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.

Phylogenetic patterns of trait and trait plasticity evolution: Insights from amphibian embryos.

Environmental variation favors the evolution of phenotypic plasticity. For many species, we understand the costs and benefits of different phenotypes, but we lack a broad understanding of how plastic traits evolve across large clades. Using identical experiments conducted across North America, we examined prey responses to predator cues. We quantified five life-history traits and the magnitude of their plasticity for 23 amphibian species/populations (spanning three families and five genera) when exposed to no cues, crushed-egg cues, and predatory crayfish cues. Embryonic responses varied considerably among species and phylogenetic signal was common among the traits, whereas phylogenetic signal was rare for trait plasticities. Among trait-evolution models, the Ornstein-Uhlenbeck (OU) model provided the best fit or was essentially tied with Brownian motion. Using the best fitting model, evolutionary rates for plasticities were higher than traits for three life-history traits and lower for two. These data suggest that the evolution of life-history traits in amphibian embryos is more constrained by a species' position in the phylogeny than is the evolution of life history plasticities. The fact that an OU model of trait evolution was often a good fit to patterns of trait variation may indicate adaptive optima for traits and their plasticities.

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.

The influence of breeding phenology on the genetic structure of four pond-breeding salamanders.

Understanding metapopulation dynamics requires knowledge about local population dynamics and movement in both space and time. Most genetic metapopulation studies use one or two study species across the same landscape to infer population dynamics; however, using multiple co-occurring species allows for testing of hypotheses related to different life history strategies. We used genetic data to study dispersal, as measured by gene flow, in three ambystomatid salamanders (Ambystoma annulatum, A. maculatum, and A. opacum) and the Central Newt (Notophthalmus viridescens louisianensis) on the same landscape in Missouri, USA. While all four salamander species are forest dependent organisms that require fishless ponds to reproduce, they differ in breeding phenology and spatial distribution on the landscape. We use these differences in life history and distribution to address the following questions: (1) Are there species-level differences in the observed patterns of genetic diversity and genetic structure? and (2) Is dispersal influenced by landscape resistance? We detected two genetic clusters in A. annulatum and A. opacum on our landscape; both species breed in the fall and larvae overwinter in ponds. In contrast, no structure was evident in A. maculatum and N. v. louisianensis, species that breed during the spring. Tests for isolation by distance were significant for the three ambystomatids but not for N. v. louisianensis. Landscape resistance also contributed to genetic differentiation for all four species. Our results suggest species-level differences in dispersal ability and breeding phenology are driving observed patterns of genetic differentiation. From an evolutionary standpoint, the observed differences in dispersal distances and genetic structure between fall breeding and spring breeding species may be a result of the trade-off between larval period length and size at metamorphosis which in turn may influence the long-term viability of the metapopulation. Thus, it is important to consider life history differences among closely related and ecologically similar species when making management decisions.

Stoichiometry and Life-History Interact to Determine the Magnitude of Cross-Ecosystem Element and Biomass Fluxes.

Ecosystems are linked through the transfer of materials and energy. Studies examining material fluxes across habitat boundaries frequently quantify unidirectional flows of nutrients and energy. However, material fluxes can be multidirectional, and we lack a conceptual framework to describe how their quantity and stoichiometry influence the net transfer of individual elements between ecosystems. Here we develop a zero net transfer isocline (ZNTI) framework that integrates the relative mass and stoichiometry of fluxes into and out of an ecosystem. We then use case studies with amphibians and salmon to elucidate how life history, ontogenetic shifts in stoichiometry, and trophic interactions shape relative fluxes of nutrients between aquatic and terrestrial ecosystems. Because they increase in both size and Ca content from ova to metamorphs, amphibian life histories strongly bias them toward net Ca export into the terrestrial environment. Because amphibian biomass, C, P, and Ca ZNTIs do not overlap, there is no value of survivorship where the net flux of biomass, C, P, and Ca are simultaneously balanced between terrestrial and aquatic habitats. The degree of iteroparity and semelparity in salmon strongly affects both the magnitude of net biomass and P flux between riverine and marine environments. While the net direction of biomass flux generally remains strongly biased toward import into the riverine system, net P flux can reach net export into the marine environment because of increasing adult breeding survival leading to reduced mass and %P of what they deposit in rivers (e.g., ova vs. whole carcasses). These examples highlight how ontogenetic shifts in body size and stoichiometry result in asymmetric fluxes of elements and biomass that can lead to simultaneous net imports and exports of different elements within the same system. Furthermore, they demonstrate how changes in life-history characteristics and stage-specific survivorship can lead to changes in net elemental transport between ecosystems.

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.

Reducing bias in population and landscape genetic inferences: the effects of sampling related individuals and multiple life stages.

