Joint effects of resources and amphibians on pond ecosystems.

Primary production can be controlled through bottom-up (e.g., resources) or top-down (e.g., predators) constraints. Two key bottom-up resources in small aquatic systems are light and nutrients, and forest canopy cover heavily influences these factors, whereas amphibian and invertebrate colonizers exert top-down pressure as grazers and predators. We designed our experiment to specifically manipulate two different top-down and bottom-up factors. We manipulated resources by altering light (low/high) and nutrient (low/high) availability; omnivores with the presence/absence of southern leopard frog tadpoles (Lithobates sphenocephalus); and predators with the presence/absence of spotted salamander larvae (Ambystoma maculatum) in a full-factorial experiment conducted over 14 weeks. We observed that both bottom-up and top-down effects were important in predicting lower trophic level biomass. We found a significant top-down effect of salamanders on Daphnia, but tadpoles had the strongest overall effect on the food web, influencing phytoplankton (+), periphyton (-), and chironomids (-). None of our models were good predictors of phytoplankton biomass, but both shading and nutrient availability relatively equally boosted periphyton biomass. We also found large temporal differences in food-web dynamics. Our results underscore the need for more information into how ecosystem functioning could be altered by land use, amphibian extirpation, and climate change.

Variation in phenology and density differentially affects predator-prey interactions between salamanders.

Variation in the timing of breeding (i.e., phenological variation) can affect species interactions and community structure, in part by shifting body size differences between species. Body size differences can be further altered by density-dependent competition, though synergistic effects of density and phenology on species interactions are rarely evaluated. We tested how field-realistic variation in phenology and density affected ringed salamander (Ambystoma annulatum) predation on spotted salamanders (Ambystoma maculatum), and whether these altered salamander dynamics resulted in trophic cascades. In outdoor mesocosms, we experimentally manipulated ringed salamander density (low/high) and breeding phenology (early/late) of both species. Ringed salamander body size at metamorphosis, development, and growth were reduced at higher densities, while delayed phenology increased hatchling size and larval development, but reduced relative growth rates. Survival of ringed salamanders was affected by the interactive effects of phenology and density. In contrast, spotted salamander growth, size at metamorphosis, and survival, as well as the biomass of lower trophic levels, were negatively affected primarily by ringed salamander density. In an additional mesocosm experiment, we isolated whether ringed salamanders could deplete shared resources prior to their interactions with spotted salamanders, but instead found direct interactions (e.g., predation) were the more likely mechanism by which ringed salamanders limited spotted salamanders. Overall, our results indicate the effects of phenological variability on fitness-related traits can be modified or superseded by differences in density dependence. Identifying such context dependencies will lead to greater insight into when phenological variation will likely alter species interactions.

Top predators and habitat complexity alter an intraguild predation module in pond communities.

Predator diversity and habitat complexity frequently influence species interactions at lower trophic levels, yet their joint investigation has been performed infrequently despite the demonstrated importance of each individual factor. We investigated how different top predators and varying habitat complexity influence the function of an intraguild predation module consisting of two larval salamanders, intraguild predator Ambystoma annulatum and intraguild prey A. maculatum. We manipulated predator food webs and habitat complexity in outdoor mesocosms. Top predators significantly influenced body condition and survival of A. annulatum, but habitat complexity had minimal effects on either response. A three-way interaction among the covariates top predator identity, habitat complexity and A. annulatum survival influenced body condition and survival of A. maculatum via a density-mediated indirect effect. Different top predator combinations had variable effects in different habitat complexity treatments on intraguild predator (A. annulatum) survival that subsequently influenced intraguild prey (A. maculatum) body condition and survival. Future work should consider how different top predators influence other food web components, which should vary due to predator attributes and the physical environments in which they co-occur.

Non-additive response of larval ringed salamanders to intraspecific density.

Conditions experienced in early developmental stages can have long-term consequences for individual fitness. High intraspecific density during the natal period can affect juvenile and eventually adult growth rates, metabolism, immune function, survival, and fecundity. Despite the important ecological and evolutionary effects of early developmental density, the form of the relationship between natal density and resulting juvenile phenotype is poorly understood. To test competing hypotheses explaining responses to intraspecific density, we experimentally manipulated the initial larval density of ringed salamanders (Ambystoma annulatum), a pond-breeding amphibian, over 11 densities. We modeled the functional form of the relationship between natal density and juvenile traits, and compared the relative support for the various hypotheses based on their goodness of fit. These functional form models were then used to parameterize a simple simulation model of population growth. Our data support non-additive density dependence and presents an alternate hypothesis to additive density dependence, self-thinning and Allee effects in larval amphibians. We posit that ringed salamander larvae may be under selective pressure for tolerance to high density and increased efficiency in resource utilization. Additionally, we demonstrate that models of population dynamics are sensitive to assumptions of the functional form of density dependence.

