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Copper (Cu) appears to be consistently more toxic to anuran species relative to other vertebrate taxa. There are limited Cu toxicity data for salamanders; of the few studies conducted on salamanders, most examined Cu effects on the embryonic, but not the larval, stage. We performed acute toxicity experiments, to quantify LC50s, on Harrison stage 46 larvae (free swimming hatchlings with egg yolk completely absorbed) of three ambystomatid salamander species. Each LC50 experiment used exposure concentrations of 10, 20, 30, 40, 50, and 60 µg/L with 10 replicates per concentration each containing one larva. We found very high toxicity for all species compared to previously published research on the embryo stage. Specifically, the 4-d LC50s for Ambystoma tigrinum and A. opacum were 35.3 and 18.73 µg/L, respectively. The same Cu concentrations caused similar toxicity to A. talpoideum (LC50 = 47.88 µg/L), but exposures required up to 48 d to elicit the same level of mortality. A time-to-event analysis indicated that time to mortality was significantly affected by Cu concentration. Additionally, for A. talpoideum, we observed that elevated levels of Cu decreased growth rate. Comparisons with previously reported Cu toxicity for embryos suggest that, as with fish, Cu may be more toxic to larval salamander stages than for embryos. Further, our data suggest that Cu is an important environmental contaminant that deserves increased scrutiny on the potential for population-level effects where contamination has occurred in wetlands and streams inhabited by salamanders.

Ecological stoichiometry uses information on a few key biological elements (C, N, and P) to explain complex ecological patterns. Although factors driving variation in these elements are well-established, expanding stoichiometric principles to explore dynamics of the many other essential elements comprising biological tissues (i.e., the ionome) is needed to determine their metabolic relationships and better understand biological control of elemental flows through ecosystems. 2. In this paper, we report observations of ionomic variation in two species of salamander (Ambystoma opacum and A. talpoideum) across ontogenic stages using specimens from biological collections of two wetlands sampled over a 30-year period. This unique data set allowed us to explore the extent of ionomic variation between species, among ontogenic stages, between sites, and through time. 3. We found species- and to a lesser extent site-specific differences in C, N, and P along with 13 other elements forming salamander ionomes but saw no evidence of temporal changes. Salamander ionomic composition was most strongly related to ontogeny with relatively higher concentrations of many elements in adult males (i.e., Ca, P, S, Mg, Zn, and Cu) compared to metamorphic juveniles, which had greater amounts of C, Fe, and Mn. 4. In addition to patterns of individual elements, covariance among elements was used to construct multi-elemental nutrient balances, which revealed differences in salamander elemental composition between species and sites and changes in elemental proportions across ontogenic development. These multi-elemental balances distinguished among species-site-ontogenic stage groups better than using only C, N, and P. 5. Overall, this study highlights the responsiveness of consumer ionomes to life-history and environmental variation while reflecting underlying relationships among elements tied to biological function. As such, ionomic studies can provide important insights into factors shaping consumer elemental composition and for predicting how these changes might affect higher-order ecological processes.

Chemical contamination is often suggested as an important contributing factor to amphibian population declines, but direct links are rarely reported. Population modeling provides a quantitative method to integrate toxicity data with demographic data to understand the long-term effects of contaminants on population persistence. In this study we use laboratory-derived embryo and larval toxicity data for two anuran species to investigate the potential for toxicity to contribute to population declines. We use the southern toad (Anaxyrus terrestris) and the southern leopard frog (Lithobates sphenocephalus) as model species to investigate copper (Cu) toxicity. We use matrix models to project populations through time and quantify extinction risk (the probability of quasi-extinction in 35 yr). Life-history parameters for toads and frogs were obtained from previously published literature or unpublished data from a long-term (>35 yr) data set. In addition to Cu toxicity, we investigate the role of climate change on amphibian populations by including the probability of early pond drying that results in catastrophic reproductive failure (CRF, i.e., complete mortality of all larval individuals). Our models indicate that CRF is an important parameter for both species as both were unable to persist when CRF probability was >50% for toads or 40% for frogs. Copper toxicity alone did not result in significant effects on extinction risk unless toxicity was very high (>50% reduction in survival parameters). For toads, Cu toxicity and high probability of CRF both resulted in high extinction risk but no synergistic (or greater than additive) effects between the two stressors occurred. For leopard frogs, in the absence of CRF survival was high even under Cu toxicity, but with CRF Cu toxicity increased extinction risk. Our analyses highlight the importance of considering multiple stressors as well as species differences in response to those stressors. Our models were consistently most sensitive to juvenile and adult survival, further suggesting the importance of terrestrial stages to population persistence. Future models will incorporate multiple wetlands with different combinations of stressors to understand if our results for a single wetland result in a population sink within the landscape.