In population or landscape genetics studies, an unbiased sampling scheme is essential for generating accurate results, but logistics may lead to deviations from the sample design. Such deviations may come in the form of sampling multiple life stages. Presently, it is largely unknown what effect sampling different life stages can have on population or landscape genetic inference, or how mixing life stages can affect the parameters being measured. Additionally, the removal of siblings from a data set is considered best-practice, but direct comparisons of inferences made with and without siblings are limited. In this study, we sampled embryos, larvae, and adult Ambystoma maculatum from five ponds in Missouri, and analyzed them at 15 microsatellite loci. We calculated allelic richness, heterozygosity and effective population sizes for each life stage at each pond and tested for genetic differentiation (F ST and D C ) and isolation-by-distance (IBD) among ponds. We tested for differences in each of these measures between life stages, and in a pooled population of all life stages. All calculations were done with and without sibling pairs to assess the effect of sibling removal. We also assessed the effect of reducing the number of microsatellites used to make inference. No statistically significant differences were found among ponds or life stages for any of the population genetic measures, but patterns of IBD differed among life stages. There was significant IBD when using adult samples, but tests using embryos, larvae, or a combination of the three life stages were not significant. We found that increasing the ratio of larval or embryo samples in the analysis of genetic distance weakened the IBD relationship, and when using D C , the IBD was no longer significant when larvae and embryos exceeded 60% of the population sample. Further, power to detect an IBD relationship was reduced when fewer microsatellites were used in the analysis.

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.

Intermediate pond sizes contain the highest density, richness, and diversity of pond-breeding amphibians.

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

Partitioning detectability components in populations subject to within-season temporary emigration using binomial mixture models.

Detectability of individual animals is highly variable and nearly always < 1; imperfect detection must be accounted for to reliably estimate population sizes and trends. Hierarchical models can simultaneously estimate abundance and effective detection probability, but there are several different mechanisms that cause variation in detectability. Neglecting temporary emigration can lead to biased population estimates because availability and conditional detection probability are confounded. In this study, we extend previous hierarchical binomial mixture models to account for multiple sources of variation in detectability. The state process of the hierarchical model describes ecological mechanisms that generate spatial and temporal patterns in abundance, while the observation model accounts for the imperfect nature of counting individuals due to temporary emigration and false absences. We illustrate our model's potential advantages, including the allowance of temporary emigration between sampling periods, with a case study of southern red-backed salamanders Plethodon serratus. We fit our model and a standard binomial mixture model to counts of terrestrial salamanders surveyed at 40 sites during 3-5 surveys each spring and fall 2010-2012. Our models generated similar parameter estimates to standard binomial mixture models. Aspect was the best predictor of salamander abundance in our case study; abundance increased as aspect became more northeasterly. Increased time-since-rainfall strongly decreased salamander surface activity (i.e. availability for sampling), while higher amounts of woody cover objects and rocks increased conditional detection probability (i.e. probability of capture, given an animal is exposed to sampling). By explicitly accounting for both components of detectability, we increased congruence between our statistical modeling and our ecological understanding of the system. We stress the importance of choosing survey locations and protocols that maximize species availability and conditional detection probability to increase population parameter estimate reliability.

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.

Effects of tannin source and concentration from tree leaves on two species of tadpoles.

Vegetation in and around freshwater ecosystems can affect aquatic organisms through the production of secondary compounds, which are retained in leaves after senescence and are biologically active. Tannins can be toxic to tadpoles, but the plant source of tannins and tannin concentration have been confounded in experimental designs in previous studies. To examine the effects of the concentration and source of tannins (tree species), we examined the effects of 4 factors on tadpole survival, growth, and development: tannin source (red oak [Quercus rubra], white oak [Quercus alba], or sugar maple [Acer saccharum]); tannin concentration (including a control); diet protein level; and tadpole species (American toad [Anaxyrus americanus] and spring peepers [Pseudacris crucifer]). Tannin source and concentration affected spring peeper survival, but American toads had uniformly high survival. Spring peepers had a lower survival rate in high tannin concentrations of oak leachate but a high survival rate in both concentrations of sugar maple leachate. These differences in survival did not correspond with changes in dissolved oxygen, and no effect of dietary protein level on tadpole performance was observed. The presence of plant leachate resulted in increased tadpole growth in both species, but the mechanism for this finding is unclear. The results of the present study show that tannin concentration and source are important factors for tadpole performance, adding further evidence that plant chemistry can affect aquatic organisms.

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 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.

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.

Context-dependent movement behavior of woodland salamanders (Plethodon) in two habitat types.

Animal movement is critical to the maintenance of functional connectivity at the landscape scale and can play a key role in population persistence and metapopulation dynamics. The permeability of habitat to animal movement may vary as a result of either differential mortality, physical resistance, or simply the behavioral responses of organisms to perceived habitat quality. Understanding how and when animal movement behavior varies among habitat types is critical for identifying barriers to dispersal and predicting species distributions in relation to landscape features. We conducted an experimental translocation study and compared the movement success and behavioral strategies of plethodontid salamanders in both forest and open-canopy habitat. We found that individuals in closed-canopy forest oriented more strongly towards their home ranges and moved significantly farther on their release night. In spite of the clear differences in movement paths, the ultimate movement success of homing salamanders did not appear to vary with habitat type. Our study contributes to a growing body of literature suggesting the importance of recognizing the context dependence of animal movement behavior. Because the movement rates of displaced salamanders were significantly reduced in open-canopy, dispersal rates of plethodontid salamanders in open-canopy habitat are likely lower than in control forest. Further mechanistic studies focusing on habitat-specific movement behavior and survival costs will be valuable for effectively identifying and mitigating barriers to animal movement.

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.