Unravelling the correlates of species richness and ecological uniqueness in a metacommunity of urban pond insects

City ponds have the potential to harbour a rich biodiversity of aquatic insects despite being located in an urban landscape. However, our current knowledge on the correlates of pond biodiversity is limited and even less is known about the factors that influence the ecological uniqueness of urban ponds. The multiple environmental gradients, at different spatial scales, that may affect biodiversity and ecological uniqueness of urban ponds can thus be seen both as an opportunity and as a challenge for a study. In this study, we aimed to fill this gap by focusing on aquatic insect assemblages in 51 ponds in the Swedish city of Stockholm, using a metacommunity perspective. We found that species richness was primarily determined by the density of aquatic insects, water depth and proportion of buildings around the pond. The uniqueness of ponds was estimated as local contributions to beta diversity (LCBD), and it was primarily related to the proportion of arable land and industry around the ponds. With regard to the metacommunity we found two interesting patterns. First, there was a negative relationship between richness and LCBD. Second, biodiversity was spatially independent, suggesting that spatially-patterned dispersal did not structure species richness or LCBD. These last two patterns are important when considering conservation efforts of biodiversity in city ponds. We hence suggest that the conservation of insect biodiversity in urban pond should consider the surroundings of the ponds, and that high-richness ponds are not necessarily those that require most attention because they are not ecologically the most unique.     

The response of amphibian communities to fish and habitat features in Mediterranean permanent ponds

Generalized linear models were used to test the effect of fish, using ponds with and without fish and habitat features as covariates, on richness and abundance of amphibian species. Five fish species and six amphibian species were recorded in 60 permanent ponds located in central Italy. The choice of covariates (macrophyte cover and pond surface area) was made after studying the correlations. The richness of amphibian species was not significantly affected by fish presence or macrophyte cover, in line with previous studies, since almost all the fish species were non-predatory. However, abundance of urodeles (newts) was negatively affected by fish and positively affected by macrophyte cover. Although fish may strongly influence the abundance and composition of amphibian communities, the results indicate that the cover of aquatic macrophytes may increase the available habitat for amphibians and therefore their abundance. Anuran species preferred ponds where fish were present, since both groups preferred larger ponds. Concordance between fish and amphibian species composition was not found by the Mantel and Partial Mantel tests. This indicates that the fish assemblages do not predict which amphibian species occur in the pond.     

Evaluating the role of regional and local processes in structuring a larval trematode metacommunity of Helisoma trivolvis

Metacommunity theory has advanced our understanding of how local and regional processes affect the structure of ecological communities. While parasites have largely been omitted from metacommunity research, parasite communities can provide the large sample sizes and discrete boundaries often required for evaluating metacommunity patterns. Here, we used assemblages of flatworm parasites that infect freshwater snails (Helisoma trivolvis) to evaluate three questions: 1) what factors affect individual host infections within ponds? 2) Is the parasite metacommunity structured among ponds? And 3) what is the relative role of local versus regional processes in determining metacommunity structure and species richness among ponds? We examined 10 821 snails from 96 sites in five park complexes in the San Francisco Bay area, California, and found 953 infections from six parasite groups. At the within‐pond level, infection status of host snails correlated positively with individual snail size and pond infection prevalence for all six parasite groups. Using an ordination method to test for metacommunity structure, we found that the parasite metacommunity was organized in a non‐random pattern with species responding individually along an environmental gradient. Based on a model selection approach involving local and regional predictors, parasite species richness and metacommunity structure correlated with both local abiotic (pH and total dissolved nitrogen) and biotic (non‐host mollusk density, and H. trivolvis biomass) factors, with little support for regional predictors. Overall, this trematode metacommunity most closely followed the predictions from the species sorting or mass effects metacommunity paradigm, in which community diversity is filtered by local site characteristics.     

