Aquatic and terrestrial drivers of dragonfly (Odonata) assemblages within and among

north-temperate lakes
Author(s): Alysa J. Remsburg and Monica G. Turner
Source: Journal of the North American Benthological Society, 28(1):44-56. 2009.
Published By: North American Benthological Society
DOI: 10.1899/08-004.1
URL: http://www.bioone.org/doi/full/10.1899/08-004.1

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J. N. Am. Benthol. Soc., 2009, 28(1):44–56
Ó 2009 by The North American Benthological Society
DOI: 10.1899/08-004.1
Published online: 18 November 2008

Aquatic and terrestrial drivers of dragonfly (Odonata) assemblages

within and among north-temperate lakes

Alysa J. Remsburg1 AND Monica G. Turner2

University of Wisconsin-Madison, Department of Zoology, Madison, Wisconsin 53706 USA

Abstract. The physical structure of vegetation influences diversity, interactions, movement, and
thermoregulation of animals. Vegetation structure might be a good indicator of habitat requirements of
generalist predators, such as dragonflies and damselflies (order Odonata), and thereby affect species
diversity. Odonates use aquatic and terrestrial habitats during larval and adult life stages, respectively, but
the relative importance of vegetation in these habitats is poorly understood. We compared how aquatic and
riparian habitat variables affected odonate larvae from 41 sites (each 30 m in shoreline length) on 17 lakes in
northern Wisconsin. We used principal components analyses to reduce multiple habitat variables to 2 lake-
level axes (lake size and development, lake wetlands and predators), 2 site-level littoral axes (littoral
macrophytes, littoral muckiness), and 2 site-level riparian axes (riparian structural complexity, riparian tall
wetland plants). Most (61.6%) of the variance in larval species richness occurred at the site level. Density of
the most abundant family, Gomphidae, was positively related to riparian tall wetland plants, whereas
species richness was positively correlated with abundance of littoral macrophytes (on the basis of multiple
linear regression with an information theoretic approach). Surveys in 18 paired littoral microsites in 9 lakes
indicated that larvae from the clasper and sprawler behavioral guilds were most abundant in microsites
with submerged macrophytes. However, predation risk, assessed by tethering larvae in patches of
submerged macrophytes, did not differ between habitats with and without macrophytes. We tested whether
shoreline plants affected recruitment from the adult stage by comparing adult odonate behaviors in response
to 2 riparian vegetation treatments. Adult damselfly abundance was higher where we placed potted wetland
plants than at manicured lawns without tall vegetation. Our results indicate that odonate larvae might be
influenced by vegetation structure in both aquatic and riparian habitats and demonstrate how animals with
complex life histories link aquatic and terrestrial communities.

Key words: dragonflies, damselflies, riparian plants, habitat selection, insect behavior, vegetation struc-
ture, predator habitat, aquatic macrophytes, lakeshore development, complex life history, benthic insects.

Understanding how vegetation structure affects Animals that use multiple habitats for different
animal abundance and diversity has long been a behaviors or life stages might respond to vegetation
fundamental objective in ecology (e.g., MacArthur structure in each of the habitats they occupy. Studies
1965, Karr and Roth 1971). The physical structure of comparing the importance of aquatic and terrestrial
vegetation influences predator–prey interactions, ther- habitat features for animals that inhabit both of these
moregulation, landscape permeability (ease of move- ecosystems generally have focused on amphibians
ment), and mate detection (McCoy and Bell 1991). (Marnell 1998, Scribner et al. 2001, Ficetola et al. 2004,
Vegetation structure can successfully describe habitat Becker et al. 2007). Research investigating how
suitability for predacious species on land (e.g., Pianka Odonata (dragonfly and damselfly) species respond
to both aquatic and terrestrial habitat structure is
1967, Brose 2003, De Souza and Martins 2005, Muntifer-
lacking, particularly for their larval phase. The long
ing et al. 2006) and in the water (e.g., Werner et al. 1983,
aquatic phase (1–4 y) of many odonates in temperate
Carr 1991, Tupper and Boutilier 1995, Cheruvelil et al.
regions might increase the importance of aquatic
2002, Sass 2004, Milner and Gloyne-Phillips 2005).
habitat features for odonates relative to amphibians
1
Present address: Biodiversity Center, Unity College, and other aquatic insects with shorter aquatic phases.
Unity, Maine 04988 USA. E-mail: [email protected] Land–water ecotones are often structurally complex
2
E-mail address: [email protected] and provide critical habitat for many animals (De-
44

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2009] AQUATIC AND RIPARIAN PLANTS INFLUENCE DRAGONFLIES 45

camps et al. 2004). Structural complexity arises from and Magnuson 2000), fish predation might exert less
the combination of abiotic characteristics (e.g., sub- control on lacustrine odonate populations than is
strate, slope), littoral vegetation, and riparian vegeta- suggested by stream studies. Among lakes, we
tion. Human activity also interacts with natural drivers expected that odonate abundance and species richness
of structural complexity. Housing development is would decline with increasing fish abundance, lake
expanding rapidly in many riparian areas (Burger depth, and water clarity (Aksnes and Giske 1993,
2000, Hansen et al. 2005), leading to simplification of Davies-Colley and Smith 2001).
vegetation structure by removal of the shrub layer We used surveys of larval odonates at a variety of
(Racey and Euler 1982, Elias and Meyer 2003), floating lake and site conditions and field experiments to
macrophytes (Jennings et al. 2003, Radomski and address 3 questions: 1) How do abundance and
Goeman 2001), and coarse wood (Marburg et al. community composition of larval odonates vary with
2006). Those structural changes might have conse- habitat structure among and within lakes? 2) Does
quences for a variety of animals in both the riparian local macrophyte abundance within lakes affect
and littoral (near-shore aquatic) zones (Bryan and predation on odonate larvae? 3) Does presence of tall
Scarnecchia 1992, Schindler et al. 2000, Lindsay et al. riparian vegetation influence use of riparian sites by
2002, Woodford and Meyer 2003, Butler and DeMay- adult dragonflies?
nadier 2008, Henning and Remsburg, in press). The
relative influence of natural and anthropogenic drivers Methods
on Odonata communities is not well known.
Odonates generally show little response to particu- Study area
lar plant species (terrestrial or aquatic) (Buchwald We conducted our study in Vilas County in
1992, Foote and Hornung 2005), but adult odonate northeastern Wisconsin, USA, in the center of the
diversity and abundance often are correlated positive- Northern Highlands Lake District (NHLD). Curtis
ly with local abundance of vegetation (reviewed in (1959) classified the vegetation as northern wet forest
Remsburg 2007). Vegetation provides adults with and boreal forest. The forested landscape is interrupt-
perching structures for thermoregulation, foraging, ed by a high density of glacial lakes (.1300 lakes in the
territory defense, mate attraction, copulation, noctur- 2639-km2 county), which cover 13% of the NHLD
nal roosting, and protection from adverse weather surface area (Martin 1965). Lakeshores in the area have
(Buchwald 1992, Wildermuth 1993, McKinnon and experienced unprecedented housing development
May 1994, Rouquette and Thompson 2007), thereby (Schnaiberg et al. 2002, Gonzalez-Abraham et al. 2007).
potentially affecting their density and diversity. We surveyed larval Odonata assemblages at 41 lake
Vegetation and other physical structures can interact sites. We defined sites as 30-m segments along the
with top-down controls on odonate larvae or on shoreline, including the adjacent littoral zone to a
odonate tenerals (immature adults that have recently water depth of 1 m and the riparian zone 6 m inland
emerged from the water) to influence population sizes. from the ordinary high water mark. We distributed
Riparian vegetation, for example, might harbor both sites among 17 lakes in Vilas County, Wisconsin; the
emerging odonates and their predators (Duffy 1994, number of sites/lake ranged from 1 to 5. We selected
Whitaker et al. 2000). Mesocosm experiments have sites with qualitative differences in littoral or riparian
shown that odonate larvae find refuge from predators vegetation structure when sites were on the same lake.
in aquatic macrophytes (Crowder and Cooper 1982, We selected ½ of the sites from developed areas and
Thompson 1987, Dionne et al. 1990, Diehl 1992). the rest from undeveloped forested areas. All sites
However, local predator densities in lakes also tend were at least somewhat sandy, had only moderate
to increase where macrophytes offer protection or riparian slopes (,208), and were separated from each
higher prey densities (Werner et al. 1983, Weaver et al. other by 120 m (with only 2 pairs of sites ,500 m
1997, Gamboa-Perez and Schmitter-Soto 1999). Littoral apart).
coarse wood or mucky substrates also might provide
refugia from predators for some larval odonate species Aquatic and terrestrial drivers of Odonata larval density
(Schmude 1998, Alzmann et al. 1999, Burcher and and richness
Smock 2002). In addition, predation on odonates can
vary among water bodies as a result of differences in At each site, we collected odonate larvae with a D-
water chemistry or fish communities (Johnson and frame net by scooping 12 m2 of benthic material (0.6
Crowley 1980, McPeek 1990, Johansson and Brodin m2/scoop) on 1 d/site during June to July 2004.
2003). For example, if littoral fish diversity and Samples were collected along four 16-m-long transects
densities are lower in lakes than in streams (Willis that paralleled the shore. We centered transects along