Life history differences influence the impacts of drought on two pond-breeding salamanders.

Drought is a strong density-independent environmental filter that contributes to population regulation and other ecological processes. Not all species respond similarly to drought, and the overall impacts can vary depending on life histories. Such differences can necessitate management strategies that incorporate information on individual species to maximize conservation success. We report the effects of a short-term drought on occupancy and reproductive success of two pond-breeding salamanders that differ in breeding phenology (fall vs. spring breeder) across an active military base landscape in Missouri, USA: We surveyed ~200 ponds for the presence of eggs, larvae, and metamorphs from 2011 to 2013. This period coincided with before, during, and after a severe drought that occurred in 2012. The two species showed contrasting responses to drought, where high reproductive failure (34% of ponds) was observed for the spring breeder during a single drought year. Alternatively, the fall breeder only showed a cumulative 8% failure over two years. The number of breeding ponds available for use in the fall decreased during the drought due to pond drying and/or a lack of re-filling. Estimates of occupancy probability declined for the fall-breeding salamander between 2012 and 2013, whereas occupancy probability estimates of the spring breeder increased post-drought. The presence of fish, hydroperiod, the amount of forest cover surrounding ponds, and canopy cover were all found to affect estimates of occupancy probabilities of each species. Pond clustering (distance to nearest pond and the number of ponds within close proximity), hydroperiod, forest cover, and canopy cover influenced both estimates of colonization and extinction probabilities. Our results show life history variation can be important in determining the relative susceptibility of a species to drought conditions, and that sympatric species experiencing the same environmental conditions can respond differently. Consideration of the spatial network and configuration of habitat patches that act as refuges under extreme environmental conditions will improve conservation efforts, such as the placement of permanent ponds for aquatic organisms. A better awareness of species-specific tolerances to environmental filters such as drought can lead to improved management recommendations to conserve and promote habitat for a greater diversity of species across landscapes of spatially connected populations.

Habitat traits and species interactions differentially affect abundance and body size in pond-breeding amphibians.

In recent studies, habitat traits have emerged as stronger predictors of species occupancy, abundance, richness and diversity than competition. However, in many cases, it remains unclear whether habitat also mediates processes more subtle than competitive exclusion, such as growth, or whether intra- and interspecific interactions among individuals of different species may be better predictors of size. To test whether habitat traits are a stronger predictor of abundance and body size than intra- and interspecific interactions, we measured the density and body size of three species of larval salamanders in 192 ponds across a landscape. We found that the density of larvae was best predicted by models that included habitat features, while models incorporating interactions among individuals of different species best explained the body size of larvae. Additionally, we found a positive relationship between focal species density and congener density, while focal species body size was negatively related to congener density. We posit that salamander larvae may not experience competitive exclusion and thus reduced densities, but instead compensate for increased competition behaviourally (e.g. reduced foraging), resulting in decreased growth. The discrepancy between larval density and body size, a strong predictor of fitness in this system, also highlights a potential shortcoming in using density or abundance as a metric of habitat quality or population health.

Abundance and phenology patterns of two pond-breeding salamanders determine species interactions in natural populations.

Phenology often determines the outcome of interspecific interactions, where early-arriving species often dominate interactions over those arriving later. The effects of phenology on species interactions are especially pronounced in aquatic systems, but the evidence is largely derived from experimental studies. We examined whether differences in breeding phenology between two pond-breeding salamanders (Ambystoma annulatum and A. maculatum) affected metamorph recruitment and demographic traits within natural populations, with the expectation that the fall-breeding A. annulatum would negatively affect the spring-breeding A. maculatum. We monitored populations of each species at five ponds over 4 years using drift fences. Metamorph abundance and survival of A. annulatum were affected by intra- and interspecific processes, whereas metamorph size and date of emigration were primarily influenced by intraspecific effects. Metamorph abundance, snout-vent length, date of emigration and survival for A. maculatum were all predicted by combinations of intra- and interspecific effects, but often showed negative relationships with A. annulatum metamorph traits and abundance. Size and date of metamorphosis were strongly correlated within each species, but in opposite patterns (negative for A. annulatum and positive for A. maculatum), suggesting that the two species use alternative strategies to enhance terrestrial survival and that these factors may influence their interactions. Our results match predictions from experimental studies that suggest recruitment is influenced by intra- and interspecific processes which are determined by phenological differences between species. Incorporating spatiotemporal variability when modeling population dynamics is necessary to understand the importance of phenology in species interactions, especially as shifts in phenology occur under climate change.