Little is known about the propensity of crocodilians to bioaccumulate trace elements as a result of chronic dietary exposure. We exposed 36 juvenile alligators (Alligator mississippiensis) to one of four dietary treatments that varied in the relative frequency of meals containing prey from coal combustion waste (CCW)-contaminated habitats vs. prey from uncontaminated sites, and evaluated tissue residues and growth rates after 12 mo and 25 mo of exposure. Hepatic and renal concentrations of arsenic (As), cadmium (Cd) and selenium (Se) varied significantly among dietary treatment groups in a dose-dependent manner and were higher in kidneys than in livers. Exposure period did not affect Se or As levels but Cd levels were significantly higher after 25 mo than 12 mo of exposure. Kidney As and Se levels were negatively correlated with body size but neither growth rates nor body condition varied significantly among dietary treatment groups. Our study is among the first to experimentally examine bioaccumulation of trace element contaminants in crocodilians as a result of chronic dietary exposure. A combination of field surveys and laboratory experiments will be required to understand the effects of different exposure scenarios on tissue residues, and ultimately link these concentrations with effects on individual health.

Many metals are acutely toxic to aquatic organisms at high concentrations and for some metals, such as copper (Cu), even low-level chronic contamination may be cause for conservation concern. Amphibian susceptibility to Cu has been examined in only a few species, and susceptibility is highly variable. The lethal and sublethal effects were examined of chronic aqueous Cu exposure on embryonic and larval eastern narrowmouth toads, Gastrophryne carolinensis. Copper levels as low as 10 μg Cu/L reduced embryonic and larval survival. Embryonic survivorship varied within- and between-source populations, with embryos derived from uncontaminated-wetland parents having greater survival at lower Cu levels than embryos from parents from a metal-contaminated constructed wetland. At 30 μg/L, embryos from the contaminated site had greater survival. Overall survival from oviposition to metamorphosis was 68.9% at 0 μg/L and 5.4% at 10 μg/L. Similarly, embryos exposed to ≥50 μg/L demonstrated developmental delays in transition from embryo to free-swimming larva. These results demonstrate a negative population-specific response to environmentally relevant levels of Cu. Environ Toxicol Chem 2015;34:575–582. © 2014 SETAC

Stable isotope enrichment techniques can aid in understanding dispersal of animals. Pond-breeding amphibians often have spatially disjunct populations that depend on immigration for persistence, yet obtaining direct estimates of dispersal rates among wetlands is challenging. We enriched aquatic mesocosms with 15N to “mark” amphibian larvae and determine the feasibility of using enrichment techniques to study dispersal in pond-breeding amphibians. Because newly metamorphosed amphibians in mark-release-recapture studies may not be recaptured until adulthood, we estimated persistence of the 15N enrichment signature up to 3 years post-metamorphosis. We reared larval marbled salamanders (Ambystoma opacum) in artificial mesocosms dosed with 15NH4Cl, and maintained metamorphs on unlabeled prey for 7 months to estimate the biological half life (BHL) of 15N in tissue. Metamorphs in spiked treatments attained δ15N levels >1000 times higher than reference animals (5 ± 1‰), and levels remained ~225 times higher than controls after 7 months. The average 15N BHL was 2.49 ± 0.24 months, indicating that the elevated signature should be discernible for a minimum of 20–28 months after metamorphosis. Our results suggest that 15N enrichment is feasible for field studies of amphibian dispersal, as metamorphs will retain isotope-enriched tissues that persist until at least the second year of breeding.