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46 A. J. REMSBURG AND M. G. TURNER [Volume 28

the 30-m site shoreline and placed them at 0.25-, 0.50-, ranked by 2nd-order criterion Akaike’s information
0.75-, and 1.00-m water depths; transect distance from criterion values (Sugiura 1978). Strength of evidence
shore depended on littoral slope. Pilot studies sug- for each candidate model can be compared using
gested that odonate densities were highest in water Akaike weights, wi, the log likelihood of model i
.0.50 m deep, so we took 6 scoops along the 0.75- and divided by the sum of log likelihoods for all candidate
1.00-m transects and 4 scoops along the 0.25- and 0.50- models (Burnham and Anderson 2002). We used all
m transects. Larvae of most species (except some candidate models to estimate the relative importance
Zygoptera and smaller Libellulidae) were present of each predictor by summing the Akaike weights
throughout the summer because odonates generally (Rw) over all models in which a variable appeared
are semivoltine in northern Wisconsin lakes (R. (Burnham and Anderson 2002).
DuBois, Wisconsin Department of Natural Resources,
personal communication). We sorted larvae from Macrophyte effects on larval densities within sites
benthic material by hand, preserved them in 70%
ethanol, and identified them to species or family (for Submerged macrophtye biomass effects.—To evaluate
suborder Zygoptera) using the keys of Needham et al. effects of submerged macrophytes on odonate larval
(2000) and Westfall and May (1996). densities at a finer scale than the site-level surveys, we
We characterized 12 lake-level variables, including sampled larvae at 18 microsites (5–10 m2) in 9 lakes in
fish abundance, human activity, vegetation, and water July 2006. In each lake, we chose 1 microsite with
chemistry in each lake. We recorded 9 littoral and 10 dense submerged macrophytes and 1 adjacent micro-
riparian habitat variables at each site (Table 1). We site with no macrophytes. The macrophyte and non-
conducted separate principal components analyses macrophyte microsites were separated by 4 m within
(PCA; PC-ORD, version 4; MjM, Gleneden Beach, each lake. Paired microsites had similar substrates and
Oregon) on lake, littoral, and riparian variables to distances from man-made structures. Within each
reduce these predictor variables to a smaller set of microsite, we used a D-net to collect benthos enclosed
orthogonal variables (Remsburg 2007). We based by a cylinder (diameter ¼ 0.55 m) at 5 adjacent
PCAs on centered, standardized matrices of Pearson subsampling locations at 0.3-m depth and 5 adjacent
correlation coefficients. We examined PCA eigenvalues subsampling locations at 0.5-m depth. We pooled the
and habitat variable correlations with the PCA scores, 10 subsamples and counted all larvae in 2.4 m2 of
and chose to use the first 2 PCA axes from each benthos at each microsite as replicates.
category of variables in mixed-effects multiple linear We used a paired Wilcoxon test to compare densities
regressions of larval densities. of 3 larval behavioral guilds between microsites with
We used mixed-effects multiple linear regression and without macrophytes. Behavioral guilds provided
models to compare the effects of lake-level and littoral more informative response variables than families
and riparian site-level habitat variables on odonate because densities of some families were very low
density, density of the most common family (Gomphi- within the microsites. Guilds included burrowers,
dae), and species richness (=[x] transformed to meet which hide in detritus or under the sand; sprawlers,
normality assumptions) across the 41 sites. We which lie on top of detritus; and claspers, which climb
assigned lake as a random effect because specific lakes on the leaves, stems, and roots of macrophytes
were not of interest, but sites on the same lakes were (Appendix).
not independent. We used similar mixed-effects Tethering experiment.—In August 2006, we conduct-
logistic models, including a fixed-effects term for area ed a predation experiment at 42 microsites from 21
sampled, to compare habitat effects on presence or lakes (including the 18 microsites used for surveys).
absence of the less-abundant Zygoptera suborder and We paired microsites within lakes. Macrophytes varied
the 4 other Anisoptera families observed. We present among lakes, so we recorded macrophyte species and
R2 values for linear regressions using the most designated macrophyte biomass categories (none, low,
influential single independent variable. All analyses moderate, or high). At each microsite, we used
were completed using R software (version 2.4.1; R Supergluet to attach monofilament fishing lines (30–
Core Development Team, Vienna, Austria). 40 cm in length) onto the abdomens of 8 live larvae (in
We used an information theoretic approach, which later instars) collected from nearby lakes or ponds. We
avoids arbitrary cutoff values (e.g., a ¼ 0.05) and used a variety of Anisoptera species and larval instars,
Bonferroni problems of multiple tests (Burnham and but most larvae were Cordulia shurtleffi (2 cm long). We
Anderson 2002), to compare alternative linear models controlled for variations in larval behavior and
containing 0 to 6 PCA scores. The 64 candidate models vulnerability to predation by matching larval species
(permutations for all possible combinations) were and sizes (range: 15–30-mm length) for paired micro-

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2009] AQUATIC AND RIPARIAN PLANTS INFLUENCE DRAGONFLIES 47

TABLE 1. Pearson correlation coefficients between habitat variables and the best 2 axes generated by 3 separate principal
components analysis (PCA) procedures on lake-level, site-level littoral, and site-level riparian habitat descriptors at 17 lakes in Vilas
County, Wisconsin (Remsburg 2007). Littoral sites consisted of 30 m along the lakeshore contour and the adjacent littoral zone with
water depth ,1 m. Riparian sites were 30 m along the shore 3 6 m inland, but % cover variables were measured within 0.5 m of the
ordinary high water mark. Descriptive labels provided for each axis and analysis refer to the major habitat features captured by the
axis. CPUE ¼ catch per unit effort.