Ecological resistance surfaces predict fine-scale genetic differentiation in a terrestrial woodland salamander.

Landscape genetics has seen tremendous advances since its introduction, but parameterization and optimization of resistance surfaces still poses significant challenges. Despite increased availability and resolution of spatial data, few studies have integrated empirical data to directly represent ecological processes as genetic resistance surfaces. In our study, we determine the landscape and ecological factors affecting gene flow in the western slimy salamander (Plethodon albagula). We used field data to derive resistance surfaces representing salamander abundance and rate of water loss through combinations of canopy cover, topographic wetness, topographic position, solar exposure and distance from ravine. These ecologically explicit composite surfaces directly represent an ecological process or physiological limitation of our organism. Using generalized linear mixed-effects models, we optimized resistance surfaces using a nonlinear optimization algorithm to minimize model AIC. We found clear support for the resistance surface representing the rate of water loss experienced by adult salamanders in the summer. Resistance was lowest at intermediate levels of water loss and higher when the rate of water loss was predicted to be low or high. This pattern may arise from the compensatory movement behaviour of salamanders through suboptimal habitat, but also reflects the physiological limitations of salamanders and their sensitivity to extreme environmental conditions. Our study demonstrates that composite representations of ecologically explicit processes can provide novel insight and can better explain genetic differentiation than ecologically implicit landscape resistance surfaces. Additionally, our study underscores the fact that spatial estimates of habitat suitability or abundance may not serve as adequate proxies for describing gene flow, as predicted abundance was a poor predictor of genetic differentiation.

Effects of fine-scale forest habitat quality on movement and settling decisions in juvenile pond-breeding salamanders.

A better understanding of how individuals respond to variation in habitat quality while moving through heterogeneous habitats is needed to predict ecological phenomena at larger scales, such as local population and metapopulation dynamics. We sought to identify how fine-scale habitat quality affects the decisions of juvenile pond-breeding salamanders (Ambystoma maculatum and A. annulatum) to cease dispersive movements away from their natal pond, select a refuge, and settle. Because of the acute susceptibility of juvenile amphibians to evaporative water loss in terrestrial habitats, we predicted that they possess mechanisms for adjusting their behavior in response to variations in fine-scale habitat quality. We used experimental field enclosures to isolate the effects of habitat quality on settling behavior and employed generalized linear mixed models to examine how manipulations in canopy cover (closed or open) and microhabitat (control, compacted soils, high coarse woody debris, high burrow density), along with environmental variables (rainfall and air temperature), affect the individual's probability of settling. Our results indicated that A. maculatum and A. annulatum had a 10% and 30% decreased probability of settling in open-canopy clearcut habitat, respectively, compared to closed-canopy forest habitat. In addition, A. annulatum were 24% less likely to settle in compacted soil treatments. Although the settlement probability of A. annulatum did not depend on refuge availability, A. maculatum were 18% and 25% more likely to settle under conditions of high burrow density and high coarse woody debris, respectively. These findings make a unique contribution to our understanding of amphibian movement ecology by demonstrating how the interplay of external factors and individual behavior produce observed patterns of movement and habitat selection.

Effects of subsidy quality on reciprocal subsidies: how leaf litter species changes frog biomass export.

Spatial subsidies are resources transferred from one ecosystem to another and which can greatly affect recipient systems. Increased subsidy quantity is known to increase these effects, but subsidy quality is likely also important. We examined the effects of leaf litter quality (varying in nutrient and tannin content) in pond mesocosms on gray treefrog (Hyla versicolor) biomass export, as well as water quality and ecosystem processes. We used litter from three different tree species native to Missouri [white oak (Quercus alba), northern red oak (Quercus rubra), and sugar maple (Acer saccharum)], one non-native tree [white pine (Pinus strobus)], and a common aquatic grass [prairie cordgrass (Spartina pectinata)]. We found that leaf litter species affected almost every variable we measured. Gray treefrog biomass export was greatest in mesocosms with grass litter and lowest with white oak litter. Differences in biomass export were affected by high tannin concentrations (or possibly the correlated variable, dissolved oxygen) via their effects on survival, and by primary production, which altered mean body mass. Effects of litter species could often be traced back to the characteristics of the litter itself: leaf nitrogen, phosphorus, and tannin content, which highlights the importance of plant functional traits in affecting aquatic ecosystems. This work and others stress that changes in forest species composition could greatly influence aquatic systems and aquatic-terrestrial linkages.