Determinant landscape‐scale factors on pond odonate assemblages

相似文献   

An expert-based landscape permeability model for assessing the impact of agricultural management on amphibian migration

The persistence of pond-breeding amphibian populations is influenced on different spatial scales ranging from the individual breeding pond to surrounding habitat patches to landscape clusters of breeding pond populations. The connectivity among breeding ponds as well as the availability of suitable terrestrial habitats surrounding the ponds plays a major role in long-term viability of amphibian species. Besides road traffic and urban structures agricultural land activity can disrupt landscape connectivity through the use of pesticides, fertilizers and physical activity such as tillage. We developed an expert-based model to assess the impact of agricultural management measures on the migration area and terrestrial habitat availability for seven amphibian species. The model is based on a Habitat Suitability Index to identify suitable terrestrial habitats and includes a landscape permeability approach. Size of migration areas, connectivity of breeding ponds and number of reachable terrestrial habitats were modeled considering species-specific migration ranges and habitat preferences. We consider how pesticides application might lead to fragmentation and isolation of amphibian breeding pond populations. Therefore the potential disrupting impact of pesticides was simulated by inflating landscape resistance to medium and high migration cost. One amphibian species showed a decrease of migration area by 48.3% and a decrease of reachable terrestrial habitats by 41.5% at high migration costs. Three additional species showed a decrease of their migration areas between 31.5 and 35.7%. At increased migration cost, some of the investigated populations were isolated at breeding pond level or restricted to pond clusters. Our model could be used to prioritize conservation efforts for pond-breeding amphibians with adequate consideration of agricultural land use and its impact on amphibian migration.     

Urban ponds as an aquatic biodiversity resource in modified landscapes

Urbanization is a global process contributing to the loss and fragmentation of natural habitats. Many studies have focused on the biological response of terrestrial taxa and habitats to urbanization. However, little is known regarding the consequences of urbanization on freshwater habitats, especially small lentic systems. In this study, we examined aquatic macro‐invertebrate diversity (family and species level) and variation in community composition between 240 urban and 782 nonurban ponds distributed across the United Kingdom. Contrary to predictions, urban ponds supported similar numbers of invertebrate species and families compared to nonurban ponds. Similar gamma diversity was found between the two groups at both family and species taxonomic levels. The biological communities of urban ponds were markedly different to those of nonurban ponds, and the variability in urban pond community composition was greater than that in nonurban ponds, contrary to previous work showing homogenization of communities in urban areas. Positive spatial autocorrelation was recorded for urban and nonurban ponds at 0–50 km (distance between pond study sites) and negative spatial autocorrelation was observed at 100–150 km and was stronger in urban ponds in both cases. Ponds do not follow the same ecological patterns as terrestrial and lotic habitats (reduced taxonomic richness) in urban environments; in contrast, they support high taxonomic richness and contribute significantly to regional faunal diversity. Individual cities are complex structural mosaics which evolve over long periods of time and are managed in diverse ways. This facilitates the development of a wide range of environmental conditions and habitat niches in urban ponds which can promote greater heterogeneity between pond communities at larger scales. Ponds provide an opportunity for managers and environmental regulators to conserve and enhance freshwater biodiversity in urbanized landscapes whilst also facilitating key ecosystem services including storm water storage and water treatment.     

Promoting dragonfly diversity in cities: major determinants and implications for urban pond design

Urbanisation is increasing and it is essential to integrate biodiversity into the spatial planning of urban areas. This requires deeper understanding of biodiversity patterns in cities. We investigated which habitat variables are major determinants of dragonfly diversity and species assemblage structure in the municipal area of Dortmund (Germany). We sampled dragonfly larvae in 33 ponds situated in city parks, commercial, residential and agricultural areas. We recorded 30 autochthonous dragonfly species with species richness ranging from zero to 17. Additionally, we surveyed a set of environmental variables including habitat size, water level, pond structures and vegetation as well as surrounding landscape and potential disturbances like waterfowl and fish. Multivariate methods were used to identify the major determinants of dragonfly diversity, abundance and assemblage structure. Analysis indicated that diversity of aquatic and terrestrial vegetation affected dragonfly diversity positively. City park ponds had low diversity, but Ischnura elegans was obviously promoted by the specific park pond conditions, including high waterfowl density. We found five assemblages mostly determined by generalistic species which were related to different pond types. Moderately disturbed ruderal and pioneer ponds in residential and agricultural areas also contained increased numbers of rare species. Our results indicate that urban ponds may have a great value for maintaining biodiversity, but various disturbances have negative impact. To promote urban biodiversity we suggest a natural design of well-vegetated ponds as well as a high diversity of different pond types and particularly a more-natural redesign of city park ponds.     