The creation of wetlands, such as urban and industrial ponds, has increased in recent decades, and these wetlands often become enriched in pollutants over time. One metal contaminant trapped in created wetlands is copper (Cu2+). Copper concentrations in sediments and overlying water may affect amphibian species that breed in created wetlands. The authors analyzed the Cu concentration in dried sediments from a contaminated wetland and the levels of aqueous Cu released after flooding the sediments with different volumes of water, mimicking low, medium, and high pond-filling events. Eggs and larvae of Ambystoma opacum Gravenhorst, a salamander that lays eggs on the sediments in dry pond beds that hatch on pond-filling, were exposed to a range of Cu concentrations that bracketed potential aqueous Cu levels in created wetlands. Embryo survival varied among clutches, but increased Cu levels did not affect embryo survival. At Cu concentrations of 500 µg/L or greater, however, embryos hatched earlier, and the aquatic larvae died shortly after hatching. Because Cu concentrations in sediments increase over time in created wetlands, even relatively tolerant species such as A. opacum may be affected by Cu levels in the posthatching environment. Environ Toxicol Chem 2014;33:1631–1637. © 2014 SETAC

Terrestrial habitats surrounding isolated wetlands are a critical resource for many pond-breeding amphibian species, yet few studies have examined the terrestrial distribution of post-metamorphic juveniles and adults. We used an encircling drift fence at a breeding pond in conjunction with partial fences at 90, 172, and 332 m from the wetland to estimate the terrestrial distribution of adult marbled salamanders (Ambystoma opacum; four breeding seasons) and mole salamanders (A. talpoideum; two seasons), as well as the dispersion of newly metamorphosed A. opacum (one summer). For newly metamorphosed A. opacum, 79% emigrated ,90 m from the wetland, and 8% moved beyond 172 m; movement distance was unrelated to body size. Distribution of adult A. opacum varied among years, with an average of 28% (range 23-31%) occurring beyond 172 m in all years. Averaged across two years, 51% of adult A. talpoideum occurred beyond 172 m. Lognormal models provided a good fit to both the juvenile and adult ambystomatid distributions, and parameters differed between age classes, sexes, species, and years within species. For adult A. opacum a buffer radius of 300 m or 340 m, depending on the year, is estimated to include 95% of adults; for A. talpoideum the estimate is 464 m or 501 m. A reanalysis of distribution data for seven ambystomatid species shows that a previous estimate of a 164-m radius to protect 95% of a population underestimates the needed buffer radius by 185 m. Because our study wetland requires a nearly 500 m wide radius to protect 95% of its ambystomatid adults, preservation of similar communities may require much more surrounding terrestrial habitat than previously thought.

Environmental contaminants are implicated in the global decline of amphibian populations. Copper (Cu) is a widespread contaminant that can be toxic at concentrations just above the normal physiological range. In the present study we examined the effects of chronic Cu aqueous exposure on embryos and larvae of southern toads, Anaxyrus (Bufo) terrestris. Measurable levels of Cu were found in larvae, with tissue concentrations up to 27.5 μg Cu/g dry mass. Aqueous concentrations of Cu as low as 10 μg/L significantly reduced survival to the free-swimming stage and no larvae reached metamorphosis at concentrations above 15 μg/L. Clutches from populations with prior Cu exposure had the lowest survivorship. Among several populations there was significant variation in survivorship at different levels of Cu. More data are needed to understand the underlying causes of within- and among-population resilience to anthropogenic stressors.

In the southeastern U.S., changes in temperature and precipitation over the last three decades have been the most dramatic in winter and spring seasons. Continuation of these trends could negatively impact pond-breeding amphibians, especially those that rely on winter and spring rains to fill seasonal wetlands, trigger breeding, and ensure reproductive success. From 2009 to 2012, we monitored Spring and Fall presence of aquatic stages (larval and paedomorphic, gilled adult) of a winter-breeding amphibian (the mole salamander, Ambystoma talpoideum) and used multi-season models to estimate occupancy, local colonization and extinction. Seasonal estimates of occupancy, corrected for imperfect detection, declined from 22.3 % of ponds in Spring 2009 to 9.9 % in Fall 2012. Our best supported model suggested that changes in occupancy were driven by increased rates of extinction that corresponded with drought-related drying of ponds. Based on uncertainty in climate change projections for the Southeast, we present a conceptual model of predicted changes in wetland hydroperiods across a landscape with projected decreases and increases in future precipitation. Such precipitation changes could alter wetland hydroperiods, facilitate extinctions of species adapted to short, intermediate or long hydroperiod environments and, ultimately, modify the composition of amphibian communities within freshwater wetland ecosystems.

