Habitat and landscape characteristics underlying anuran community structure along an urban-rural gradient

Urbanization has been cited as an important factor in worldwide amphibian declines, and although recent work has illustrated the important influence of broad-scale ecological patterns and processes on amphibian populations, little is known about the factors structuring amphibian communities in urban landscapes. We therefore examined amphibian community responses to wetland habitat availability and landscape characteristics along an urban-rural gradient in central Iowa, USA, a region experiencing rapid suburban growth. We conducted call surveys at 61 wetlands to estimate anuran calling activity, and quantified wetland habitat structure and landscape context. We used canonical correspondence analysis (CCA) to examine patterns in anuran community structure and identify the most important variables associated with those patterns. Urban density at the landscape scale had a significant negative influence on overall anuran abundance and diversity. While every species exhibited a decrease in abundance with increasing urban density, this pattern was especially pronounced for species requiring post-breeding upland habitats. Anurans most affected by urbanization were those associated with short hydroperiods, early breeding activity, and substantial upland habitat use. We suggest that broad-scale landscape fragmentation is an important factor underlying anuran community structure in this region, possibly due to limitations on the accessibility of otherwise suitable habitat in fragmented urban landscapes. This study underscores the importance of a regional approach to amphibian conservation in urban and urbanizing areas; in fragmented landscapes, a network of interconnected wetland and upland habitats may be more likely to support a successful, diverse anuran community than will isolated sites.     

Remarkable Amphibian Biomass and Abundance in an Isolated Wetland: Implications for Wetland Conservation

Abstract:  Despite the continuing loss of wetland habitats and associated declines in amphibian populations, attempts to translate wetland losses into measurable losses to ecosystems have been lacking. We estimated the potential productivity from the amphibian community that would be compromised by the loss of a single isolated wetland that has been protected from most industrial, agricultural, and urban impacts for the past 54 years. We used a continuous drift fence at Ellenton Bay, a 10-ha freshwater wetland on the Savannah River Site, near Aiken, South Carolina (U.S.A.), to sample all amphibians for 1 year following a prolonged drought. Despite intensive agricultural use of the land surrounding Ellenton Bay prior to 1951, we documented 24 species and remarkably high numbers and biomass of juvenile amphibians (>360,000 individuals; >1,400 kg) produced during one breeding season. Anurans (17 species) were more abundant than salamanders (7 species), comprising 96.4% of individual captures. Most (95.9%) of the amphibian biomass came from 232095 individuals of a single species of anuran (southern leopard frog [Rana sphenocephala ]). Our results revealed the resilience of an amphibian community to natural stressors and historical habitat alteration and the potential magnitude of biomass and energy transfer from isolated wetlands to surrounding terrestrial habitat. We attributed the postdrought success of amphibians to a combination of adult longevity (often >5 years), a reduction in predator abundance, and an abundance of larval food resources. Likewise, the increase of forest cover around Ellenton Bay from <20% in 1951 to >60% in 2001 probably contributed to the long-term persistence of amphibians at this site. Our findings provide an optimistic counterpoint to the issue of the global decline of biological diversity by demonstrating that conservation efforts can mitigate historical habitat degradation.     

Amphibian occurrence is influenced by current and historic landscape characteristics.

Human-induced habitat loss and degradation are major threats to wetland species as reflected in the fact that wetlands have declined by more than 50% in Europe and North America during the last century. Both current and historic land-use patterns are likely to be significant determinants of wetland species' distributions; however their relative importance is often unknown. We studied the importance of local (study pond) and landscape (current and 18th-century landscape) characteristics in explaining the occurrence and species richness of amphibians (Rana arvalis, Bufo bufo, and Triturus vulgaris) on the Swedish island of Gotland, where more than 40% of wetlands have been lost since the 18th century. Current local habitat characteristics were the strongest determinants of occurrence for all study species. Additionally, species occurrence was related to current and historic landscape characteristics, which generally explained equal amounts of the variation in species-occurrence data. The proportions of both current and historic arable land were negative determinants of amphibian occurrence and species richness, indicating that agricultural land use may have an overall negative impact on amphibians, and that amphibians may occur less frequently in areas with a long agricultural history. Likewise, historic forest area was positively related to B. bufo occurrence and species richness, whereas current forests had no significant effects, suggesting that there may be a lag in the response of amphibians to agriculture-mediated habitat loss. Our results suggest that historic land-use patterns may influence current amphibian populations and that inclusion of historic land-use information could be a valuable tool in future studies on amphibian-habitat relations.     

