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.     

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.     

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.     

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.     

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.     

Effects of Landscape Composition and Wetland Fragmentation on Frog and Toad Abundance and Species Richness in Iowa and Wisconsin, U.S.A.

Abstract: Management of amphibian populations to reverse recent declines will require defining high-quality habitat for individual species or groups of species, followed by efforts to retain or restore these habitats on the landscape. We examined landscape-level habitat relationships for frogs and toads by measuring associations between relative abundance and species richness based on survey data derived from anuran calls and features of land-cover maps for Iowa and Wisconsin. The most consistent result across all anuran guilds was a negative association with the presence of urban land. Upland and wetland forests and emergent wetlands tended to be positively associated with anurans. Landscape metrics that represent edges and patch diversity also had generally positive associations, indicating that anurans benefit from a complex of habitats that include wetlands. In Iowa the most significant associations with relative abundance were the length of the edge between wetland and forest ( positive) and the presence of urban land (negative). In Wisconsin the two most significant associations with relative abundance were forest area and agricultural area ( both positive). Anurans had positive associations with agriculture in Wisconsin but not in Iowa. Remnant forest patches in agricultural landscapes may be providing refuges for some anuran species. Differences in anuran associations with deep water and permanent wetlands between the two states suggest opportunities for management action. Large-scale maps can contribute to predictive models of amphibian habitat use, but water quality and vegetation information collected from individual wetlands will likely be needed to strengthen those predictions. Landscape habitat analyses provide a framework for future experimental and intensive research on specific factors affecting the health of anurans.     

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.     

Relative influence of environmental factors and fishing on coral reef fish assemblages

Understanding whether assemblages of species respond more strongly to bottom-up (availability of trophic resources or habitats) or top-down (predation pressure) processes is important for effective management of resources and ecosystems. We determined the relative influence of environmental factors and predation by humans in shaping the density, biomass, and species richness of 4 medium-bodied (10–40 cm total length [TL]) coral reef fish groups targeted by fishers (mesopredators, planktivores, grazer and detritivores, and scrapers) and the density of 2 groups not targeted by fishers (invertivores, small fish ≤10 cm TL) in the central Philippines. Boosted regression trees were used to model the response of each fish group to 21 predictor variables: 13 habitat variables, 5 island variables, and 3 fishing variables (no-take marine reserve [NTMR] presence or absence, NTMR size, and NTMR age). Targeted and nontargeted fish groups responded most strongly to habitat variables, then island variables. Fishing (NTMR) variables generally had less influence on fish groups. Of the habitat variables, live hard coral cover, structural complexity or habitat complexity index, and depth had the greatest effects on density, biomass, and species richness of targeted fish groups and on the density of nontargeted fishes. Of the island variables, proximity to the nearest river and island elevation had the most influence on fish groups. The NTMRs affected only fishes targeted by fishers; NTMR size positively correlated with density, biomass, and species richness of targeted fishes, particularly mesopredatory, and grazing and detritivorous fishes. Importantly, NTMRs as small as 15 ha positively affected medium-bodied fishes. This finding provides reassurance for regions that have invested in small-scale community-managed NTMRs. However, management strategies that integrate sound coastal land-use practices to conserve adjacent reef fish habitat, strategic NTMR placement, and establishment of larger NTMRs will be crucial for maintaining biodiversity and fisheries.     

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.     