Bioaccumulation of contaminants and subsequent maternal transfer to offspring are important factors that affect the reproductive success of wildlife. However, maternal transfer of contaminants has rarely been investigated in amphibians. We examined maternal transfer of trace elements in southern toads ( Bufo[Anaxyrus] terrestris) residing in two locations: (1) an active coal combustion waste (CCW) disposal basin and adjacent 40-ha floodplain contaminated with CCW over 35 years ago and (2) an uncontaminated reference site. Our study is among the few to document tissue concentration-dependent maternal transfer of contaminants and associated adverse effects in amphibians. We found that females collected from the CCW-contaminated area had elevated concentrations of Ni, Se, and Sr; these females also transferred elevated levels of Cu, Pb, Se, and Sr to their eggs compared to females from the reference site. Overall reproductive success, estimated as a function of clutch size and offspring viability, was reduced by 27% in clutches collected from parents from the contaminated site compared to the reference site. Offspring viability negatively correlated with female and/or egg concentrations of Se and Ni. Reproductive success negatively correlated with Se and Cu concentrations in females, and Se concentrations in eggs. Our study highlights how exposure to CCW can negatively affect amphibian reproduction.

Natural and anthropogenic stressors typically do not occur in isolation; therefore, understanding ecological risk of contaminant exposure should account for potential interactions of multiple stressors. Realistically, common contaminants can also occur chronically in the environment. Because parental exposure to stressors may cause transgenerational effects on offspring, affecting their ability to cope with the same or novel environmental stressors, the exposure histories of generations preceding that being tested should be considered. To examine multiple stressor and parental exposure effects we employed a 2 × 2 × 2 factorial design in outdoor 1000-L mesocosms (n = 24). Larval southern toads (Anaxyrus terrestris), bred from parents collected from reference and metal-contaminated sites, were exposed to two levels of both an anthropogenic (copper—0, 30 µg/L Cu) and natural (predator cue − present/absent) stressor and reared to metamorphosis. Toads from the metal-contaminated parental source population were smaller at metamorphosis and had delayed development; i.e., a prolonged larval period. Similarly, larval Cu exposure also reduced size at metamorphosis and prolonged the larval period. We, additionally, observed a significant interaction between larval Cu and predator-cue exposure on larval period, wherein delayed emergence was only present in the 30-µg/L Cu treatments in the absence of predator cues. The presence of parental effects as well as an interaction between aquatic stressors on commonly measured endpoints highlight the importance of conducting multistressor studies across generations to obtain data that are more relevant to field conditions in order to determine population-level effects of contaminant exposure.

Exposure to environmental contaminants contributes to the global decline of amphibian populations. The impacts of organic contaminants on amphibians are well documented. However, substantially less is known concerning the potential effects of metals on amphibian populations. Copper (Cu) is an essential element, but it can be toxic at concentrations only slightly higher than the normal physiological range. The present study examines the effects of chronic Cu exposure on embryos and larvae of southern leopard frogs, Lithobates (Rana) sphenocephalus. Groups of eggs from multiple clutches were collected from two wetlands and exposed to a range of Cu concentrations (0-150 mg/L) until they reached the free-swimming stage, and then individual larvae were reared to metamorphosis. Higher Cu concentrations significantly reduced embryo survival to the free-swimming stage but did not further reduce survival to metamorphosis. Larval period was affected by Cu treatment, but the clutch from which larvae originated (i.e., parentage) explained a higher proportion of the variation. Embryo survival to hatching varied significantly among clutches, ranging from 42.9 to 79.2%. Measurable levels of Cu were found in larvae with body burdens up to 595 mg Cu/g dry mass in the 100 mg/L treatment, and larval Cu body burdens were higher than in metamorphs. The present study also demonstrated that higher initial egg density ameliorated embryo mortality at higher Cu levels and should be accounted for in future studies. Environ. Toxicol. Chem. 2012;31:1587-1594. # 2012 SETAC Keywords—Lithobates (Rana) sphenocephalus Copper Amphibian Ecotoxicology Metal toxicity

We conducted a mesocosm study to assess the individual and interactive effects of previous maternal exposure and larval exposure to trace element-laden sediments on southern toads (Bufo terrestris). Previous maternal exposure to coal combustion wastes (CCW) reduced larval survival to metamorphosis up to 57% compared to larvae of unexposed females. Larvae reared on CCW accumulated significant concentrations of trace elements resulting in extended larval periods, reduced growth rates, and reduced mass at metamorphosis. However, the effects were dependent on age of sediments, suggesting the effects of contaminants from CCW may be partially ameliorated over time through the reduced bioavailability of trace elements in aged CCW. Most importantly, maternal exposure to contaminants coupled with larval exposure to fresh CCW interacted to reduce survival to metamorphosis by 85% compared to reference conditions. Our study yields further evidence that disposal of CCW in aquatic basins potentially creates ecological traps for some amphibian populations.