Fish assemblages in temporary ponds adjacent to 'terra-firme' streams in Central Amazonia

1. The effect of habitat structural features and physicochemical characteristics of the water on the composition and richness of fish assemblages in temporary ponds near streams were examined at three spatial scales: among ponds, among streams and between drainage basins, in a ‘terra‐firme’ (not subject to long‐term flooding) forest reserve in Central Amazonia. 2. The fish assemblage in temporary ponds was composed of subsets of 18 small‐bodied species widely distributed in the reserve. The assemblages had a nested subset structure, where smaller ponds contained subgroups of the species found in larger ponds. 3. Species composition and richness in temporary ponds were similar between drainage basins, although the fish assemblages in streams differed between basins. 4. Fish assemblage structure was influenced by local factors related to habitat structure, such as pond area and depth, canopy cover and hydroperiod. Physicochemical characteristics of the water in the ponds were similar between drainage basins and had little detectable effect on the structure of pond fish assemblages. 5. No correspondence was found between the composition, richness or abundance of fishes in the ponds and in stretches of the streams adjacent to the ponds. Therefore, it is not possible to predict the composition of these temporary pond fish assemblages from the fish assemblages found in adjacent streams.     

The effect of fish and aquatic habitat complexity on amphibians

Fish introductions are considered one of the most widespread anthropogenic threats to aquatic ecosystems. Their negative impact on native amphibian communities has received increasing attention in recent years. We investigated the relationship between the introduced fish, emergent vegetation cover and native amphibians in man-made ponds generated by regulation and dam building along the Tarnava Mare Valley (Romania) during the last 40 years. We inventoried amphibians and fish inhabiting 85 permanent ponds and estimated habitat complexity focusing on emergent vegetation cover. Four amphibian species were found to be negatively associated with the presence of predatory fish. Species richness of ponds without fish and ponds without predatory fish did not differ significantly, whereas ponds containing only predatory fish had significantly lower amphibian richness. A significant positive relationship was found between the emergent vegetation cover and pond occupancy of six amphibian species and amphibian species richness. As a management recommendation, we suggest the restriction of fish introductions to non predatory fish and the maintenance of high emergent vegetation cover in the ponds. Handling editor: S. Declerck     

Fish species richness in spring‐fed ponds: effects of habitat size versus isolation in temporally variable environments

1. Species richness in a habitat patch is determined by immigration (regional) and extinction (local) processes, and understanding their relative importance is crucial for conservation of biodiversity. In this study, we applied the Island Biogeography concept to spring ponds connected to a river in southwestern Japan to examine how immigration and extinction processes interact to determine fish species richness in temporally variable environments. 2. Fish censuses were conducted 15 times in 13 study ponds at 1–4 month intervals from August 1998 through October 2000. Effects of habitat size (pond area), isolation (distance from the river) and temporal environmental variability (water level fluctuation) on (i) species richness, (ii) immigration and extinction rates and (iii) population size and persistence of each fish species were assessed. 3. The results revealed predominant effects of distance on species richness, immigration/extinction rates and population size and persistence. Species richness decreased with increasing distance but was not related to either pond area or water level fluctuation. A negative effect of distance on immigration rate was detected, while neither pond area nor water level fluctuation had significant effects on extinction rate. Further, population size and persistence of four species increased with decreasing distance, suggesting that, in ponds close to the river, immigrants from the river reduce the probability of extinction (i.e. provide a rescue effect), contributing to the maintenance of high species richness. 4. Overall results emphasise the importance of immigration processes, rather than extinction, in shaping patterns of species richness in our system. The predominant importance of immigration was probably because of (i) high temporal variability that negates habitat‐size effects and (ii) continuous immigration that easily compensates for local extinctions. Our results suggest that consideration of regional factors (e.g. connectivity, locations of source populations and barriers to colonisation) is crucial for conservation and restoration of local habitats.     