Pearson correlation coefficients

Habitat descriptors PCA axis 1 PCA axis 2
Lake-level Lake size and development Lake wetlands and predators
Insectivorous fish CPUEa 0.20 0.73
Total fish CPUEa 0.24 0.34
Conductance (lS/cm)b 0.67 0.18
Secchi depth (m)b 0.16 0.07
Morphometryb 0.60 0.54
Area (ha)b 0.75 0.14
Maximum depth (m)b 0.49 0.22
Perimeter (km)b 0.87 0.30
Length of wetlands (m)c 0.15 0.85
% perimeter with wetlandsc 0.15 0.78
Building density (/km)d 0.56 0.40
Number of buildingsd 0.89 0.03
Site-level littoral Littoral macrophytes Littoral muckiness
Sand (ordinal variable: 0, 1, 2)e 0.23 0.77
Rock (ordinal variable: 0, 1, 2)e 0.44 0.21
Gravel (ordinal variable: 0, 1, 2)e 0.63 0.10
Muck (ordinal variable: 0, 1, 2)e 0.53 0.64
Mean macrophyte richnessf 0.90 0.09
Mean macrophyte % coverf 0.86 0.03
Mean frequency of emergent macrophytesf 0.66 0.28
Coarse wood (number/50 m)e 0.03 0.60
Slope (8) 0.48 0.47
Site-level riparian Riparian structural complexity Riparian tall wetland plants
Canopy (ordinal variable: 1, 2, 3) 0.70 0.03
Slope (8) 0.35 0.10
Length of shore with shrubs (m) 0.86 0.04
Length of shore with tall wetland plants (m)g 0.17 0.84
Vegetation distance from water (m) 0.46 0.26
Maximum understory foliage height (m) 0.68 0.25
Mean structural diversity (number of structure types
intersected) 0.73 ,0.01
Mean foliage height diversity (number of layers intercepted) 0.92 0.08
% shore herbaceous 0.12 0.82
% shore shrubs 0.56 0.58
a
CPUE based on electrofishing, minnow traps, and crayfish traps in 2001 to 2004 (Carpenter et al. 2006)
b
Data collected 2001 to 2003 (Carpenter et al. 2006)
c
Gergel (1996)
d
Schnaiberg et al. (2002)
e
Littoral substrate dominance and coarse wood abundance estimated at 0.5-m water depth in 2001 to 2004 (Marburg et al. 2006)
f
Macrophyte species richness and % cover estimated in 2001 to 2003 along a transect extending 0 to 2-m depth (Alexander 2005)
g
Carex, Typha, or Iris species

sites within the same lake. Laboratory observations exposure to predation at all microsites (timing paired
indicated that fish predation occurred on tethered by lake). We did all tethering experiments during the
larvae. We assumed that empty lines indicated afternoon (between 1200 and 1900 h), when fish
predation on odonates because larvae never became densities in macrophytes are probably greatest (Hall
unglued from fishing lines in the laboratory. We et al. 1979, Gliwicz et al. 2006). We used larval
attached tethered larvae just above the benthos by mortality (proportion of larvae missing after exposure)
tying fishing lines to submerged metal stakes. We to estimate relative predation rates.
retrieved tethered larvae remaining after 3.5 to 6 h of We used analysis of variance (ANOVA) with lake as

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48 A. J. REMSBURG AND M. G. TURNER [Volume 28

a random effect to compare mortality of tethered tion period to improve detection of the less active
larvae among 42 microsites with different macrophyte odonates perched on shore. We recorded the total
biomass categories. Variance partitioning indicated number of individuals present within the plot during
that a large portion of the variance occurred at the 10 min of observation. We estimated instantaneous
lake level. Therefore, we also used mixed-effects linear odonate abundance within the plot by counting all
regressions to test effects of secchi depth and conduc- individuals visible at the end of 10 min.
tance at the lake level on larval mortality (fish catch We used blocked mixed-effects ANOVAs to compare
per unit effort for these lakes was not available). Zygoptera and Anisoptera abundance estimates
among treatments, where observation day was treated
Effects of riparian vegetation on adult abundance as a random effect. We pooled responses by suborder
because odonate abundances were low overall, and the
We experimentally investigated whether riparian major differences in behaviors corresponded with
vegetation affects adult site use, which could help suborder.
explain larval distributions within lakes. We recorded
adult Odonata behavioral responses to riparian vege- Results
tation treatments in 2 blocks of lakeshore home sites.
One block had very dense aquatic macrophytes Aquatic and terrestrial drivers of Odonata larval density
(dense-macrophyte block), whereas the other had very and richness
few (sparse-macrophyte block). Both blocks were on We found larvae from 23 Odonata species at the 41
bays in the same lake (Little Saint Germain), had stone survey sites (Appendix). Species richness ranged from
seawalls separating trimmed lawns from the water, 0 to 10 species per site (mean ¼ 5.2 species). Gomphus
similar aspect, and riparian slope ’ 0. Ten meters spicatus was the most abundant (0.45 6 0.86 [SD]
separated the 2 treatment plots; plots were 15 m along larvae/m2) and frequently occurring species (present
the shoreline 3 2 m inland. at 67% of the sites). Gomphus, which included 3 other
The 2 riparian vegetation treatments applied to each species (Gomphus exilis, Gomphus lividus, and Gomphus
block were: 1) trimmed lawns with 20 dwarf cattail fraternus), was by far the most prevalent genus
(Typha minima) plants in pots added at the shoreline (occurring at 86% of the sites). The family Gomphidae
(Typha plots), and 2) trimmed lawns without potted or occurred at 87.8% of sites, whereas Zygoptera, Macro-
other tall plants (no-Typha plots). We chose the dwarf minae, Cordulinae, Libellulinae, and Aeshnidae oc-
cattails because they closely resembled the height (0.6 curred at 44, 34, 22, 20, and 17% of sites, respectively.
m) and rigidity of native sedges. We replicated both Correlations of habitat variables with the 6 PCA
treatments on 21 d at the dense macrophyte block and axes used in the multivariate regressions had straight-
on 19 d at the sparse macrophyte block. This design forward interpretations (Table 1). Total variance of the
might have allowed repeated observations of some 12 lake-level variables summarized by the first 2 axes
individuals, but we treated separate observation days of the PCA was 52.4% (axis 1 ¼ 30.4% [lake size and
as replicates because adults from the genera we development]; axis 2 ¼ 22.0% [lake wetlands and
observed move away from the shoreline to roosting predators]). The first 2 axes of the PCAs from the 9 site-
sites each evening (Askew 1982, Bried and Ervin 2006) level littoral and 10 site-level riparian variables
and probably do not demonstrate site fidelity across captured 53.9% (axis 1 ¼ 34.9% [littoral macrophytes];
multiple days (Jacobs 1955, Logan 1971). At both axis 2 ¼ 19.0% [littoral muckiness]) and 56.4% (axis 1 ¼
blocks, we alternated the 2 treatments each day 37.8% [riparian structural complexity]; axis 2 ¼ 18.6%
between the 2 plots to separate Typha treatment effects [riparian tall wetland plants]) of the variance, respec-
from littoral or location-specific influences. tively. The 3rd lake-level axis represented 14.6% of the
On each observation day, we set up the Typha variance (more of the variance than the 3rd site-level
treatments 3 h before recording odonate behaviors in axes represented) and was correlated almost exclu-
both treatments within the same block. To capture the sively with secchi depth (r ¼ 0.9). However, we did
period of greatest odonate activity, we made observa- not add the 3rd lake-level axis to the set of candidate
tions from 25 June to 18 August 2006 between 1200 models because secchi depth was not correlated with
and 1600 h on days with relatively low wind and odonate density or diversity. Scores from the 3
cloud cover. A single observer (AJR) stood at 1 corner separate PCAs generally were not correlated with
of the plot to survey odonates, including those over the each other (Pearson correlation coefficients , 0.3),
water adjacent to the riparian plot (within ;5 m of except for the 1st lake axis and 1st riparian axis (r ¼
shore), in the whole plot simultaneously. The observer 0.41).
also walked slowly across the plot twice per observa- More of the variance in density and richness of

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2009] AQUATIC AND RIPARIAN PLANTS INFLUENCE DRAGONFLIES 49

TABLE 2. Relative importance (sums of Akaike weights) of lake- and site-level summary habitat descriptors in predicting
Odonata larval densities at 41 Wisconsin lake sites. The 6 predictors are synthetic axes derived from 3 separate principal
components analyses of 12 lake-level, 9 site-level littoral, and 10 site-level riparian variables (see Table 1). Bold font indicates
statistical significance (p , 0.05).