Contaminants often occur as mixtures in the environment, but investigations into toxicity usually employ a single chemical. Metal contaminant mixtures from anthropogenic activities such as mining and coal combustion energy are widespread, yet relatively little research has been performed on effects of these mixtures on amphibians. Considering that amphibians tend to be highly sensitive to copper (Cu) and that metal contaminants often occur as mixtures in the environment, it is important to understand the interactive effects that may result from multiple metals. Interactive effects of Cu and zinc (Zn) on amphibians have been reported as antagonistic and, conversely, synergistic. The goal of our study was to investigate the role of Zn in Cu toxicity to amphibians throughout the embryonic developmental period. We also considered maternal effects and population differences by collecting multiple egg masses from contaminated and reference areas for use in four experiments across three species. We performed acute toxicity experiments with Cu concentrations that cause toxicity (10–200 μg/L) in the absence of other contaminants combined with sublethal concentrations of Zn (100 and 1000 μg/L). Our results suggest very few effects of Zn on Cu toxicity at these concentrations of Zn. As has been previously reported, we found that maternal effects and population history had significant influence on Cu toxicity. The explanation for a lack of interaction between Cu and Zn in this experiment is unknown but may be due to the use of sublethal Zn concentrations when previous experiments have used Zn concentrations associated with acute toxicity. Understanding the inconsistency of amphibian Cu/Zn mixture toxicity studies is an important research direction in order to create generalities that can be used to understand risk of contaminant mixtures in the environment.

In organisms that have complex life cycles, factors in the larval environment may affect both larval and adult traits. For amphibians, the postmetamorphic transition from the aquatic environment to terrestrial habitat may be a period of high juvenile mortality. We hypothesized that lipid stores at metamorphosis may affect an animal's success during this critical transition period. We examined variation in total lipid levels among years and sites in recently metamorphosed individuals of two pond-breeding salamander species, the marbled salamander (Ambystoma opacum) and the mole salamander (A. talpoideum), with limited data for one anuran species (southern leopard frog, Rana sphenocephala). Lipid levels were allometrically related to body size and ranged from 1.9 to 23.8% of body dry mass. The two salamander species differed in lipid allocation patterns, with A. opacum apportioning a higher percentage of total lipid reserves into fat bodies than A. talpoideum. Species differences in lipid allocation patterns may primarily reflect that large metamorphs will mature as one-year olds, and, regardless of species, will alter lipid compartmentalization accordingly. We used mark-recapture data obtained at drift fences encircling breeding ponds for 13 A. opacum cohorts to estimate the proportion of postmetamorphic individuals that survived to breed (age 1-4) and the mean age at first reproduction. Regression models indicated that size-corrected lipid level at metamorphosis (i.e., lipid residuals), and to a lesser extent rainfall following metamorphosis, was positively related to adult survival. Snout-vent length at metamorphosis was negatively related to age at first reproduction. We suggest that lipid stores at metamorphosis are vital to juvenile survival in the months following the transition from aquatic to terrestrial habitat, and that a trade-off shaped by postmetamorphic selection in the terrestrial habitat exists between allocation to energy stores versus structural growth in the larval environment.