Anuran tadpole assemblages in riparian areas of the Middle Paraná River, Argentina

We investigated habitat variables influencing the species richness and composition of anuran tadpole assemblages in ponds from riparian areas of the Middle Paraná River. Tadpoles were collected at seven ponds that differed in hydroperiod and 10 habitat variables (water temperature, pH, conductivity, dissolved oxygen, turbidity, depth, size, time of isolation, macrophyte richness, and level of natural or anthropic disturbances) during a 5 year period. The presence/absence of potential tadpole predators was also recorded. Although we did not examine tadpole diets, we categorized anurans into ecomorphological guilds. We found that dissolved oxygen, maximum depth, pond size, macrophyte richness, and disturbances were significantly related to tadpole species richness. We also found that the richness of potential tadpole predator increased with longer pond hydroperiod. Thus, the number of tadpole species was higher in temporary and semi-permanent ponds without potential tadpole predators. The results of our study provide the first quantitative data on the tadpole assemblages in ponds associated with riparian areas of the Middle Paraná River Floodplain based on multi-year intensive sampling, a finding of both ecological interest and practical significance for future conservation management of anurans of Paraná River ecosystems.     

General relationships between consumer dispersal,resource dispersal and metacommunity diversity

One of the central questions of metacommunity theory is how dispersal of organisms affects species diversity. Here, we show that the diversity–dispersal relationship should not be studied in isolation of other abiotic and biotic flows in the metacommunity. We study a mechanistic metacommunity model in which consumer species compete for an abiotic or biotic resource. We consider both consumer species specialised to a habitat patch, and generalist species capable of using the resource throughout the metacommunity. We present analytical results for different limiting values of consumer dispersal and resource dispersal, and complement these results with simulations for intermediate dispersal values. Our analysis reveals generic patterns for the combined effects of consumer and resource dispersal on the metacommunity diversity of consumer species, and shows that hump‐shaped relationships between local diversity and dispersal are not universal. Diversity–dispersal relationships can also be monotonically increasing or multimodal. Our work is a new step towards a general theory of metacommunity diversity integrating dispersal at multiple trophic levels.     

Direct and indirect effects of predation on tadpole community structure in the Amazon rainforest

Abstract The relationship between the distribution of predators (fish, odonates and water beetles) and prey assemblages (amphibian larvae) was investigated in the tropical rainforest of central Amazonas, Brasil. The anuran community uses a variety of waterbodies for reproduction, ranging from streams and streamside ponds to isolated forest ponds. Predators in this system include fish in streams and streamside ponds, and invertebrates (primarily odonate naiads and beetles) in forest ponds. Tadpole species richness and assemblage structure were related to fish density and species richness. No relationships between tadpole assemblages and abiotic pond characteristics were detected. The presence offish explained much of the variation in both species composition and species richness within and among ponds. Some species of tadpole were consistently found to coexist with high densities of fish. Path analyses suggest that while fish have a strong direct effect on tadpole associations and species richness, they also have an indirect effect through invertebrate predators (odonate larvae and coleopteran beetles). Prey survival-strategies such as palatability and behaviour may explain how tadpole species composition is affected by predators at the community level. These findings suggest that the observed patterns of habitat use by larval anurans may be structured in response to the distribution of key predators (fish) in this system.     

Urbanization Level and Woodland Size Are Major Drivers of Woodpecker Species Richness and Abundance

Urbanization is a process globally responsible for loss of biodiversity and for biological homogenization. Urbanization may have a direct negative impact on species behaviour and indirect effects on species populations through alterations of their habitats, for example patch size and habitat quality. Woodpeckers are species potentially susceptible to urbanization. These birds are mostly forest specialists and the development of urban areas in former forests may be an important factor influencing their richness and abundance, but documented examples are rare. In this study we investigated how woodpeckers responded to changes in forest habitats as a consequence of urbanization, namely size and isolation of habitat patches, and other within-patch characteristics. We selected 42 woodland patches in a gradient from a semi-natural rural landscape to the city centre of Poznań (Western Poland) in spring 2010. Both species richness and abundance of woodpeckers correlated positively to woodland patch area and negatively to increasing urbanization. Abundance of woodpeckers was also positively correlated with shrub cover and percentage of deciduous tree species. Furthermore, species richness and abundance of woodpeckers were highest at moderate values of canopy openness. Ordination analyses confirmed that urbanization level and woodland patch area were variables contributing most to species abundance in the woodpecker community. Similar results were obtained in presence-absence models for particular species. Thus, to sustain woodpecker species within cities it is important to keep woodland patches large, multi-layered and rich in deciduous tree species.     

Dispersal and the metapopulation paradigm in amphibian ecology and conservation: are all amphibian populations metapopulations?