Sums of Akaike weights from all candidate models with the predictor

Lake Littoral Riparian

Model type/ Size and Wetlands and Structural Tall wetland
dependent variable development predators Macrophytes Muckiness complexity plants
Linear regression
Odonata density 0.33 0.20 0.54 0.20 0.21 0.63
Species richness 0.29 0.22 0.60 0.21 0.26 0.28
Gomphidae density 0.40 0.20 0.42 0.20 0.22 0.73
Logistic regression
Zygoptera presence 0.23 0.22 0.73 0.33 0.31 0.34
Libellulinae presence 0.38 0.23 0.58 0.22 0.32 0.21
Macrominae presence 0.25 0.45 0.25 0.34 0.59 0.25
Cordulinae presence 0.73 0.23 0.24 0.87 0.32 0.25
Aeshnidae presence 0.34 0.20 0.35 0.95 0.45 0.65

odonates occurred at the site level (56.5 and 61.6%, increasing littoral macrophyte abundance, which was
respectively) than at the lake level (34.6 and 29.4%, also the best predictor for presence of Zygoptera and
respectively), and all of the variance in Gomphidae Libellulinae (Table 2). Littoral muckiness was the best
densities occurred at the site level. The information predictor for presence of Aeshnidae and Cordulinae
theoretic approach for model selection enabled us to (Table 2).
assess the importance of predictor variables across all
possible models, rather than only from a somewhat- Macrophyte effects on larval densities within sites
arbitrary best model. Density of all odonates combined Macrophyte biomass effects.—At the 18 paired micro-
was best explained by increasing riparian tall wetland sites, we found most of the same species as in our 2004
plants, followed by increasing littoral macrophytes survey and 3 new Libellulids: Leucorhinia hudsonica,
(Table 2). Importance of the riparian tall wetland plant Leucorhinia proxima, and Tramea carolina. Unlike in the
axis was even greater for the Gomphidae family, and initial survey, the most abundant species was Epitheca
the model fit was better than that of all odonates. The cynosura (0.45 6 1.0 larvae/m2), even though it
strongest predictor for Odonata species richness was occurred only at macrophyte microsites (Fig. 1).
Gomphus species occurred at 67% of the microsites.
We found many more odonates in microsites with
submerged macrophytes (5.2 6 3.8 larvae/m2) than in
microsites without (0.8 6 1.0 larvae/m2; Wilcoxon
rank sum test, W ¼ 74, p ¼ 0.004). Clasper and sprawler
larval behavioral guilds were significantly more
abundant in microsites with than without submerged
macrophytes (claspers: W ¼ 66, p ¼ 0.02; sprawlers: W ¼
65, p ¼ 0.03), whereas the burrower larval behavioral
guild did not differ significantly between microsites
with and without submerged macrophytes (p . 0.1;
Fig. 2). We observed higher Anisoptera species
richness at microsites with (5.22 6 3.35) than without
submerged macrophytes (2.11 6 1.83; Wilcoxon rank
sum test W ¼ 63, p ¼ 0.05).
FIG. 1. Rank-abundance curves showing mean (61 SE) Tethering experiment.—Tethered larvae mortality
number of individual larvae for each odonate species found ranged from 0 to 100% among the 42 microsites (53%
in 2.4 m2 lake microsites with and without dense submerged 6 32%). On separate occasions, we observed 2
macrophytes. bluegills and 1 small painted turtle feeding on tethered

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50 A. J. REMSBURG AND M. G. TURNER [Volume 28

FIG. 2. Mean densities (61 SE) of 3 larval behavioral

guilds among paired 7-m2 lake microsites with and without
submerged macrophytes.

larvae in the field. Mortality did not vary with

submerged macrophyte biomass category (F1,20 ¼
1.19, p ¼ 0.3) or with macrophyte presence (paired t-
test: t20 ¼ 1.13, p ¼ 0.3). Mean larval mortality was
47% (6 32%) at microsites with submerged macro-
phytes and 56% (6 35%) at microsites without
submerged macrophytes. Much of the total variance
in mortality (39.1%) occurred at the lake level, but
mortality was not correlated significantly with lake
clarity or conductance (p . 0.1).

Effects of riparian vegetation on adult abundance

FIG. 3. Mean (61 SE) number of adult Anisoptera (A) and
We observed adult odonates from 15 different Zygoptera (B) within 15- 3 7-m experimental plots at 2
genera at the experimental plots but only Enallagma blocks of lakeshore home sites in northern Wisconsin (1 with
(Zygoptera:Coenagrionidae), Leucorrhinia, and Libellula sparse and 1 with dense emergent macrophytes in the littoral
spp. (Anisoptera:Libellulinae) were present on .50% zone). Typha treatments consisted of 20 potted Typha minima
placed on the seawalls. Typha treatments were rotated
of the observation days (96%, 65% and 56%, respec-
between 2 plots/site on 21 d during June to August 2006.
tively). Abundances of Gomphidae adults at these
plots were much lower than those of Libellulidae
animals with complex life histories provides an
adults, probably because Gomphidae spend more time
additional link between terrestrial and aquatic ecosys-
perched in treetops than do Libellulidae. Anisoptera
tems. A growing body of research on agricultural
abundance was greatest at the dense-macrophyte
systems has demonstrated that vegetation structure in
block and did not differ significantly between Typha
one ecosystem affects insect abundances in an adjacent
treatments (p . 0.1; Fig. 3A). Zygoptera abundance
system (Langellotto and Denno 2004, Tscharntke et al.
was significantly higher at the Typha than at the no-
2005, Bianchi et al. 2006). In our study, site-level
Typha treatment (F1,51 ¼ 9.2, p ¼ 0.004; Fig. 3B).
riparian vegetation was a better predictor of total
larval Odonata density, Gomphidae density, and
Discussion
Macrominae presence than site- or lake-level aquatic
Riparian vegetation influenced larval odonate as- habitat features. In particular, larval density of the
semblages in the adjacent aquatic ecosystem. Previous most abundant family, Gomphidae, increased with the
studies on ecological links between riparian land- amount of riparian tall wetland plants adjacent to—
scapes and aquatic communities have focused primar- but not in—the lakes. Previous studies have linked
ily on detrital, nutrient, and structural subsidies abundance of adult odonates to riparian sedge or rush
(Wallace et al. 1997, Nakano et al. 1999, Bastow et al. abundance (Van Buskirk 1986, Clark and Samways
2002, Sass 2004, Baxter et al. 2005, Roth et al. 2007). 1996, Foote and Hornung 2005, Hofmann and Mason
Our results suggest that adult habitat selection by 2005b, Rouquette and Thompson 2007, Butler and