Effects of Agricultural Cultivation on Demographics of Southern High Plains Amphibians

Abstract:  Anthropogenic disturbance of landscapes surrounding wetlands is considered a factor in local and global amphibian declines. Few data exist on the effects of agricultural cultivation of wetland watersheds on amphibians, and results from previous studies are contradictory. Our objective was to test the effects of general anthropogenic land use (cultivation vs. grassland) on the demographics of seven species and three age classes of amphibians in the Southern High Plains of Texas. We partially enclosed 16 playa wetlands (4 per land use per year) with drift fences and pitfall traps and monitored relative daily abundance and diversity from 16 May to 17 October 1999 and 19 April to 18 August 2000. In general, relative abundance (i.e., average daily capture) of New Mexico and plains spadefoots (  Spea multiplicata, S. bombifrons ) was greater at cropland than grassland playas; the abundance of other species and diversity of the amphibian assemblage was not affected by land use. Also, abundance generally was greater in 1999 than 2000 for metamorph spadefoots and barred tiger salamanders ( Ambystoma tigrinum mavortium ). Differences in spadefoot abundance between land-use types may have been related to low species-specific vagility, resulting in increased nestedness within disturbed landscapes and reduced abundance of a potential keystone intraguild predator in cropland playas. The yearly difference in amphibian abundance was likely related to annual precipitation, which influenced wetland hydroperiod. Agricultural cultivation surrounding wetlands is associated with the increased abundance of some amphibian species, but other demographic and fitness parameters—such as temporal demographics, body size, and diet diversity—may be negatively affected.     

Demographic Consequences of Terrestrial Habitat Loss for Pool‐Breeding Amphibians: Predicting Extinction Risks Associated with Inadequate Size of Buffer Zones

Abstract: Much of the biodiversity associated with isolated wetlands requires aquatic and terrestrial habitat to maintain viable populations. Current federal wetland regulations in the United States do not protect isolated wetlands or extend protection to surrounding terrestrial habitat. Consequently, some land managers, city planners, and policy makers at the state and local levels are making an effort to protect these wetland and neighboring upland habitats. Balancing human land‐use and habitat conservation is challenging, and well‐informed land‐use policy is hindered by a lack of knowledge of the specific risks of varying amounts of habitat loss. Using projections of wood frog (Rana sylvatica) and spotted salamander (Ambystoma maculatum) populations, we related the amount of high‐quality terrestrial habitat surrounding isolated wetlands to the decline and risk of extinction of local amphibian populations. These simulations showed that current state‐level wetland regulations protecting 30 m or less of surrounding terrestrial habitat are inadequate to support viable populations of pool‐breeding amphibians. We also found that species with different life‐history strategies responded differently to the loss and degradation of terrestrial habitat. The wood frog, with a short life span and high fecundity, was most sensitive to habitat loss and isolation, whereas the longer‐lived spotted salamander with lower fecundity was most sensitive to habitat degradation that lowered adult survival rates. Our model results demonstrate that a high probability of local amphibian population persistence requires sufficient terrestrial habitat, the maintenance of habitat quality, and connectivity among local populations. Our results emphasize the essential role of adequate terrestrial habitat to the maintenance of wetland biodiversity and ecosystem function and offer a means of quantifying the risks associated with terrestrial habitat loss and degradation.     