Effects of Land‐Use Change on Community Composition of Tropical Amphibians and Reptiles in Sulawesi,Indonesia

Abstract: Little is known about the effects of anthropogenic land‐use change on the amphibians and reptiles of the biodiverse tropical forests of Southeast Asia. We studied a land‐use modification gradient stretching from primary forest, secondary forest, natural‐shade cacao agroforest, planted‐shade cacao agroforest to open areas in central Sulawesi, Indonesia. We determined species richness, abundance, turnover, and community composition in all habitat types and related these to environmental correlates, such as canopy heterogeneity and thickness of leaf litter. Amphibian species richness decreased systematically along the land‐use modification gradient, but reptile richness and abundance peaked in natural‐shade cacao agroforests. Species richness and abundance patterns across the disturbance gradient were best explained by canopy cover and leaf‐litter thickness in amphibians and by canopy heterogeneity and cover in reptiles. Amphibians were more severely affected by forest disturbance in Sulawesi than reptiles. Heterogeneous canopy cover and thick leaf litter should be maintained in cacao plantations to facilitate the conservation value for both groups. For long‐term and sustainable use of plantations, pruned shade trees should be permanently kept to allow rejuvenation of cacao and, thus, to prevent repeated forest encroachment.     

A test of the substitution–habitat hypothesis in amphibians

Most examples that support the substitution‐habitat hypothesis (human‐made habitats act as substitutes of original habitat) deal with birds and mammals. We tested this hypothesis in 14 amphibians by using percentage occupancy as a proxy of habitat quality (i.e., higher occupancy percentages indicate higher quality). We classified water body types as original habitat (no or little human influence) depending on anatomical, behavioral, or physiological adaptations of each amphibian species. Ten species had relatively high probabilities (0.16–0.28) of occurrence in original habitat, moderate probability of occurrence in substitution habitats (0.11–0.14), and low probability of occurrence in refuge habitats (0.05–0.08). Thus, the substitution–habitat hypothesis only partially applies to amphibians because the low occupancy of refuges could be due to the negligible human persecution of this group (indicating good conservation status). However, low occupancy of refuges could also be due to low tolerance of refuge conditions, which could have led to selective extinction or colonization problems due to poor dispersal capabilities. That original habitats had the highest probabilities of occupancy suggests amphibians have a good conservation status in the region. They also appeared highly adaptable to anthropogenic substitution habitats.     

Value of Artificial Habitats for Amphibian Reproduction in Altered Landscapes

Abstract:  Installation and maintenance of stormwater ponds to detain and treat runoff from impervious surfaces is a common method of stormwater control in developed areas. That these ponds capture pollutants, however, is of concern for wildlife species that use the ponds, particularly pond-breeding amphibians. To assess the relative contribution of stormwater ponds to the persistence of amphibian populations in suburban landscapes, we compared amphibian use of stormwater ponds and other available wetlands in suburban and forested watersheds. We surveyed three suburban and three primarily forested first-order watersheds to identify all potential wetlands that might serve as breeding sites for pond-breeding amphibians. We performed call, egg-mass, and larval surveys to measure breeding effort at each wetland in spring and summer 2007 and 2008. In suburban watersheds most (89%) of the wetlands that had breeding activity were either stormwater ponds or otherwise artificial. This pattern was also evident in the forested watersheds, where amphibians were primarily found breeding in wetlands created by past human activity. Late-stage larvae were found only in anthropogenic wetlands in all study areas because the remaining natural wetlands did not hold water long enough for larvae to complete development. Our results suggest that in urban and suburban landscapes with naturally low densities of wetlands, wetlands created by current or historic land uses may be as important to amphibian conservation as natural wetlands or pools and that management strategies directed at urban and suburban landscapes should recognize and incorporate human-created wetlands.     

Effects of Urbanization on the Distribution and Abundance of Amphibians and Invasive Species in Southern California Streams