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2009] AQUATIC AND RIPARIAN PLANTS INFLUENCE DRAGONFLIES 51

DeMaynadier 2008), but our study shows that the Aeshnidae also might respond to littoral macrophytes
abundance of larval odonates is correlated with at a larger spatial or temporal scale than do other
abundance of terrestrial herbaceous vegetation. odonates.
The stronger relationship between larval abundance High species richness at sites with more littoral
and riparian tall wetland plants than with riparian macrophytes can be attributed to more Zygoptera and
structural complexity might reflect the role of the plant Libellulinae occurrences at those sites, consistent with
structure. Riparian tall wetland plants (Carex, Typha, previous studies showing the strongest associations of
and Iris spp.) are rigid and provide stable vertical odonate richness with macrophyte biomass (Weather-
structures on which emerging larvae complete hard- head and James 2001, Michaletz et al. 2005, Osborn
ening of the wings (Corbet 1999). However, Gomphi- 2005, Butler and DeMaynadier 2008). Zygopterans
dae (unlike most other Anisoptera families) also can require emergent macrophytes for endophytic ovipo-
emerge on horizontal surfaces (Eda 1963). A cause– sition, and both Zygoptera and Libellulinae larvae use
effect relationship between the availability of suitable macrophytes. Last, all Zygoptera and Libellulinae
emergence structures and odonate population abun- species are perchers as adults, so emergent macro-
dances remains to be demonstrated. Adult odonates phytes also might serve as important perching
might prefer rigid plants for perching because their structures for adults at oviposition sites (McKinnon
height and open structure facilitate thermoregulation and May 1994, Switzer and Walters 1999, De Marco
(May 1976, Pezalla 1979) or mate attraction (McKinnon and Resende 2004).
and May 1994). Riparian vegetation also could affect Predation risk for tethered larvae did not help
adult abundances if it were correlated with terrestrial explain why larval densities were highest at microsites
prey densities (Baird and May 1997, Whitaker et al. with dense macrophytes. Our results differ from those
2000, Garono and Kooser 2001, Henning and Rems- of mesocosm predation experiments in other systems
burg, in press). that have shown the importance of macrophytes in
The correlations we observed between larvae and predator avoidance by odonate larvae (Crowder and
riparian vegetation also might reflect adult selection Cooper 1982, Thompson 1987, Dionne et al. 1990,
for oviposition sites with highest survivorship of eggs Diehl 1992). Tethering can interfere with predator
and larvae (Storch and Frynta 1999). Adults cannot avoidance behaviors (Curran and Able 1998, Kneib
assess all aspects of aquatic habitat suitability for and Scheele 2000), but field and aquarium observa-
larvae (Wildermuth 1993). However, adult odonates tions gave us no reason to suspect that effects of
might use emergent aquatic plants or riparian plants tethering on larval behavior differed among microsite
as proximate cues of macrophyte, substrate, prey treatments. Nevertheless, unnaturally high predation
availability, or predator conditions for larvae in littoral rates resulting from tethering might have over-
sites (Buchwald 1992, Wildermuth 1992, McKinnon whelmed the effects of vegetation in our experiment.
and May 1994). Adult riparian site use generally Differences among fish communities among lakes also
corresponds with oviposition locations (Corbet 1999), might have obscured the predator avoidance effect in
and larvae are unlikely to move .20 m from the our field experiment. Alternatively, competition or
littoral site where eggs were laid (Ubukata 1984, foraging behaviors of odonate larvae might better
Schaffner and Anholt 1998, but see Alzmann et al. explain their associations with macrophytes than
1999), so adult habitat use could help explain larval predator avoidance (Johansson 1991, Schmude 1998,
distributions. Other studies have demonstrated indi- Elkin and Baker 2000, Hofmann and Mason 2005a),
rectly that adding artificial perches can lead to higher potentially because of higher predator densities inside
adult odonate densities (Wolf and Waltz 1988, Rehfeldt than outside macrophyte beds (Werner et al. 1983,
1990, Baird and May 1997, May and Baird 2002, De Weaver et al. 1997, Gamboa-Perez and Schmitter-Soto
Marco and Resende 2004). Our field experiment 1999). Additional field experiments, such as fish
provides some support for a link between adult site exclusion cages, will be necessary to clarify the specific
selection and larval densities at sites with riparian role of macrophytes for odonate larvae.
wetland plants, but more observations of Gomphidae Our findings that both riparian and littoral vegeta-
oviposition are needed to resolve the question directly. tion structure influence lentic Odonata assemblages
Aeshnidae, which are endophytic species, were most have several implications for conservation. Odonate
likely to occur at sites with greater littoral muckiness abundances and diversity could decline as a conse-
and riparian tall wetland plants; but their presence quence of the vegetation simplification and removal of
was not predicted by littoral macrophytes, possibly macrophytes and coarse wood that often accompanies
because we observed so few Aeshnidae larvae in lakeshore development (Racey and Euler 1982, Ra-
surveys. As the strongest fliers of all odonates, domski and Goeman 2001, Elias and Meyer 2003,

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52 A. J. REMSBURG AND M. G. TURNER [Volume 28

Marburg et al. 2006). Our observation that site-level SELYS 1840 (Insecta; Odonata; Gomphidae) from a still
vegetation structure affects abundance and diversity of water habitat. International Review of Hydrobiology 84:
charismatic insects potentially could motivate home- 299–313.
owners to maintain or restore wetland and littoral ASKEW, R. 1982. Roosting and resting site selection by
coenagrionid damselflies. Advances in Odonatology 1:
vegetation. Furthermore, invasion by rusty crayfish
1–8.
(Orconectes rusticus) leads to severe macrophyte BAIRD, J. M., AND M. L. MAY. 1997. Foraging behavior of
reduction in north-temperate lakes (Wilson et al. Pachydiplax longipennis (Odonata: Libellulidae). Journal
2004) and might indirectly reduce odonate diversity of Insect Behavior 10:655–678.
and density. BASTOW, J. L., J. L. SABO, J. C. FINLAY, AND M. E. POWER. 2002. A
Last, Odonata abundances can influence many other basal aquatic-terrestrial trophic link in rivers: algal
components of aquatic and riparian communities subsidies via shore-dwelling grasshoppers. Oecologia
because Odonata are predators of smaller inverte- (Berlin) 131:261–268.
brates throughout their complex life cycle (Thorp and BAXTER, C. V., K. D. FAUSCH, AND W. C. SAUNDERS. 2005.
Tangled webs: reciprocal flows of invertebrate prey link
Cothran 1984, Johnson et al. 1987, Van Buskirk 1988).
streams and riparian zones. Freshwater Biology 50:201–
High predation by adult odonates on terrestrial
220.
pollinators, including bees, moths, and flies, might BECKER, C. G., C. R. FONSECA, C. F. B. HADDAD, R. F. BATISTA,
even alter riparian plant reproduction (Knight et al. AND P. I. PRADO. 2007. Habitat split and the global decline
2005). At critical times, odonates make up a significant of amphibians. Science 318:1775–1777.
portion of the diets of fish (Crowder and Cooper 1982, BIANCHI, F., C. J. H. BOOIJ, AND T. TSCHARNTKE. 2006.
Johnson et al. 1995, Sass 2004) and birds (Orians and Sustainable pest regulation in agricultural landscapes: a
Horn 1969, Wissinger 1988), so habitat variables that review on landscape composition, biodiversity and
drive odonate abundances could lead to widespread natural pest control. Proceedings of the Royal Society
changes in both aquatic and terrestrial food webs. of London B: Biological Sciences 273:1715–1727.
BRIED, J. T., AND G. N. ERVIN. 2006. Abundance patterns of
dragonflies along a wetland buffer. Wetlands 26:878–883.
Acknowledgements
BROSE, U. 2003. Bottom-up control of carabid beetle commu-
This research was supported financially by the nities in early successional wetlands: mediated by
National Science Foundation (NSF) Biocomplexity vegetation structure or plant diversity? Oecologia
(Berlin) 135:407–413.
Program DEB-0083545, North-temperate lakes Long-
BRYAN, M. D., AND D. L. SCARNECCHIA. 1992. Species richness,
Term Ecological Research (LTER) site, an NSF Grad- composition, and abundance of fish larvae and juveniles
uate Research Fellowship, the Garden Club of Amer- inhabiting natural and developed shorelines of a glacial
ica, and the Animal Behavior Society. Tim Kratz, M. Iowa lake. Environmental Biology of Fishes 35:329–341.
Woodford, S. van Egeren, and the University of BUCHWALD, R. 1992. Vegetation and dragonfly fauna—
Wisconsin Trout Lake station staff provided invaluable characteristics and examples of biocenological field
logistical support. C. Ane offered statistical advice. We studies. Vegetatio 101:99–107.
thank W. Smith, R. DuBois, K. Tennessen, and R. BURCHER, C. L., AND L. A. SMOCK. 2002. Habitat distribution,
Bohanan for sharing their expertise on Odonata dietary composition and life history characteristics of
odonate nymphs in a blackwater coastal plain stream.
behaviors and taxonomy. B. Peckarsky, A. Ives, J.
American Midland Naturalist 148:75–89.
Zedler, C. Gratton, and S. Carpenter helped guide and
BURGER, J. 2000. Landscapes, tourism, and conservation.
critique the manuscript. Field and laboratory assistants Science of the Total Environment 249(1–3):39–49.
included M. Theis, P. Winkler, B. Henning, J. Batten, J. BURNHAM, K. P., AND D. R. ANDERSON. 2002. Model selection
Estreen, D. Hong, J. Iacarella, C. Leugers, A. Duong, and multimodel inference: a practical information-
and A. Olson. The manuscript was reviewed by A. theoretic approach. 2nd edition. Springer, New York.
Olson and 2 anonymous referees. BUTLER, R. G., AND P. G. DEMAYNADIER. 2008. The significance
of littoral and shoreline habitat integrity to the conser-
Literature Cited vation of lacustrine damselflies (Odonata). Journal of
Insect Conservation 12:23–36.
AKSNES, D. L., AND J. GISKE. 1993. A theoretical model of CARPENTER, S. R., T. K. KRATZ, J. KITCHELL, J. J. MAGNUSON, AND
aquatic visual feeding. Ecological Modelling 67:233–250. M. J. VANDER ZANDEN. 2006. Biocomplexity; coordinated
ALEXANDER, M. L. 2005. Lake ecosystem variation across field studies. National Biological Information Infra-
landscape position and human development gradients in structure Metadata Clearinghouse. (Available from:
Vilas County, Wisconsin. PhD Thesis, University of http://mercury.ornl.gov/metadata/lter/html/lter/
Wisconsin, Madison, Wisconsin. metacat.lternet.edu_knb_metacat_action_read_qformat_
ALZMANN, N., B. KOHLER, AND G. MAIER. 1999. Spatial xml_docid_knb-lter-ntl.126.2_5.html)
distribution, food and activity of Gomphus pulchellus CARR, M. H. 1991. Habitat selection and recruitment of an