The importance of defining focal assemblages when evaluating amphibian and reptile responses to land use

Habitat loss and degradation are primary threats to amphibians and reptiles, but the relative effects of common land uses on assemblages and the mechanisms that underlie faunal responses are poorly studied. We reviewed the effects of four prevalent types of habitat alteration (urbanization, agriculture, livestock grazing, and silviculture) on amphibian and reptile species richness and abundance by summarizing reported responses in the literature and by estimating effect sizes across studies for species richness in each land‐use type. We then used a multinomial model to classify species as natural habitat specialists, generalists, and disturbed habitat specialists and examined variation in effect sizes for each land‐use type according to habitat specialization categories. There were mixed conclusions from individual studies, some reporting negative, neutral, or positive effects of land use on species richness and total abundance. A large proportion of studies reported species‐specific effects of individual species abundance. However, in our analysis of effect sizes, we found a general trend of negative effects of land use on species richness. We also demonstrate that habitat associations of common species and species turnover can explain variation in the effect of land use on herpetofauna. Our review highlights the pervasive negative effects of common land uses on amphibians and reptiles, the importance of identifying groups vulnerable to land‐use change (e.g., forest‐associated species) in conservation studies, and the potential influence of disturbance‐associated species on whole assemblage analyses.     

Effects of river-flow regulation on anuran occupancy and abundance in riparian zones

The natural flow regimes of rivers worldwide have been heavily altered through anthropogenic activities, and dams in particular have a pervasive effect on riverine ecosystems. Flow-regulation effects of dams negatively affect species diversity and abundance of a variety of aquatic animals, including invertebrates and fishes. However, the effects on semiaquatic animals are relatively unknown. We conducted anuran calling surveys at 42 study locations along the Broad and Pacolet Rivers in South Carolina to address the potential effects of flow regulation by damming on anuran occupancy and abundance. We estimated occupancy and abundance with Program PRESENCE. Models incorporated distance upstream and downstream from the nearest dam as covariates and urbanization pressure as an alternative stressor. Distance from dam was associated with occupancy of 2 of the 9 anuran species in our analyses and with abundance of 6 species. In all cases, distance downstream from nearest dam was a better predictor of occupancy and abundance than distance upstream from nearest dam. For all but one species, distance downstream from nearest dam was positively correlated with both occupancy and abundance. Reduced occupancy and abundance of anurans likely resulted from downstream alterations in flow regime associated with damming, which can lead to reduced area of riparian wetlands that serve as anuran breeding habitat. Our results showed that damming has a strong negative effect on multiple anuran species across large spatial extents and suggest that flow regulation can affect semiaquatic animals occupying riparian zones.     

Testing wetland features to increase amphibian reproductive success and species richness for mitigation and restoration

Aquatic habitat features can directly influence the abundance, species richness, and quality of juvenile amphibians recruited into adult populations. We examined the influences of within-wetland slope, vegetation, and stocked mosquito fish (Gambusia affinis) on amphibian metamorph production and species richness during the first two years post-construction at 18 experimental wetlands in northeast Missouri (U.S.A.) grasslands. We used an information theoretic approach (AICc) to rank regression models representing total amphibian metamorph production, individual amphibian species metamorph production, and larval amphibian species richness. Total amphibian metamorph production was greatest at shallow-sloped, fish-free wetlands during the first year, but shallow-sloped wetlands with high vegetation cover were best the second year. Species richness was negatively associated with fish and positively associated with vegetation in both survey years. Leopard frog (Rana blairi/sphenocephala complex) metamorph quality, based on average metamorph size, was influenced by slope and the number of cohorts in the wetland. However, the tested variables had little influence on the size of American toads (Bufo americanus) or boreal chorus frogs (Pseudacris maculata). Our results indicate that wetlands designed to act as functional reproductive habitat for amphibians should incorporate shallows, high amounts of planted or naturally established vegetation cover, and should be fish-free.     