Abstract:  Urbanization negatively affects natural ecosystems in many ways, and aquatic systems in particular. Urbanization is also cited as one of the potential contributors to recent dramatic declines in amphibian populations. From 2000 to 2002 we determined the distribution and abundance of native amphibians and exotic predators and characterized stream habitat and invertebrate communities in 35 streams in an urbanized landscape north of Los Angeles (U.S.A.). We measured watershed development as the percentage of area within each watershed occupied by urban land uses. Streams in more developed watersheds often had exotic crayfish ( Procambarus clarkii ) and fish, and had fewer native species such as California newts ( Taricha torosa ) and California treefrogs ( Hyla cadaverina ). These effects seemed particularly evident above 8% development, a result coincident with other urban stream studies that show negative impacts beginning at 10–15% urbanization. For Pacific treefrogs ( H. regilla ), the most widespread native amphibian, abundance was lower in the presence of exotic crayfish, although direct urbanization effects were not found. Benthic macroinvertebrate communities were also less diverse in urban streams, especially for sensitive species. Faunal community changes in urban streams may be related to changes in physical stream habitat, such as fewer pool and more run habitats and increased water depth and flow, leading to more permanent streams. Variation in stream permanence was particularly evident in 2002, a dry year when many natural streams were dry but urban streams were relatively unchanged. Urbanization has significantly altered stream habitat in this region and may enhance invasion by exotic species and negatively affect diversity and abundance of native amphibians.     

Traits, not origin, explain impacts of plants on larval amphibians

Managing habitats for the benefit of native fauna is a priority for many government and private agencies. Often, these agencies view nonnative plants as a threat to wildlife habitat, and they seek to control or eradicate nonnative plant populations. However, little is known about how nonnative plant invasions impact native fauna, and it is unclear whether managing these plants actually improves habitat quality for resident animals. Here, we compared the impacts of native and nonnative wetland plants on three species of native larval amphibians; we also examined whether plant traits explain the observed impacts. Specifically, we measured plant litter quality (carbon : nitrogen : phosphorus ratios, and percentages of lignin and soluble phenolics) and biomass, along with a suite of environmental conditions known to affect larval amphibians (hydroperiod, temperature, dissolved oxygen, and pH). Hydroperiod and plant traits, notably soluble phenolics, litter C:N ratio, and litter N:P ratio, impacted the likelihood that animals metamorphosed, the number of animals that metamorphosed, and the length of larval period. As hydroperiod decreased, the likelihood that amphibians achieved metamorphosis and the percentage of tadpoles that successfully metamorphosed also decreased. Increases in soluble phenolics, litter N:P ratio, and litter C:N ratio decreased the likelihood that tadpoles achieved metamorphosis, decreased the percentage of tadpoles metamorphosing, decreased metamorph production (total metamorph biomass), and increased the length of larval period. Interestingly, we found no difference in metamorphosis rates and length of larval period between habitats dominated by native and nonnative plants. Our findings have important implications for habitat management. We suggest that to improve habitats for native fauna, managers should focus on assembling a plant community with desirable traits rather than focusing only on plant origin.     

Frog occupancy of polluted wetlands in urban landscapes

Urban sprawl and the rising popularity of water-sensitive urban design of urban landscapes has led to a global surge in the number of wetlands constructed to collect and treat stormwater runoff in cities. However, contaminants, such as heavy metals and pesticides, in stormwater adversely affect the survival, growth, and reproduction of animals inhabiting these wetlands. A key question is whether wildlife can identify and avoid highly polluted wetlands. We investigated whether pond-breeding frogs are attempting to breed in wetlands that affect the fitness of their offspring across 67 urban wetlands in Melbourne, Australia. Frog species richness and the concentration of contaminants (heavy metals and pesticides) were not significantly related, even in the most polluted wetlands. The proportion of fringing vegetation at a wetland had the greatest positive influence on the number of frog species present and the probability of occurrence of individual species, indicating that frogs inhabited wetlands with abundant vegetation, regardless of their pollution status. These wetlands contained contaminant levels similar to urban wetlands around the world at levels that reduce larval amphibian survival. These results are, thus, likely generalizable to other areas, suggesting that urban managers could inadvertently be creating ecological traps in countless cities. Wetlands are important tools for the management of urban stormwater runoff, but their construction should not facilitate declines in wetland-dependent urban wildlife.     

Birds in agricultural mosaics: the influence of landscape pattern and countryside heterogeneity.