W:/xml/jnabs/jnbs_2801/jnbs-28-01-05.3d  Wednesday, 12 November 2008  10:35 am  Allen Press, Inc.  Page 52

2009] AQUATIC AND RIPARIAN PLANTS INFLUENCE DRAGONFLIES 53

assemblage of temperate zone reef fishes. Journal of aquatic and terrestrial habitat for the European pond
Experimental Marine Biology and Ecology 146:113–137. turtle (Emys orbicularis): implications for conservation
CHERUVELIL, K. S., P. A. SORANNO, J. D. MADSEN, AND M. J. planning and management. Canadian Journal of Zoolo-
ROBERSON. 2002. Plant architecture and epiphytic macro- gy 82:1704–1712.
invertebrate communities: the role of an exotic dissected FOOTE, A. L., AND C. L. R. HORNUNG. 2005. Odonates as
macrophyte. Journal of the North American Bentholog- biological indicators of grazing effects on Canadian
ical Society 21:261–277. prairie wetlands. Ecological Entomology 30:273–283.
CLARK, T. E., AND M. J. SAMWAYS. 1996. Dragonflies (Odonata) G AMBOA -P EREZ , H. C., AND J. J. SCHMITTER-S OTO. 1999.
as indicators of biotope quality in the Kruger National Distribution of cichlid fishes in the littoral of Lake
Park, South Africa. Journal of Applied Ecology 33:1001– Bacalar, Yucatan Peninsula. Environmental Biology of
1012. Fishes 54:35–43.
CORBET, P. S. 1999. Dragonflies: behavior and ecology of GARONO, R. J., AND J. G. KOOSER. 2001. The relationship
Odonata. Cornell University Press, Ithaca, New York. between patterns in flying adult insect assemblages and
CROWDER, L. B., AND W. E. COOPER. 1982. Habitat structural vegetation structure in wetlands of Ohio and Texas. Ohio
complexity and the interaction between bluegills and Journal of Science 101:12–21.
their prey. Ecology 63:1802–1813. GERGEL, S. E. 1996. Scale-dependent landscape effects on
CURRAN, M. C., AND K. W. ABLE. 1998. The value of tethering north temperate lakes and rivers. MS Thesis, University
fishes (winter flounder and tautog) as a tool for assessing of Wisconsin, Madison, Wisconsin.
predation rates. Marine Ecology Progress Series 163:45– GLIWICZ, Z. M., J. SLON, AND I. SZYNKARCZYK. 2006. Trading
51. safety for food: evidence from gut contents in roach and
CURTIS, J. T. 1959. The vegetation of Wisconsin. University of bleak captured at different distances offshore from their
Wisconsin Press, Madison, Wisconsin. daytime littoral refuge. Freshwater Biology 51:823–839.
DAVIES-COLLEY, R. J., AND D. G. SMITH. 2001. Turbidity, GONZALEZ-ABRAHAM, C. E., V. C. RADELOFF, R. B. HAMMER, T. J.
suspended sediment, and water clarity: a review. Journal HAWBAKER, S. I. STEWARD, AND M. K. CLAYTON. 2007.
of the American Water Resources Association 37:1085–
Building patterns and landscape fragmentation in
1101.
northern Wisconsin, USA. Landscape Ecology 22:217–
DECAMPS, H., G. PINAY, R. J. NAIMAN, G. E. PETTS, M. E.
230.
MCCLAIN, A. HILLBRICHT-ILKOWSKA, T. A. HANLEY, R. M.
HALL, D. J., E. E. WERNER, J. F. GILLIAM, G. G. MITTELBACH, D.
HOLMES, J. QUINN, J. GIBERT, A. M. P. TABACCHI, F. SCHIEMER,
HOWARD, C. G. DONER, J. A. DICKERMAN, AND A. J. STEWART.
E. TABACCHI, AND M. ZALEWSKI. 2004. Riparian zones:
1979. Diel foraging behavior and prey selection in the
where biogeochemistry meets biodiversity in manage-
Golden Shiner (Notemigonus crysoleucas). Journal of the
ment practice. Polish Journal of Ecology 52:3–18.
Fisheries Research Board of Canada 36:1029–1039.
DE MARCO, P., AND D. C. RESENDE. 2004. Cues for territory
HANSEN, A. J., R. L. KNIGHT, J. M. MARZLUFF, S. POWELL, K.
choice in two tropical dragonflies. Neotropical Entomol-
BROWN, P. H. GUDE, AND A. JONES. 2005. Effects of exurban
ogy 33:397–401.
development on biodiversity: patterns, mechanisms, and
DE SOUZA, A. L. T., AND R. P. MARTINS. 2005. Foliage density of
branches and distribution of plant-dwelling spiders. research needs. Ecological Applications 15:1893–1905.
Biotropica 37:416–420. HENNING, B. M., AND A. J. REMSBURG. Effects of lakeshore
DIEHL, S. 1992. Fish predation and benthic community vegetation structure on avian and amphibian abundance
structure: the role of omnivory and habitat complexity. in northern Wisconsin. American Midland Naturalist (in
Ecology 73:1646–1661. press).
DIONNE, M., M. BUTLER, AND C. FOLT. 1990. Plant-specific HOFMANN, T. A., AND C. F. MASON. 2005a. Competition,
expression of antipredator behavior by larval damsel- predation and microhabitat selection of Zygoptera larvae
flies. Oecologia (Berlin) 83:371–377. in a lowland river. Odonatologica 34:27–36.
DUFFY, W. G. 1994. Demographics of Lestes disjunctus HOFMANN, T. A., AND C. F. MASON. 2005b. Habitat character-
disjunctus (Odonata, Zygoptera) in a riverine wetland. istics and the distribution of Odonata in a lowland river
Canadian Journal of Zoology 72:910–917. catchment in eastern England. Hydrobiologia 539:137–
EDA, S. 1963. Emergence of Epophlebia superstes Selys. Tombo 147.
5:1–3 (in Japanese) (reviewed by Corbet 1999). JACOBS, M. E. 1955. Studies on territorialism and sexual
ELIAS, J. E., AND M. W. MEYER. 2003. Comparisons of selection in dragonflies. Ecology 36:566–586.
undeveloped and developed shorelands, northern Wis- JENNINGS, M. J., E. E. EMMONS, G. R. HATZENBELER, C. EDWARDS,
consin, and recommendations for restoration. Wetlands AND M. A. BOZEK. 2003. Is littoral habitat affected by
23:800–816. residential development and land use in watersheds of
ELKIN, C. M., AND R. BAKER. 2000. Lack of preference for low- Wisconsin lakes? Lake and Reservoir Management 19:
predation-risk habitats in larval damselflies explained by 272–279.
costs of intraspecific interactions. Animal Behaviour 60: JOHANSSON, F. 1991. Foraging modes in an assemblage of
511–521. odonate larvae—effects of prey and interference. Hydro-
FICETOLA, G. F., E. PADOA-SCHIOPPA, A. MONTI, R. MASSA, F. DE biologia 209:79–87.
BERNARDI, AND L. BOTTONI. 2004. The importance of JOHANSSON, F., AND T. BRODIN. 2003. Effects of fish predators