Effect of Human Disturbance on Bee Communities in a Forested Ecosystem

Abstract:  It is important for conservation biologists to understand how well species persist in human-dominated ecosystems because protected areas constitute a small fraction of the Earth's surface and because anthropogenic habitats may offer more opportunities for conservation than has been previously thought. We investigated how an important functional group, pollinators (bees; Hymenoptera: Apiformes), are affected by human land use at the landscape and local scales in southern New Jersey (U.S.A.). We established 40 sites that differed in surrounding landscape cover or local habitat type and collected 2551 bees of 130 species. The natural habitat in this ecosystem is a forested, ericaceous heath. Bee abundance and species richness within forest habitat decreased, not increased, with increasing forest cover in the surrounding landscape. Similarly, bee abundance was greater in agricultural fields and suburban and urban developments than in extensive forests, and the same trend was found for species richness. Particular species groups that might be expected to show greater sensitivity to habitat loss, such as floral specialists and bees of small or large body size, did not show strong positive associations with forest habitat. Nevertheless, 18 of the 130 bee species studied were positively associated with extensive forest. One of these species is a narrow endemic that was last seen in 1939. Our results suggest that at least in this system, moderate anthropogenic land use may be compatible with the conservation of many, but not all, bee species.     

Amphibian Breeding Distribution in an Urbanized Landscape

Abstract:  Amphibians commonly use wetlands for breeding habitat, and given the concern about their ongoing global declines, the effects of urbanization on the breeding distribution of amphibians need to be quantified. Thus, we conducted a survey of the larval amphibian community in central Pennsylvania (U.S.A.) wetlands along an urbanization gradient. Wetlands in urban areas had less surrounding forest and wetlands and greater road density than rural wetlands. Urbanization was also associated with increases in hydroperiod (i.e., wetland permanency) and the presence of fish predators. Moreover, urban wetlands had lower larval amphibian species richness than rural wetlands. This decrease in richness was attributable to a decrease in occurrence of wood frogs ( Rana sylvatica ) and ambystomatid salamanders ( Ambystoma maculatum and A. jeffersonianum ) in urban sites. Wood frogs and ambystomatid salamanders were positively associated with the amount of forest surrounding sites and negatively associated with hydroperiod. As a result, we hypothesize that these species are sensitive to the effects of urban development. The remaining species in this study appear fairly resilient to the effects of urbanization. These data demonstrate the importance of quantifying both local and landscape attributes when describing the factors that limit the breeding distribution of amphibians. We recommend that to preserve amphibian biodiversity in urbanized landscapes, it is best to focus on regional diversity, which protects a variety of sites that encompass various hydroperiods, have adequate buffer habitat, and are connected by dispersal routes.     

Effects of Forest Fragmentation and Habitat Degradation on West African Leaf-Litter Frogs

Abstract:  Habitat degradation alters the dynamics and composition of anuran assemblages in tropical forests. The effects of forest fragmentation on the composition of anuran assemblages are so far poorly known. We studied the joint influence of forest fragmentation and degradation on leaf-litter frogs. We specifically asked whether the processes structuring leaf-litter anuran assemblages in fragmented forests are the same as those in continuous forests. We analyzed anuran assemblages with respect to habitat characteristics, including fragmentation and degradation parameters. In comparison with continuous forests, species richness and diversity were lower and assemblage composition was altered in forest fragments. These changes seemed to be mainly caused by habitat degradation rather than forest fragmentation. Availability of aquatic sites for breeding, vegetation structure (including those variables indicating degradation), and leaf-litter cover had the most influence on the presence of single species. The comparatively small impact of fragmentation on anurans might be due to the location of the study area; it still possessed large tracts of continuous forest. These forest blocks may stabilize the regional rainforest climate and thus weaken the effects of fragmentation .     

Effects of Oil‐Palm Plantations on Diversity of Tropical Anurans

Agriculturally altered vegetation, especially oil‐palm plantations, is rapidly increasing in Southeast Asia. Low species diversity is associated with this commodity, but data on anuran diversity in oil‐palm plantations are lacking. We investigated how anuran biological diversity differs between forest and oil‐palm plantation, and whether observed differences in biological diversity of these areas is linked to specific environmental factors. We hypothesized that biological diversity is lower in plantations and that plantations support a larger proportion of disturbance‐tolerant species than forest. We compared species richness, abundance, and community composition between plantation and forest areas and between site types within plantation and forest (forest stream vs. plantation stream, forest riparian vs. plantation riparian, forest terrestrial vs. plantation terrestrial). Not all measures of biological diversity differed between oil‐palm plantations and secondary forest sites. Anuran community composition, however, differed greatly between forest and plantation, and communities of anurans in plantations contained species that prosper in disturbed areas. Although plantations supported large numbers of breeding anurans, we concluded the community consisted of common species that were of little conservation concern (commonly found species include Fejervarya limnocharis, Microhyla heymonsi, and Hylarana erythrea). We believe that with a number of management interventions, oil‐palm plantations can provide habitat for species that dwell in secondary forests. Efectos de las Plantaciones de Palma de Aceite sobre la Diversidad de Anuros Tropicales Faruk et al.     