Despite their potential vulnerability to contaminants from exposure at multiple life stages, amphibians are one of the least studied groups of vertebrates in ecotoxicology, and research on radiation effects in amphibians is scarce. We used multiple endpoints to assess the radiosensitivity of the southern toad (Anaxyrus [Bufo] terrestris) during its pre-terrestrial stages of development -embryonic, larval, and metamorphic. Toads were exposed, from several hours after oviposition through metamorphosis (up to 77 days later), to four low dose rates of 137Cs at 0.13, 2.4, 21, and 222 mGy d-1, resulting in total doses up to 15.8 Gy. Radiation treatments did not affect hatching success of embryos, larval survival, or the length of the larval period. The individual family variation in hatching success of embryos was larger than the radiation response. In contrast, newly metamorphosed individuals from the higher dose-rate treatments had higher mass and mass/length body indices, a measure which may relate to higher post-metamorphic survival. The increased mass and index at higher dose rates may indicate that the chronic, low dose rate radiation exposures triggered secondary responses. Additionally, the increases in growth were linked to a decrease in DNA damage (as measured by the Comet Assay) in red blood cells at a dose rate of 21 mGy d-1 and a total dose of 1.1 Gy. In conclusion, the complex effects of low dose rates of ionizing radiation may trigger growth and cellular repair mechanisms in amphibian larvae.

Amphibian populations at the Savannah River Site (SRS), South Carolina, USA, have been censused consistently for 35 years, and this provides a time series to examine the causes of population fluctuations. We examined archived museum specimens of 15 anuran species collected at wetlands on the SRS for the presence of the causative agent (Batrachochytrium dendrobatidis ) of chytridiomycosis, an emerging disease asso- ciated with population declines elsewhere. Infections were present in three out of 137 (2.18%) individuals; the pathogen was detected in two Rana catesbeianaand a single Rana sphenocephala, all collected between 1978 and 1981. Lesions were not consistent with the later stages of fatal chytridiomycosis. Analysis of population trajectories of nine amphibian species over 26 years at SRS showed that four species declined significantly over this period, including R. sphenocephala. However, we demonstrate that these declines are more likely caused by an increase in the number of years with insufficient rainfall and a shortened hydroperiod at the breeding site than by chytrid epidemics. This pattern appears to be linked to a drying trend at SRS through the 1990s, although it is unclear whether this was caused by climate change. This study demonstrates that the presence of B. dendrobatidis in am- phibian communities where some species are declining does not always implicate chytrids as a cause of the decline. Like many other emerging pathogens, the outcome of infection can vary among individuals and populations, depending on life history traits, environmental conditions, and virulence factors of the pathogen. Our report also demonstrates the use- fulness of archived museum specimens and long-term population monitoring in studying the host-parasite ecology of emerging diseases.

In aquatic communities, the phenology of breeding may influence species interactions. In the early-breeding marbled salamander, Ambystoma opacum, timing of pond filling may determine whether interactions among larvae are competitive or predatory. The objectives of our studies were to determine how time of egg hatching affected size, larval period, and survival to metamorphosis in A. opacum, and if early-hatching in A. opacum influenced the competitive and predator-prey relationships with smaller larvae of the mole salamander, Ambystoma talpoideum. Salamander larvae were reared from hatching through metamorphosis in large, outdoor enclosures located in a natural temporary pond in Aiken County, South Carolina, in two experiments. In study 1, we reared early- and late-hatching A. opacum larvae separately from hatching through metamorphosis. In study 2, we examined how early- versus late-hatching A. opacum affected a syntopic species, A. talpoideum. In general, early-hatching A. opacum were larger an...

Created ponds were built as an experiment in mitigating the loss of a wetland to construction. We monitored amphibian breeding population sizes and juvenile recruitment at these “created ponds” for 8.5 years and compared the populations to those observed at the original wetland, Sun Bay (≤600 m from the created ponds), and at an undisturbed reference wetland, Rainbow Bay. Some amphibians continued breeding migrations to Sun Bay even after it was filled with soil. Few of the anuran colonists of the created ponds had been previously captured at Sun Bay, but many of the salamander colonists had been collected. The created ponds became permanent, whereas Sun Bay and Rainbow Bay were temporary ponds. Juveniles of two salamander species and 10 species of frogs and toads metamorphosed and emigrated from the created ponds during the study. By the final years of the study, the community structure of adult and juvenile amphibians differed among the three created ponds, as well as between these ponds and the prior amphibian community at the filled wetland and the contemporaneous community at the reference wetland. Mean size at metamorphosis was smaller at the created ponds than at the reference site for two species of frogs, whereas the opposite was true for two salamanders. We conclude that the created ponds provided partial mitigation for the loss of the natural amphibian breeding habitat. Differences between the created ponds and the natural wetlands were likely related to differences in their hydrologic regimes, size, substrates, vegetation, and surrounding terrestrial habitats and to the limited availability of colonists of some species.