W:/xml/jnabs/jnbs_2801/jnbs-28-01-05.3d  Wednesday, 12 November 2008  10:35 am  Allen Press, Inc.  Page 53

54 A. J. REMSBURG AND M. G. TURNER [Volume 28

and abiotic factors on dragonfly community structure. MCKINNON, B., AND M. L. MAY. 1994. Mating habitat choice
Journal of Freshwater Ecology 18:415–423. and reproductive success of Pachydiplax longipennis
JOHNSON, D., AND P. CROWLEY. 1980. Habitat and seasonal (Burmeister) (Anisoptera: Libellulidae). Advances in
segregation among coexisting odonate larvae. Odonato- Odonatology 6:59–77.
logica 9:297–308. MCPEEK, M. A. 1990. Determination of species composition in
JOHNSON, D. M., T. H. MARTIN, M. MAHATO, L. B. CROWDER, AND the Enallagma damselfly assemblages of permanent lakes.
P. H. CROWLEY. 1995. Predation, density dependence, and Ecology 71:83–98.
life histories of dragonflies: a field experiment in a MICHALETZ, P. H., K. E. DOISY, AND C. F. RABENI. 2005.
freshwater community. Journal of the North American Influences of productivity, vegetation, and fish on
Benthological Society 14:547–562. macroinvertebrate abundance and size in Midwestern
JOHNSON, D. M., C. L. PIERCE, T. H. MARTIN, C. N. WATSON, R. E. USA impoundments. Hydrobiologia 543:147–157.
BOHANAN, AND P. H. CROWLEY. 1987. Prey depletion by MILNER, A. M., AND I. T. GLOYNE-PHILLIPS. 2005. The role of
odonate larvae—combining evidence from multiple field riparian vegetation and woody debris in the develop-
experiments. Ecology 68:1459–1465. ment of macroinvertebrate assemblages in streams. River
KARR, J. R., AND R. R. ROTH. 1971. Vegetation structure and Research and Applications 21:403–420.
avian diversity in several New World areas. American MUNTIFERING, J. R., A. J. DICKMAN, L. M. PERLOW, T. HRUSKA, P.
Naturalist 105:423–435. G. RYAN, L. L. MARKER, AND R. M. JEO. 2006. Managing the
KNEIB, R. T., AND C. E. H. SCHEELE. 2000. Does tethering of matrix for large carnivores: a novel approach and
mobile prey measure relative predation potential? An perspective from cheetah (Acinonyx jubatus) habitat
empirical test using mummichogs and grass shrimp. suitability modelling. Animal Conservation 9:103–112.
Marine Ecology Progress Series 198:181–190. NAKANO, S., H. MIYASAKA, AND N. KUHARA. 1999. Terrestrial–
KNIGHT, T. M., M. W. MCCOY, J. M. CHASE, K. A. MCCOY, AND R. aquatic linkages: riparian arthropod inputs alter trophic
D. HOLT. 2005. Trophic cascades across ecosystems. cascades in a stream food web. Ecology 80:2435–2441.
Nature 437:880–883. NEEDHAM, J. G., M. J. WESTFALL, AND M. L. MAY. 2000.
LANGELLOTTO, G. A., AND R. F. DENNO. 2004. Responses of Dragonflies of North America. Scientific Publishers,
invertebrate natural enemies to complex-structured Gainesville, Florida.
habitats: a meta-analytical synthesis. Oecologia (Berlin) ORIANS, G. H., AND H. S. HORN. 1969. Overlap in foods and
139:1–10. foraging of four species of blackbirds in the potholes of
LINDSAY, A. R., S. S. GILLUM, AND M. W. MEYER. 2002. Influence of central Washington. Ecology 50:930–938.
lakeshore development on breeding bird communities in a OSBORN, R. 2005. Odonata as indicators of habitat quality at
mixed northern forest. Biological Conservation 107:1–11. Lakes in Louisiana, United States. Odonatologica 34:259–
LOGAN, E. R. 1971. A comparative ecological and behavioral 270.
study of two species of damselflies, Enallagma boreale PEZALLA, V. M. 1979. Behavioral ecology of the dragonfly
(Selys) and Enallagma carunculatum Morse (Odonata: Libellula pulchella Drury (Odonata:Anisoptera). American
Coenagrionidae). PhD Thesis, Washington State Uni- Midland Naturalist 102:1–22.
versity, Pullman, Washington. PIANKA, E. R. 1967. Lizard species diversity. Ecology 48:333–
MACARTHUR, R. H. 1965. Patterns of species diversity. 351.
Biological Reviews 40:510–533. RACEY, G. D., AND D. L. EULER. 1982. Small mammal and
MARBURG, A. E., M. G. TURNER, AND T. K. KRATZ. 2006. Natural habitat response to shoreline cottage development in
and anthropogenic variation in coarse wood among and central Ontario. Canadian Journal of Zoology 60:865–
within lakes. Journal of Ecology 94:558–568. 880.
MARNELL, F. 1998. Discriminant analysis of the terrestrial and RADOMSKI, P., AND T. J. GOEMAN. 2001. Consequences of human
aquatic habitat determinants of the smooth newt lakeshore development on emergent and floating-leaf
(Triturus vulgaris) and the common frog (Rana temporaria) vegetation abundance. North American Journal of
in Ireland. Journal of Zoology 244:1–6. Fisheries Management 21:46–61.
MARTIN, L. 1965. The physical geography of Wisconsin. 3rd REHFELDT, G. E. 1990. Anti-predator strategies in oviposition
edition. University of Wisconsin Press, Madison, Wis- site selection of Pyrrhosoma nymphula (Zygoptera: Odo-
consin. nata). Oecologia (Berlin) 85:233–237.
MAY, M. L. 1976. Thermoregulation and adaptation to REMSBURG, A. J. 2007. Aquatic and terrestrial vegetation
temperature in dragonflies (Odonata: Anisoptera). Eco- influence lacustrine dragonfly (order Odonata) assem-
logical Monographs 46:1–32. blages at multiple life stages. PhD Thesis, University of
MAY, M. L., AND J. M. BAIRD. 2002. A comparison of foraging Wisconsin, Madison, Wisconsin.
behavior in two ‘‘percher’’ dragonflies, Pachydiplax long- ROTH, B. M., I. C. KAPLAN, G. G. SASS, P. T. JOHNSON, A. E.
ipennis and Erythemis simplicicollis (Odonata: Libelluli- MARBURG, A. C. YANNARELL, T. D. HAVLICEK, T. V. WILLIS, M.
dae). Journal of Insect Behavior 15:765–778. G. TURNER, AND S. R. CARPENTER. 2007. Linking terrestrial
MCCOY, E. D., AND S. S. BELL. 1991. Habitat structure. Pages 3– and aquatic ecosystems: the role of woody habitat in lake
27 in S. S. Bell, E. D. McCoy, and H. R. Mushinsky food webs. Ecological Modelling 203:439–452.
(editors). Population and community biology series. ROUQUETTE, J. R., AND D. J. THOMPSON. 2007. Roosting site
Chapman and Hall, New York. selection in the endangered damselfly, Coenagrion mer-

W:/xml/jnabs/jnbs_2801/jnbs-28-01-05.3d  Wednesday, 12 November 2008  10:35 am  Allen Press, Inc.  Page 54