Assessing Risks to Biodiversity from Future Landscape Change

We examined the impacts of possible future land development patterns on the biodiversity of a landscape. Our landscape data included a remote sensing derived map of the current habitat of the study area and six maps of future habitat distributions resulting from different land development scenarios. Our species data included lists of all bird, mammal, reptile, and amphibian species in the study area, their habitat associations, and area requirements for each. We estimated the area requirements using home ranges, sampled population densities, or genetic area requirements that incorporate dispersal distances. Our measures of biodiversity were species richness and habitat abundance. We calculated habitat abundance in two ways. First, we computed the total habitat area for each species in each landscape. Second, we calculated the number of habitat units for each species in each landscape by dividing the size of each habitat patch in the landscape by the area requirement and summing over all patches. Species richness was based on presence of habitat. Species became extinct in the landscape if they had no habitat area or no habitat units, respectively. We then computed ratios of habitat abundance in each future landscape to habitat abundance in the present for each species. We also computed the ratio of future to present species richness. We then calculated summary statistics across all species. Species richness changed little from present to future. There were distinctly greater risks to habitat abundance in landscapes that extrapolated from present trends or zoning patterns, however, as opposed to landscapes in which land development activities followed more constrained patterns. These results were stable when tested using Monte Carlo simulations and sensitivity tests on the area requirements. We conclude that this methodology can begin to discriminate the effects of potential changes in land development on vertebrate biodiversity.     

Functional Equivalency between Rice Fields and Seminatural Wetland Habitats

Abstract: Evaluating the potential for anthropogenic habitats to act as surrogates for the natural habitats they replace is a key issue in conservation biology. In California, flooded rice fields are used by numerous aquatic birds during winter. If this habitat functions similarly to more natural wetlands, increased flooding may help replace the extensive wetlands that occurred in the region prior to agricultural development. I tested whether food abundance, perceived predation threat, foraging performance, and the way in which birds allocate their time to different behaviors differed between flooded rice fields and seminatural wetlands for several species of aquatic bird. When appropriate, I also compared flooded and unflooded fields. Invertebrate densities did not differ among habitats. Seminatural wetlands had less rice grain but more seeds from other plants than the two rice habitats. The frequency with which predators passed over a feeding area was lower in flooded fields than in unflooded fields or seminatural wetlands. Most differences in feeding performance and time allocation among habitats were small and statistically insignificant. For some species, feeding efficiency was greater in seminatural wetlands than in flooded fields. Increasing attack rates and the amount of time spent feeding when in flooded fields, however, may allow birds to compensate for reduced efficiency. Multivariate analyses showed that group size, predation threat, time of day, date, and water depth often were associated with behaviors, but that these variables rarely accounted for habitat differences. Flooded fields apparently provide equivalent foraging habitat to seminatural wetlands and, because of reduced predation threat, may be a safer habitat for waterbirds. Thus, if managed appropriately, one of the world's dominant forms of agriculture can provide valuable waterbird habitat.     

