the
Ontario Ministry of Natural Resources through the Institute for
Watershed Science, Trent University and from Canadas Natural
Sciences and Engineering Research Council (NSERC) Discovery
grant and NSERC University Faculty Award to M.A.X.

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Associate editor: Alexander D. Huryn

Received: 23 November 2009
Accepted: 15 June 2010
Amended: 27 July 2010