2009] AQUATIC AND RIPARIAN PLANTS INFLUENCE DRAGONFLIES 55

curiale, and implications for habitat design. Journal of VAN BUSKIRK, J. 1988. Interactive effects of dragonfly
Insect Conservation 11:187–193. predation in experimental pond communities. Ecology
SASS, G. G. 2004. Fish community and food web responses to 69:857–867.
a whole-lake removal of coarse woody habitat. PhD WALLACE, J. B., S. L. EGGERT, J. L. MEYER, AND J. R. WEBSTER.
Thesis, University of Wisconsin, Madison, Madison, 1997. Multiple trophic levels of a forest stream linked to
Wisconsin. terrestrial litter inputs. Science 277:102–104.
SCHAFFNER, A. K., AND B. R. ANHOLT. 1998. Influence of WEATHERHEAD, M. A., AND M. R. JAMES. 2001. Distribution of
predator presence and prey density on behavior and macroinvertebrates in relation to physical and biological
growth of damselfly larvae (Ischnura elegans) (Odonata: variables in the littoral zone of nine New Zealand lakes.
Zygoptera). Journal of Insect Behavior 11:793–809. Hydrobiologia 462:115–129.
SCHINDLER, D. E., S. I. GEIB, AND M. R. WILLIAMS. 2000. Patterns WEAVER, M. J., J. J. MAGNUSON, AND M. K. CLAYTON. 1997.
of fish growth along a residential development gradient Distribution of littoral fishes in structurally complex
in north temperate lakes. Ecosystems 3:229–237. macrophytes. Canadian Journal of Fisheries and Aquatic
SCHMUDE, K. L. 1998. Effects of habitat complexity on Sciences 54:2277–2289.
macroinvertebrate colonization of artificial substrates in
WERNER, E. E., J. F. GILLIAM, D. J. HALL, AND G. G. MITTELBACH.
north temperate lakes. Journal of the North American
1983. An experimental test of the effects of predation risk
Benthological Society 17:73–80.
on habitat use in fish. Ecology 64:1540–1548.
SCHNAIBERG, J., J. RIERA, M. G. TURNER, AND P. R. VOSS. 2002.
WESTFALL, M. J., AND M. L. MAY. 1996. Damselflies of North
Explaining human settlement patterns in a recreational
America. Scientific Publishers, Gainesville, Florida.
lake district: Vilas County, Wisconsin, USA. Environ-
mental Management 30:24–34. WHITAKER, D. M., A. L. CARROLL, AND W. A. MONTEVECCHI.
SCRIBNER, K. T., J. W. ARNTZEN, N. CRUDDACE, R. S. OLDHAM, 2000. Elevated numbers of flying insects and insectivo-
AND T. BURKE. 2001. Environmental correlates of toad
rous birds in riparian buffer strips. Canadian Journal of
abundance and population genetic diversity. Biological Zoology 78:740–747.
Conservation 98:201–210. WILDERMUTH, H. 1992. Habitate und Habitatwahl der grossen
STORCH, D., AND D. FRYNTA. 1999. Evolution of habitat Moosjungfer (Leucorrhinia pectoralis) Charp. 1825 (Odo-
selection: stochastic acquisition of cognitive clues? nata, Libellulidae). Zeitschrift fur Okologie und Natur-
Evolutionary Ecology 13:591–600. schutz 1:3–21.
SUGIURA, N. 1978. Further analysis of the data by Akaike’s WILDERMUTH, H. 1993. Habitat selection and oviposition site
information criterion and the finite corrections. Commu- recognition by the dragonfly Aeshna juncea (L.): an
nications in Statistics: Theory and Methods 7:13–26. experimental approach in natural habitats (Anisoptera:
SWITZER, P. V., AND W. WALTERS. 1999. Choice of lookout posts Aeshnidae). Odonatologica 22:27–44.
by territorial amberwing dragonflies, Perithemis tenera WILLIS, T. V., AND J. J. MAGNUSON. 2000. Patterns in fish species
(Anisoptera: Libellulidae). Journal of Insect Behavior 12: composition across the interface between streams and
385–398. lakes. Canadian Journal of Fisheries and Aquatic
THOMPSON, D. J. 1987. Regulation of damselfly populations: Sciences 57:1042–1052.
the effects of weed density on larval mortality due to WILSON, K. A., J. J. MAGNUSON, D. M. LODGE, A. M. HILL, T. K.
predation. Freshwater Biology 17:367–371. KRATZ, W. L. PERRY, AND T. V. WILLIS. 2004. A long-term
THORP, J., AND M. COTHRAN. 1984. Regulation of freshwater rusty crayfish (Orconectes rusticus) invasion: dispersal
community structure at multiple intensities of dragonfly patterns and community change in a north temperate
predation. Ecology 65:1546–1555. lake. Canadian Journal of Fisheries and Aquatic Sciences
TSCHARNTKE, T., T. A. RAND, AND F. BIANCHI. 2005. The 61:2255–2266.
landscape context of trophic interactions: insect spillover WISSINGER, S. A. 1988. Spatial distribution, life history and
across the crop–noncrop interface. Annales Zoologici estimates of survivorship in a fourteen-species assem-
Fennici 42:421–432. blage of larval dragonflies (Odonata: Anisoptera).
TUPPER, M., AND R. G. BOUTILIER. 1995. Effects of habitat on Freshwater Biology 20:329–340.
settlement, growth, and postsettlement survival of WOLF, L. L., AND E. C. WALTZ. 1988. Oviposition site selection
Atlantic cod (Gadus morhua). Canadian Journal of
and spatial predictability of female white-faced dragon-
Fisheries and Aquatic Sciences 52:1834–1841.
flies (Leucorrhinia intacta) (Odonata: Libellulidae). Ethol-
UBUKATA, H. 1984. Oviposition site selection and avoidance of
ogy 78:306–320.
additional mating by females of the dragonfly, Cordulia
WOODFORD, J. E., AND M. W. MEYER. 2003. Impact of lakeshore
aenea amurensis Selys (Corduliidae). Researches on
development on green frog abundance. Biological
Population Ecology 26:285–301.
Conservation 110:277–284.
VAN BUSKIRK, J. 1986. Establishment and organization of
territories in the dragonfly Sympetrum rubicundulum Received: 8 January 2008
(Odonata: Libellulidae). Animal Behaviour 34:1781–1790. Accepted: 4 August 2008

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56 A. J. REMSBURG AND M. G. TURNER [Volume 28

APPENDIX. Odonata species collected as larvae from 59 littoral sites during a broad-scale survey of 41 sites (12 m2 of littoral
benthos sampled per site) in 2004 and a fine-scale survey of 18 microsites (½ with macrophytes, ½ without macrophytes; 2.4 m2 of
littoral benthos sampled per microsite) in northern Wisconsin lakes in 2006. Behavioral guild classifications are shown for larval and
adult stages. Three Libellulinae species (Leucorrhinia hudsonica, Leucorrhinia proxima, and Tramea carolina) were observed only in 2006;
all other species were found in both years.

Larval behavior Adult behavior

Taxon Clasper Sprawler Burrower Flier Percher
Aeshnidae
Basiaeschna janata X X
Boyeria vinosa X X
Gomphidae
Arigomphus furcifer X X
Dromogomphus spinosus X
Gomphus spicatus X X
Gomphus exilis X X
Gomphus lividus X X
Gomphurus fraternus X X
Hagenius brevistylus X X
Libellulinae
Celithemis elisa X X
Ladona julia X X X
Leucorrhinia hudsonica X X
Leucorrhinia proxima X X
Somatachlora cingulata X X
Tramea carolina X X
Cordulinae
Cordulia shurtleffIi X X
Drorocordulia libera X X
Epitheca cynosura X X
Epitheca princeps X X
Epitheca spinigera X X
Macrominae
Didymops transversa X X X
Macromia illinoiensis X X X
Coenagrionidae spp. X X

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