Carryover effects in amphibians: are characteristics of the larval habitat needed to predict juvenile survival?

Carryover effects occur when experiences early in life affect an individual's performance at a later stage. Many studies have shown carryover effects to be important for future performance. However, it is currently unclear whether variation in later environments could overwhelm factors from an earlier life stage. We were interested in whether similar patterns would emerge under the same experimental design with similar taxa. To examine this, we implemented a four-way factorial experimental design with different forestry practices on three species of anurans (each examined in different years) in the aquatic larval environment and terrestrial juvenile environment in outdoor mesocosms in central Missouri, USA. Using Cormack-Jolly-Seber mark-recapture models implemented in program MARK, we investigated whether one environment or both environments best predicted terrestrial juvenile survival. We found only limited evidence of carryover effects for one of three species in one time period. These were the effects of time to metamorphosis and body condition at metamorphosis predicting leopard frog (Lithobates sphenocephalus) survival. However, both effects were counterintuitive and/or very weak. For wood frogs (L. sylvaticus), all of the variables predicting survival had confidence intervals that included zero, but very low survival may have limited our ability to estimate parameters. The terrestrial environment was important for predicting survival in both American toads (Anaxyrus americanus) and southern leopard frogs. The partial harvest forest tended to increase survival relative to control forest and early-successional forest in American toads. Alternately, early-successional forest with downed wood removed increased survival for leopard frogs, but this treatment was no different from control forest for American toads. Previous studies have shown negative effects of recent clearcuts on terrestrial amphibians. It appears that vegetative regrowth after just a few years can mitigate these initial negative effects. Our study shows that variation in later environments probably can overwhelm variation from earlier environments. However, previous evidence of carryover effects suggests that more research is needed to predict when carryover effects are likely to occur.

Life history as a predictor of salamander recovery rate from timber harvest in southern appalachian forests, USA.

Forest management often represents a balance between social, economic, and ecological objectives. In the eastern United States, numerous studies have established that terrestrial salamander populations initially decline in abundance following timber harvest, yet the large-scale and long-term consequences are relatively unknown. We used count data from terrestrial survey points to examine the relation between salamander abundance and historic timber harvest while accounting for imperfect detection of individuals. Overall, stream- and terrestrial-breeding salamanders appeared to differ by magnitude of population decline, rate of population recovery, and extent of recolonization from surrounding forest. Specifically, estimated abundance of both species groups was positively associated with stand age and recovery rates were predicted to increase over time for red-legged salamanders (Plethodon shermani) and decrease in stream-breeding species. Abundance of stream-breeding salamanders was predicted to reach a peak by 100 years after timber harvest, and the population growth rate of red-legged salamanders was predicted to undergo a significant increase 100 years after harvest. Estimated abundance of stream-breeding salamanders in young forest stands was also negatively associated with the distance to adjacent forest, a result that suggests immigration has a role in the recovery of these species. Our results indicate that salamander abundance in young forest stands may be only modestly lower than in more mature forest but that full recovery from timber harvest may take a substantial amount of time and that species life history may affect patterns of recovery. Historia de Vida como un Vaticinador de la Tasa de Recuperación de una Salamandra a la Colecta de Madera en los Bosques del Sur de los Apalaches, E.U.A.

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.

Reciprocal subsidies in ponds: does leaf input increase frog biomass export?

Reciprocal subsidies occur when ecosystems are paired, both importing and exporting resources to each other. The input of subsidies increases reciprocal subsidy export, but it is unclear how this changes with other important factors, such as ambient resources. We provide a conceptual framework for reciprocal subsidies and empirical data testing this framework using a pond-forest system in Missouri, USA. Our experiment used in situ pond mesocosms and three species of anurans: wood frogs, American toads, and southern leopard frogs. We predicted that increases in ambient resources (primary productivity) and detrital subsidy input (deciduous tree leaves) into pond mesocosms would increase reciprocal export (frog biomass) to the surrounding terrestrial ecosystem. In contrast, we found that increases in primary productivity consistently decreased frog biomass, except with leaf litter inputs. With leaf inputs, primary productivity did not affect the export of frogs, indicating that leaf detritus and associated microbial communities may be more important than algae for frog production. We found that subsidy inputs tended to increase reciprocal exports, and thus partial concordance with our conceptual framework.