Amphibians are frequently characterized as having limited dispersal abilities, strong site fidelity and spatially disjunct breeding habitat. As such, pond‐breeding species are often alleged to form metapopulations. Amphibian species worldwide appear to be suffering population level declines caused, at least in part, by the degradation and fragmentation of habitat and the intervening areas between habitat patches. If the simplification of amphibians occupying metapopulations is accurate, then a regionally based conservation strategy, informed by metapopulation theory, is a powerful tool to estimate the isolation and extinction risk of ponds or populations. However, to date no attempt to assess the class‐wide generalization of amphibian populations as metapopulations has been made. We reviewed the literature on amphibians as metapopulations (53 journal articles or theses) and amphibian dispersal (166 journal articles or theses for 53 anuran species and 37 salamander species) to evaluate whether the conditions for metapopulation structure had been tested, whether pond isolation was based only on the assumption of limited dispersal, and whether amphibian dispersal was uniformly limited. We found that in the majority of cases (74%) the assumptions of the metapopulation paradigm were not tested. Breeding patch isolation via limited dispersal and/or strong site fidelity was the most frequently implicated or tested metapopulation condition, however we found strong evidence that amphibian dispersal is not as uniformly limited as is often thought. The frequency distribution of maximum movements for anurans and salamanders was well described by an inverse power law. This relationship predicts that distances beneath 11–13 and 8–9 km, respectively, are in a range that they may receive one emigrating individual. Populations isolated by distances approaching this range are perhaps more likely to exhibit metapopulation structure than less isolated populations. Those studies that covered larger areas also tended to report longer maximum movement distances – a pattern with implications for the design of mark‐recapture studies. Caution should be exercised in the application of the metapopulation approach to amphibian population conservation. Some amphibian populations are structured as metapopulations – but not all.     

Intermediate Pond Sizes Contain the Highest Density,Richness, and Diversity of Pond-Breeding Amphibians

We present data on amphibian density, species richness, and diversity from a 7140-ha area consisting of 200 ponds in the Midwestern U.S. that represents most of the possible lentic aquatic breeding habitats common in this region. Our study includes all possible breeding sites with natural and anthropogenic disturbance processes that can be missing from studies where sampling intensity is low, sample area is small, or partial disturbance gradients are sampled. We tested whether pond area was a significant predictor of density, species richness, and diversity of amphibians and if values peaked at intermediate pond areas. We found that in all cases a quadratic model fit our data significantly better than a linear model. Because small ponds have a high probability of pond drying and large ponds have a high probability of fish colonization and accumulation of invertebrate predators, drying and predation may be two mechanisms driving the peak of density and diversity towards intermediate values of pond size. We also found that not all intermediate sized ponds produced many larvae; in fact, some had low amphibian density, richness, and diversity. Further analyses of the subset of ponds represented in the peak of the area distribution showed that fish, hydroperiod, invertebrate density, and canopy are additional factors that drive density, richness and diversity of ponds up or down, when extremely small or large ponds are eliminated. Our results indicate that fishless ponds at intermediate sizes are more diverse, produce more larvae, and have greater potential to recruit juveniles into adult populations of most species sampled. Further, hylid and chorus frogs are found predictably more often in ephemeral ponds whereas bullfrogs, green frogs, and cricket frogs are found most often in permanent ponds with fish. Our data increase understanding of what factors structure and maintain amphibian diversity across large landscapes.     

A landscape ecology approach identifies important drivers of urban biodiversity

Cities are growing rapidly worldwide, yet a mechanistic understanding of the impact of urbanization on biodiversity is lacking. We assessed the impact of urbanization on arthropod diversity (species richness and evenness) and abundance in a study of six cities and nearby intensively managed agricultural areas. Within the urban ecosystem, we disentangled the relative importance of two key landscape factors affecting biodiversity, namely the amount of vegetated area and patch isolation. To do so, we a priori selected sites that independently varied in the amount of vegetated area in the surrounding landscape at the 500‐m scale and patch isolation at the 100‐m scale, and we hold local patch characteristics constant. As indicator groups, we used bugs, beetles, leafhoppers, and spiders. Compared to intensively managed agricultural ecosystems, urban ecosystems supported a higher abundance of most indicator groups, a higher number of bug species, and a lower evenness of bug and beetle species. Within cities, a high amount of vegetated area increased species richness and abundance of most arthropod groups, whereas evenness showed no clear pattern. Patch isolation played only a limited role in urban ecosystems, which contrasts findings from agro‐ecological studies. Our results show that urban areas can harbor a similar arthropod diversity and abundance compared to intensively managed agricultural ecosystems. Further, negative consequences of urbanization on arthropod diversity can be mitigated by providing sufficient vegetated space in the urban area, while patch connectivity is less important in an urban context. This highlights the need for applying a landscape ecological approach to understand the mechanisms shaping urban biodiversity and underlines the potential of appropriate urban planning for mitigating biodiversity loss.     