We report a host gender bias in haemogregarine infection characteristics in the American alligator (Alligator mississippiensis) at the Savannah River Site, South Carolina, USA. Prevalence and severity in female alligators was higher than it was in males. The reason for this pattern is not clear.

In parallel with a renewed interest in nuclear power and its possible environmental impacts, a new environmental radiation protection system calls for environmental indicators of radiological stress. However, because environmental stressors seldom occur alone, this study investigated the combined effects of an ecological stressor (larval density) and an anthropogenic stressor (ionizing radiation) on amphibians. Scaphiopus holbrookii tadpoles reared at different larval densities were exposed to four low irradiation dose rates (0.13, 2.4, 21, and 222 mGy d(-1)) from (137)Cs during the sensitive period prior to and throughout metamorphosis. Body size at metamorphosis and development rate served as fitness correlates related to population dynamics. Results showed that increased larval density decreased body size but did not affect development rate. Low dose rate radiation had no impact on either endpoint.

Climate change has had a significant impact globally on the timing of ecological events such as reproduction and migration in many species. Here, we examined the phenology of reproductive migrations in 10 amphibian species at a wetland in South Carolina, USA using a 30 year dataset. We show for the first time that two autumn-breeding amphibians are breeding increasingly later in recent years, coincident with an estimated 1.2°C increase in local overnight air temperatures during the September through February pre-breeding and breeding periods. Additionally, two winter-breeding species in the same community are breeding increasingly earlier. Four of the 10 species studied have shifted their reproductive timing an estimated 15.3 to 76.4 days in the past 30 years. This has resulted in rates of phenological change that range from 5.9 to 37.2 days per decade, providing examples of some of the greatest rates of changing phenology in ecological events reported to date. Owing to the opposing direction of the shifts in reproductive timing, our results suggest an alteration in the degree of temporal niche overlap experienced by amphibian larvae in this community. Reproductive timing can drive community dynamics in larval amphibians and our results identify an important pathway by which climate change may affect amphibian communities.

Reports of declining amphibian populations in many parts of the world are numerous, but supporting long-term census data are generally unavailable. Census data from 1979 to 1990 for three salamander species and one frog species at a breeding pond in South Carolina showed fluctuations of substantial magnitude in both the size of breeding populations and in recruitment of juveniles. Breeding population sizes exhibited no overall trend in three species and increased in the fourth. Recent droughts account satisfactorily for an increase in recruitment failures. These data illustrate that to distinguish between natural population fluctuations and declines with anthropogenic causes may require long-term studies.

Wide variation in reproductive success is common among amphibians that breed in seasonal ponds, but persistence of adults can buffer against these fluctuations, particularly for long-lived species. We hypothesized that the frequent episodes of catastrophic failure of the marbled salamander (Ambystoma opacum) enhance the importance of high terrestrial survival. At Rainbow Bay in South Carolina reproductive success was poor (< 1 metamorph/breeding female) in nearly half of the 22 years that the species bred. Complete failure occurred in 6 of 22 years. To study catastrophic failure, we adapted an age-structured, individual-based model with density-dependent growth and survival of larvae. The model was based on extensive data from local field studies and experiments. With consistently good survival in the pond stages, the simulated population required survival probabilities in the upland stages (juveniles and adults) near 0.5/year to persist and near 0.8/year to achieve the increases observed. Catastrophic failure, occurring randomly with probability 0.5/year created additional fluctuations in the population, raised the thresholds of survival required for persistence, and caused extinction under conditions that were otherwise favorable. The marbled salamander at Rainbow Bay is not at great risk of extinction because of catastrophic failure, but the risk increases dramatically if life span is decreased or frequency of failure is increased. Any reduction in terrestrial survival will have deleterious consequences by reducing the breeding populations at equilibrium, even if it does not jeopardize persistence. Our model provides assessments of risk that can be applied to poorly studied species with similar life histories, such as the endangered flatwoods salamander (A. cingulatum).