Agricultural environments are critical to the conservation of biota throughout the world. Efforts to identify key influences on the conservation status of fauna in such environments have taken complementary approaches. Many studies have focused on the role of remnant or seminatural vegetation and emphasized the influence on biota of spatial patterns in the landscape. Others have recognized that many species use diverse "countryside" elements within farmland, and emphasize the benefits of landscape heterogeneity for conservation. Here, we investigated the effect of independent measures of both the spatial pattern (extent and configuration) and heterogeneity of elements (i.e., land uses/vegetation types) on bird occurrence in farm-scale agricultural mosaics in southeastern Australia. Birds were sampled in all types of elements in 27 mosaics (each 1 x 1 km) selected to incorporate variation in cover of native vegetation and the number of different element types in the mosaic. We used an information-theoretic approach to identify the mosaic properties that most strongly influenced bird species richness. Subgroups of birds based on habitat requirements responded most strongly to the extent of preferred elements in mosaics. Woodland birds were richer in mosaics with higher cover of native vegetation while open-tolerant species responded to the extent of scattered trees. In contrast, for total species richness, mosaic heterogeneity (richness of element types) and landscape context (cover of native vegetation in surrounding area) had the greatest influence. These results showed that up to 76% of landscape-level variation in richness of bird groups is attributable to mosaic properties directly amenable to management by landowners. Key implications include (1) conservation goals for farm landscapes must be carefully defined because the richness of different faunal components is influenced by different mosaic properties; (2) the extent of native vegetation is a critical influence in agricultural environments because it drives the farm-scale richness of woodland birds and has a broader context effect on total bird richness in mosaics; (3) land-use practices that enhance the heterogeneity of farmland mosaics are beneficial for native birds; and (4) the cumulative effect of even small elements in farm mosaics contribute to the structural properties of entire landscapes.     

Habitat Split as a Cause of Local Population Declines of Amphibians with Aquatic Larvae

Abstract:  Most amphibian species have biphasic life histories and undergo an ontogenetic shift from aquatic to terrestrial habitats. In deforested landscapes, streams and forest fragments are frequently disjunct, jeopardizing the life cycle of forest-associated amphibians with aquatic larvae. We tested the impact of habitat split—defined as human-induced disconnection between habitats used by different life-history stages of a species—on four forest-associated amphibian species in a severely fragmented landscape of the Brazilian Atlantic Forest. We surveyed amphibians in forest fragments with and without streams (referred to as wet and dry fragments, respectively), including the adjacent grass-field matrix. Our comparison of capture rates in dry fragments and nearby streams in the matrix allowed us to evaluate the number of individuals that engaged in high-risk migrations through nonforested habitats. Adult amphibians moved from dry fragments to matrix streams at the beginning of the rainy season, reproduced, and returned at the end of the breeding period. Juveniles of the year moved to dry fragments along with adults. These risky reproductive migrations through nonforested habitats that expose individuals to dehydration, predation, and other hazards may cause population declines in dry fragments. Indeed, capture rates were significantly lower in dry fragments compared with wet fragments. Declining amphibians would strongly benefit from investments in the conservation and restoration of riparian vegetation and corridors linking breeding and nonbreeding areas.     

Coupling ecology and GIS to evaluate efficacy of marine protected areas in Hawaii.