Effects of Conversion of Dry Tropical Forest to Agricultural Mosaic on Herpetofaunal Assemblages

Abstract: We explored the impact of forest conversion to agricultural mosaic on anuran, lizard, snake, and turtle assemblages of Neotropical dry forests. Over 2 years, we sampled 6 small watersheds on the west coast of Mexico, 3 conserved and 3 disturbed. The disturbed watersheds were characterized by a mosaic of pastures and cultivated fields (corn, beans, squash) intermingled with patches of different successional stages of dry forest. In each watershed, we conducted 11 diurnal and nocturnal time‐constrained searches in 10 randomly established plots. We considered vulnerability traits of species in relation to habitat modification. Eighteen anuran, 18 lizard, 23 snake, and 3 turtle species were recorded. Thirty‐six species (58%) occurred in both forest conditions, and 14 (22%) and 12 species (19%) occurred only in the conserved and disturbed sites, respectively. Assemblages responded differently to disturbance. Species richness, diversity, and abundance of lizards were higher in disturbed forests. Anuran diversity and species richness were lower in disturbed forest but abundance was similar in both forest conditions. Diversity, richness, and abundance of turtles were lower in disturbed forest. The structure and composition of snake assemblages did not differ between forest conditions. We considered species disturbance sensitive if their abundance was significantly less in disturbed areas. Four anuran (22%), 2 lizard (11%), and 3 turtle (100%) species were sensitive to disturbance. No snake species was sensitive. The decline in abundance of disturbance‐sensitive species was associated with the reduction of forest canopy cover, woody stem cover, roots, and litter‐layer ground cover. Anuran species with small body size and direct embryonic development were especially sensitive to forest disturbance. An important goal for the conservation of herpetofauna should be the determination of species traits associated with extinction or persistence in agricultural mosaics.     

Spatial and Seasonal Patterns of Bird Communities in Italian Agroecosystems

Abstract:  Despite agricultural landscapes covering almost 60% of the total land area of Italy, knowledge of the effects of agriculture and its intensification on bird communities is still scarce. I analyzed the effects of land uses on bird diversity and community structure in different farmland habitats of lowland northwestern Italy. I surveyed breeding and overwintering birds with a hierarchically nested sampling design and used generalized linear and mixed models to investigate the relationships between the diversity and abundance of birds and habitat or landscapes attributes. The effects of agriculture on α avian diversity varied with season and spatial scale, whereas nonagricultural habitats (long-term fallows or woodlands) had a generally positive effect that was constant throughout time and space. As the amount of woodland habitat increased, spatial turnover (β diversity) of breeding birds decreased. Arable landscapes supported low levels of avian diversity throughout the year but were favored by emblematic farmland birds that have declined severely in Europe and in the study area. Farmland birds (40% of which are experiencing population declines) were more abundant or foraged more frequently in the less-disturbed habitat types such as fallows, grasslands, and winter stubbles and tended to avoid the prevailing cultivations (maize, vineyard, and wheat). Landscape simplification, the expansion of maize cultivation, winter plowing practices, and the conversion of highly diverse grasslands to tilled lands are likely to be responsible for the local decline of most farmland species (Skylark [Alauda arvensis ], Starling [Sturnus vulgaris ], buntings [Emberiza spp.], and wagtails [Motacilla spp.]) and for the increase of birds that are turning into agricultural pests (Hooded Crow [Corvus corone cornix ]).     

Countryside Biogeography of Moths in a Fragmented Landscape: Biodiversity in Native and Agricultural Habitats

Abstract: Studies of fragmented landscapes, especially in the tropics, have traditionally focused on the native fragments themselves, ignoring species distributions in surrounding agricultural or other human-dominated areas. We sampled moth species richness within a 227-ha forest fragment and in four surrounding agricultural habitats (coffee, shade coffee, pasture, and mixed farms) in southern Costa Rica. We found no significant difference in moth species richness or abundance among agricultural habitats, but agricultural sites within 1 km of the forest fragment had significantly higher richness and abundance than sites farther than 3.5 km from the fragment. In addition, species composition differed significantly between distance classes ( but not among agricultural habitats), with near sites more similar to forest than far sites. These results suggest that (1) different agricultural production regimes in this region may offer similar habitat elements and thus may not differ substantially in their capacities to support native moth populations and (2) that the majority of moths may utilize both native and agricultural habitats and move frequently between them, forming "halos" of relatively high species richness and abundance around forest fragments. Correlations between species richness and the amount of nearby forest cover, measured over circles of various radii around the sites, suggest that halos extend approximately 1.0–1.4 km from the forest edge. The extent of these halos likely differs among taxa and may influence their ability to survive in fragmented landscapes.     