Intersex gonads in frogs: understanding the time course of natural development and role of endocrine disruptors.

The paucity of data on sexual development of anuran amphibians has played an important role in the recent controversy over atrazine exposure. Although some studies have demonstrated the presence of abnormal gonads in control treatments, others have not, leading to varying interpretations of the effects of atrazine exposure on sexual development. However, the timing of development varies among anuran amphibians such that, at any snapshot in time, different species may exhibit different stages of sexual differentiation. We examined three species representing each of the differentiation rates (Bufo americanus=retarded rate; Hyla versicolor=basic rate; Rana sphenocephala=accelerated rate), to examine the natural time course of sexual development along with the influence of atrazine exposure. For each species, exposure to atrazine (1, 3, 10, 30 parts per billion), 17-beta-estradiol or control water occurred throughout larval life. Gonad histology was performed at 3-week intervals during the larval period or at a juvenile stage to examine the proportion of males, females, underdeveloped testes, testicular oocytes (TO; testes with 0-30% oocytes), and ovotestes (OVTs; testes with>30% oocytes). Our results illustrate that a phase of intersex gonads (TO or OVT) is normal during R. sphenocephala sexual development, a species representing the accelerated differentiation rate. Further, intersex gonads were found in juvenile stages of B. americanus and H. versicolor, representing retarded and basic rates, respectively, suggesting that a phase of intersex may be common regardless of differentiation rate. Moreover, these data highlight the importance of longitudinal studies rather than snapshots in time.

Larval responses of three midwestern anurans to chronic, low-dose exposures of four herbicides.

Low levels of agricultural herbicides often contaminate surface water and might persist throughout the growing season, potentially acting as stressors on aquatic organisms. Although low-dose, chronic exposures to agrochemicals are likely common for many nontarget organisms, studies addressing these effects using end-use herbicide formulations are rare. We exposed three common species of tadpoles to conservative levels of atrazine, S-metolachlor, and glyphosate end-use herbicide formulations throughout the larval period to test for survival differences or life-history trait alterations. Exposure to the glyphosate product Roundup WeatherMax at 572 ppb glyphosate acid equivalents (a.e.) resulted in 80% mortality of western chorus frog tadpoles, likely as a result of a unique surfactant formulation. Exposure to WeatherMax or Roundup Original Max at 572 ppb a.e. also lengthened the larval period for American toads. Chronic atrazine and S-metolachlor exposures induced no significant negative effects on survival, mass at metamorphosis, or larval period length at the levels tested. These results highlight the importance of explicitly tying chronic tests to the natural environment and considering contributions of surfactant/adjuvant components to end-use formulation toxicities, even between very similar products.

Survival costs associated with wood frog breeding migrations: effects of timber harvest and drought.

Migration presents a trade-off for individuals between the potential fitness benefits of reaching high-quality habitat vs. the potential costs of migration. Within an information-theoretic framework, we examined the costs of migration for adult wood frogs (Rana sylvatica) in response to timber harvest and annual weather conditions using Cox proportional-hazard estimates of survival. In 2004 prior to timber harvest, survival did not differ between the inside (0.75, SE = 0.078) and outside (0.73, SE = 0.235) of the circular timber harvest arrays (each 164 m radius). Following timber harvest, survival inside arrays in both 2005 and 2006 (0.22, SE = 0.065; 0.42, SE = 0.139) was lower than survival outside of the arrays and prior to harvest. Sources of mortality included predation in all years and desiccation in the drought year of 2005. The most-supported models for explaining both predation and desiccation risks reflected behaviors as opposed to timber harvest or weather conditions. Both predation and desiccation risks increased when frogs made frequent movements or were located near breeding ponds. Optimal behaviors for reducing predation and desiccation risks were the same before and after timber harvest; however, the survival consequences for not adopting these behaviors were more severe following harvest. Our results provide empirical evidence for (1) the ecological pressures that influence migratory behavior and (2) differential survival in relation to migratory behavior which reveals why frogs move relatively long distances away from breeding sites.

Post-Pleistocene differentiation in a Central Interior Highlands endemic salamander.