Local and Landscape Factors Driving the Structure of Tropical Anuran Communities: Do Ephemeral Ponds have a Nested Pattern?

The sets of species in animal and plant communities often comprise nested subsets of the species in broader communities. Although most mechanisms causing nested patterns are known and have been demonstrated for different environments and taxa, how amphibian communities are structured in ephemeral ponds in tropical disturbed landscapes remains unknown. We investigated if pond size, duration, presence of trees (local factors), and the proportion of forest cover surrounding ponds (landscape factor) affect anuran species richness and composition, and if pond assemblages showed a nested pattern. We sampled 11 ephemeral ponds in a pasture matrix near a large Atlantic Forest remnant in Brazil and measured local and landscape variables inside two buffer zones around each pond (100 and 500 m). We marked 1514 individuals from 23 anuran species, and found that richness in ponds ranged from 3–14 species. Both local and landscape factors explained frog species richness in the sampled ponds, and seemed to affect community composition. Frog communities occurred in a non‐nested pattern, contrary to our expectations: species found in poor subsets were not found in larger, more complex ponds. Local and landscape characteristics create a variety of environments in ephemeral ponds, even in impoverished ones; these characteristics restrict pond occupancy for some species, and result in a non‐nested pattern.     

Patch size determines the strength of edge effects on carabid beetle assemblages in urban remnant forests

Habitat fragmentation due to urbanization is increasing rapidly worldwide. Although patch area and edge effect are both important determinants of species diversity and the number of individuals in fragmented landscapes, studies that tested interaction between two effects were limited. Here we examined the interaction between area and edge effects on species richness and the number of individuals of carabids in highly fragmented forests in Tokyo, central Japan. We surveyed carabids in each of 26 forest patches (1.1–121.6 ha) using pitfall traps set in both edge and interior zones. First, we correlated the edge-to-interior differences of both species richness and the number of individuals with patch area. Second, we examined the interaction between patch area and distance to the edge on species richness and the number of individuals using generalized linear models. We found a significant decrease in carabid species richness and the number of individuals in edge zones. The edge-to-interior differences in both species richness and the number of individuals were positively correlated with patch area. Model selection revealed the evident interaction effects between patch area and distance to the edge: higher number of individuals was predicted in only large interior zones. Our results indicated that carabid beetle assemblages were influenced by the interaction between area and edge effects. Thus, in urban areas where small forest remnants dominate, circularizing the shape of the forest patches to maximize the core areas may be the most feasible and realistic means to preserve biodiversity.     

Fine-scale urbanization affects Odonata species diversity in ponds of a megacity (Paris,France)

Current developments in urban ecology include very few studies focused on pond ecosystems, though ponds are recognized as biodiversity hotspots. Using Odonata as an indicator model, we explored changes in species composition in ponds localized along an urban gradient of a megacity (Paris, France). We then assessed the relative importance of local- and landscape-scale variables in shaping Odonata α-diversity patterns using a model-averaging approach. Analyses were performed for adult (A) and adult plus exuviae (AE) census data. At 26 ponds, we recorded 657 adults and 815 exuviae belonging to 17 Odonata species. The results showed that the Odonata species assemblage composition was not determined by pond localization along the urban gradient. Similarly, pond characteristics were found to be similar among urban, suburban and periurban ponds. The analyses of AE census data revealed that fine-scale urbanization (i.e., increased density of buildings surrounding ponds) negatively affects Odonata α-diversity. In contrast, pond localization along the urban gradient weakly explained the α-diversity patterns. Several local-scale variables, such as the coverage of submerged macrophytes, were found to be significant drivers of Odonata α-diversity. Together, these results show that the degree of urbanization around ponds must be considered instead of pond localization along the urban gradient when assessing the potential impacts of urbanization on Odonata species diversity. This work also indicates the importance of exuviae sampling in understanding the response of Odonata to urbanization.