Relationships among Isolated Wetland Size, Hydroperiod, and Amphibian Species Richness: Implications for Wetland Regulations

Abstract: Wetland development within the United States is regulated primarily by size. Decisions concerning wetland destruction or conservation are therefore based in part on three inherent assumptions: (1) small wetlands contain water for short portions of the year; (2) small wetlands support few species; and (3) species found in small wetlands are also found in larger wetlands. We tested these assumptions using data on wetland size, relative hydroperiod (drying scores), and relative species richness of amphibians in depression wetlands of the southeastern United States. We found a significant (p = 0.03) but weak (r² = 0.05) relationship between hydroperiod and wetland size and no relationship (p = 0.48) between amphibian species richness and wetland size. Furthermore, synthetic models of lentic communities predict that short-hydroperiod wetlands support a unique group of species. Empirical investigations support this prediction. Our results indicate that hydroperiod length should be included as a primary criterion in wetland regulations. We advocate a landscape approach to wetlands regulation, focused in part on conserving a diversity of wetlands that represent the entire hydroperiod gradient.     

Multiple stressors in amphibian communities: effects of chemical contamination, bullfrogs, and fish.

A leading hypothesis of amphibian population declines is that combinations of multiple stressors contribute to declines. We examined the role that chemical contamination, competition, and predation play singly and in combination in aquatic amphibian communities. We exposed larvae of American toads (Bufo americanus), southern leopard frogs (Rana sphenocephala), and spotted salamanders (Ambystoma maculatum) to overwintered bullfrog tadpoles (R. catesbeiana), bluegill sunfish (Lepomis macrochirus), the insecticide carbaryl, and ammonium nitrate fertilizer in 1000-L mesocosms. Most significantly, our study demonstrated that the presence of multiple factors reduced survival of B. americanus and A. maculatum and lengthened larval periods of R. sphenocephala. The presence of bluegill had the largest impact on the community; it eliminated B. americanus and A. maculatum and reduced the abundance of R. sphenocephala. Chemical contaminants had the second strongest effect on the community with the insecticide, reducing A. maculatum abundance by 50% and increasing the mass of anurans (frogs and toads) at metamorphosis; the fertilizer positively influenced time and mass at metamorphosis for both anurans and A. maculatum. Presence of overwintered bullfrogs reduced mass and increased time to metamorphosis of anurans. While both bluegill and overwintered bullfrog tadpoles had negative effects on the amphibian community, they performed better in the presence of one another and in contaminated habitats. Our results indicate that predicting deleterious combinations from single-factor effects may not be straightforward. Our research supports the hypothesis that combinations of factors can negatively impact some amphibian species and could contribute to population declines.     

Anthropogenic Correlates of Species Richness in Southeastern Ontario Wetlands

We examined the relationship between the richness of four different wetland taxa (birds, mammals, herptiles, and plants) in 30 southeastern Ontario, Canada wetlands and two anthropogenic factors: road construction and forest removal/conversion on adjacent lands. Data were obtained from two sources: road densities and forest cover from 1:50,000 Government of Canada topographic maps and species lists and wetland areas from Ontario Ministry of Natural Resources wetland evaluation reports. Multiple regression analysis was used to model the relationships between species richness and wetland area, road density, and forest cover. Our results show a strong positive relationship between wetland area and species richness for all taxa. The species richness of all taxa except mammals was negatively correlated with the density of paved roads on lands up to 2 km from the wetland. Furthermore, both herptile and mammal species richness showed a strong positive correlation with the proportion of forest cover on lands within 2 km. These results provide evidence that at the landscape level, road construction and forest removal on adjacent lands pose significant risks to wetland biodiversity. Furthermore, they suggest that most existing wetland policies, which focus almost exclusively on activities within the wetland itself and/or a narrow buffer zone around the wetland perimeter, are unlikely to provide adequate protection for wetland biodiversity.