In order to properly determine the efficacy of marine protected areas (MPAs), a seascape perspective that integrates ecosystem elements at the appropriate ecological scale is necessary. Over the past four decades, Hawaii has developed a system of 11 Marine Life Conservation Districts (MLCDs) to conserve and replenish marine resources around the state. Initially established to provide opportunities for public interaction with the marine environment, these MLCDs vary in size, habitat quality, and management regimes, providing an excellent opportunity to test hypotheses concerning MPA design and function using multiple discrete sampling units. Digital benthic habitat maps for all MLCDs and adjacent habitats were used to evaluate the efficacy of existing MLCDs using a spatially explicit stratified random sampling design. Analysis of benthic cover validated the a priori classification of habitat types and provided justification for using these habitat strata to conduct stratified random sampling and analyses of fish habitat utilization patterns. Results showed that a number of fish assemblage characteristics (e.g., species richness, biomass, diversity) vary among habitat types, but were significantly higher in MLCDs compared with adjacent fished areas across all habitat types. Overall fish biomass was 2.6 times greater in the MLCDs compared to open areas. In addition, apex predators and other species were more abundant and larger in the MLCDs, illustrating the effectiveness of these closures in conserving fish populations within their boundaries. Habitat type, protected area size, and level of protection from fishing were all important determinates of MLCD effectiveness with respect to their associated fish assemblages. Although size of these protected areas was positively correlated with a number of fish assemblage characteristics, all appear too small to have any measurable influence on the adjacent fished areas. These protected areas were not designed for biodiversity conservation or fisheries enhancement yet still provide varying degrees of protection for fish populations within their boundaries. Implementing this type of biogeographic process, using remote sensing technology and sampling across the range of habitats present within the seascape, provides a robust evaluation of existing MPAs and can help to define ecologically relevant boundaries for future MPA design in a range of locations.     

Deviations from predictions of the metabolic theory of ecology can be explained by violations of assumptions

The metabolic theory of ecology (MTE) is based on models derived from the first principles of thermodynamics and biochemical kinetics. The MTE predicts that the relationship between temperature and species richness of ectotherms should show a specific slope. Testing the validity of this model, however, depends on whether empirical data do not violate assumptions and are obtained within contour conditions. When dealing with richness gradients, the MTE must be empirically tested only for ectothermic organisms at high organization levels and when their body size as well as abundance does not vary with temperature gradients. Here we evaluate whether the magnitude of the deviations in slope expected from the MTE to empirical data for New World amphibians is due to the violations of model assumptions and to lack of generality due to restricting contour conditions. We found that the MTE correctly predicted biodiversity patterns only at higher levels of organization and when assumptions of the basic model were not violated. Approximately 60% of the deviations from the MTE-predicted slope across amphibian families were due to violations of the model assumptions. The hypothesis that richness patterns are a function of environmental temperature is too restrictive and does not take complex environmental and ecological processes into account. However, our results suggest that it may be possible to obtain multiple derivations of the MTE equation if idiosyncrasies in spatial and biological/ecological issues that are essential to understanding biodiversity patterns are considered.     

Declines in Woodland Salamander Abundance Associated with Non-Native Earthworm and Plant Invasions

Abstract:  Factors that negatively affect the quality of wildlife habitat are a major concern for conservation. Non-native species invasions, in particular, are perceived as a global threat to the quality of wildlife habitat. Recent evidence indicates that some changes to understory plant communities in northern temperate forests of North America, including invasions by 3 non-native plant species, are facilitated by non-native earthworm invasion. Furthermore, non-native earthworm invasions cause a reduction in leaf litter on the forest floor, and the loss of forest leaf litter is commonly associated with declines in forest fauna, including amphibians. We conducted a mark-recapture study of woodland salamander abundance across plant invasion fronts at 10 sites to determine whether earthworm or plant invasions were associated with reduced salamander abundance. Salamander abundance declined exponentially with decreasing leaf litter volume. There was no significant relationship between invasive plant cover and salamander abundance, independent of the effects of leaf litter loss due to earthworm invasion. An analysis of selected salamander prey abundance (excluding earthworms) at 4 sites showed that prey abundance declined with declining leaf litter. The loss of leaf litter layers due to non-native earthworm invasions appears to be negatively affecting woodland salamander abundance, in part, because of declines in the abundance of small arthropods that are a stable resource for salamanders. Our results demonstrate that earthworm invasions pose a significant threat to woodland amphibian fauna in the northeastern United States, and that plant invasions are symptomatic of degraded amphibian habitat but are not necessarily drivers of habitat degradation.