This chapter describes a 16-year study, in which it has been found that pond hydroperiod is a primary source of variation in community structure for a natural community of pond-breeding amphibians. Larval competition and predation form other axes that are positioned along a continuum of hydroperiod, and the strength of their influence on the success of species is mediated by pond hydroperiod. Although hydroperiod, competition, and predation had a detectable influence on the amphibian community at Rainbow Bay, the effects of these factors were often difficult to separate. Correlative analyses are less powerful for sorting out confounding factors than manipulative experiments in replicated artificial ponds. The predictor variables used are often themselves correlated. Nevertheless, in many cases, a good year for a species is also a good year for its competitors and predators. Juvenile recruitment of all species is limited by a short hydroperiod in the driest years. In years with longer hydroperiods, the density of competitors affected the number of metamorphosing juveniles per breeding females for some species. The density of salamander larvae is also a significant predictor of per-capita juvenile recruitment for anurans. A significant negative relationship is detected for two of four anuran species analyzed, most likely due to predation by the salamander larvae on the anuran tadpoles. These results indicate that community structure, at least in terms of relative abundances, varied continuously. However, regulation of community structure within a pond occurs through the predictable interaction of rainfall, hydroperiod, competition, and predation.

The relationships among timing of metamorphosis, size at metamorphosis, and traits related to adult fitness were studied for 8 yr in the salamander Ambystoma talpoideum at a temporary pond. Among years, the modal time of metamorphosis and mean body size at metamorphosis were positively correlated with the date the pond dried. In years that the pond dried late, one group of larvae metamorphosed well before the pond dried, whereas the other group metamorphosed just before pond drying. Mean body size of late—metamorphosing individuals was not greater than that of individuals metamorphosing early. Early—metamorphosing males and females were larger at first and second reproduction than were late—metamorphosing individuals. Independent of timing of metamorphosis, larger juveniles at metamorphosis were also larger adults at first reproduction. Age at first reproduction for males was not associated with timing of or size at metamorphosis but large early—metamorphosing females reproduced at a younger age than did small early—metamorphosing females. Neither time of metamorphosis nor size at metamorphosis was associated with survival to first reproduction. These results demonstrate a direct relationship between phenotypic variation generated in the larval stage and adult traits closely associated with an individual's fitness.

An investigayion was made into the status of the Morelet's crocodile Crocodylus moreleti Dumeril & Dumeril in Belize, a Caribbean nation of Central America. Juvenile animals were found to be rather common in isolated areas, but very few adults were observed; we believe this to be an indication of over-exploitation. Certain aspects of the hide-hunting-dealing enterprise are also examined, and it is concluded that if C. moreleti is to survive in any numbers, a policy of protection should be instituted.

Numbers of successfully metamorphosing juvenile amphibians were tabulated at three wetlands in South Carolina, U.S.A. using terrestrial drift fences with pitfall traps. A relatively undisturbed Carolina bay was studied for eight years, a partially drained Carolina bay for four years, and a man-made borrow pit for three years. Annual production of juveniles at the undisturbed Carolina bay ranged from zero to 75,644 individuals of 15 species. Fewer individuals of fewer species typically metamorphosed at the borrow pit than at the undisturbed bay, with the least numbers at the partially drained Carolina bay. Both total number and species diversity of metamorphosing juveniles at each site each year showed a strong positive correlation with hydroperiod, i.e., the number of days a site contained standing water that year. Data for one common anuran species and the most common salamander species were analyzed separately by multiple regression, in addition to the community analyses. For the mole salamander, Ambystoma talpoideum, hydroperiod was a significant predictor of the number of metamorphosing juveniles, but the number of breeding females was not. For the ornate chorus frog, Pseudacris ornata, the number of breeding females was a significant predictor of the number of metamorphosing juveniles, but hydroperiod was not. Variation in the dates of wetland filling and drying interacts with other factors to determine amphibian community structure and diversity. Either increasing or decreasing the number of days a wetland holds water could increase or decrease the number and species diversity of amphibians in and around a wetland.

The third ventricular walls and floor of male and female mink (Mustela vison) were analyzed with comparative light microscopy, transmission and scanning electron microscopy. Distinct regional fine structural variations were noted in the appearance of the dorsal, middle third and lower portions of the ventricular wall and floor. These variations were characterized by an abrupt decrease in the density of ciliated ependymal surfaces in the middle one-third of the ventricular wall. The ventral walls and floor of the lateral and infundibular recesses displayed smoother contours with numerous bleb-like protrusions and blunt villiform processes. Occasionally multipolar neuron-like cells were observed to lie upon the floor of the infundibular recess. Speculation is raised relative to the differing functional capacities of these various ventricular areas.