AIM: For many endemic species with limited dispersal capacities, the relationship between landscape changes and species distributions is still unclear. We characterized the population structure of the endemic ringed salamander (Ambystoma annulatum) across its distribution in the Central Interior Highlands (CIH) of North America, an area of high species endemism, to infer the ecological and evolutionary history of the species. METHODS: We sampled 498 individuals across the species distribution and characterized the population genetic structure using nuclear microsatellite and mitochondrial DNA (mtDNA) markers. RESULTS: Ambystoma annulatum exist in two strongly supported nuclear genetic clusters across the CIH that correspond to a northern cluster that includes the Missouri Ozark populations and a southern cluster that includes the Arkansas and Oklahoma Ozarks and the Ouachita Mountains. Our demographic models estimated that these populations diverged approximately 2,700 years ago. Pairwise estimates of genetic differentiation at microsatellite and mtDNA markers indicated limited contemporary gene flow and suggest that genetic differentiation was primarily influenced by changes in the post-Pleistocene landscape of the CIH. MAIN CONCLUSIONS: Both the geologic history and post-European settlement history of the CIH have influenced the population genetic structure of A. annulatum. The low mtDNA diversity suggests a retraction into and expansion out of refugial areas in the south-central Ozarks, during temperature fluctuations of the Pleistocene and Holocene epochs. Similarly, the estimated divergence time for the two nuclear clusters corresponds to changes in the post-Pleistocene landscape. More recently, decreased A. annulatum gene flow may be a result of increased habitat fragmentation and alteration post-European settlement.

Effects of timber harvesting on pond-breeding amphibian persistence: testing the evacuation hypothesis.

Numerous studies have documented the decline of amphibians following timber harvest. However, direct evidence concerning the mechanisms of population decline is lacking and hinders attempts to develop conservation or recovery plans and solutions for forest species. We summarized the mechanisms by which abundance of amphibians may initially decline following timber harvest into three testable hypotheses: (1) mortality, (2) retreat, and (3) evacuation. Here, we tested the evacuation hypothesis within a large-scale, replicated experiment. We used drift fences with pitfall traps to capture pond-breeding amphibians moving out of experimental clearcut quadrants and into control quadrants at four replicate arrays located within the Daniel Boone Conservation Area on the upper Ozark Plateau in Warren County, Missouri, USA. During the preharvest year of 2004, only 51.6% of the 312 individuals captured were moving out of pre-clearcut quadrants, and movement did not differ from random. In contrast, during both postharvest years of 2005 and 2006, the number of captures along the quadrant edge increased, and a higher proportion of individuals (59.9% and 56.6%, respectively, by year) were moving out of clearcut quadrants than entering. Salamanders moved out of clearcuts in large percentages (Ambystoma annulatum, 78.2% in 2005, 78.2% in 2006; A. maculatum, 64.0% in 2005, 57.1% in 2006). Frogs and toads also moved out of clearcut quadrants, but in lower percentages (Bufo americanus, 59.6% in 2005, 53.3% in 2006; Rana clamitans, 52.7% in 2006). Salamanders moved out of clearcuts with low-wood treatments more than out of clearcuts with high-wood treatments. Movement of salamanders out of clearcuts was independent of sex. Estimated movement out of clearcuts represented between 8.7% and 35.0% of the total breeding adults captured for two species of salamanders. Although we recognize that some portion of the amphibian population may retreat underground for short periods and others may not survive the effects of timber harvest, these data are the first direct evidence showing that individuals are capable of leaving clearcuts and shifting habitat use.

Suitability of golf course ponds for amphibian metamorphosis when bullfrogs are removed.

Managing areas designed for human recreation so that they are compatible with natural amphibian populations can reduce the negative impacts of habitat destruction. We examined the potential for amphibians to complete larval development in golf course ponds in the presence or absence of overwintered bullfrog tadpoles (Rana catesbeiana), which are frequently found in permanent, human-made ponds. We reared larval American toads (Bufo americanus), southern leopard frogs (R. sphenocephala), and spotted salamanders (Ambystoma maculatum) with 0 or 5 overwintered bullfrog tadpoles in field enclosures located in ponds on golf courses or in experimental wetlands at a reference site. Survival to metamorphosis of American toads, southern leopard frogs, and spotted salamanders was greater in ponds on golf courses than at reference sites. We attributed this increased survival to low abundance of insect predators in golf course ponds. The presence of overwintered bullfrogs, however, reduced the survival of American toads, southern leopard frogs, and spotted salamanders reared in golf course ponds, indicating that the suitability of the aquatic habitats for these species partly depended on the biotic community present. Our results suggest that ponds in human recreational areas should be managed by maintaining intermediate hydroperiods, which will reduce the presence of bullfrog tadpoles and predators, such as fish, and which may allow native amphibian assemblages to flourish.