Dana L. Drake

A stable isotope study of a neotropical stream food web prior to the extirpation of its large amphibian community

Rapid and massive amphibian population declines have been reported throughout upland areas of the Neotropics. The abundance and species richness of Neotropical amphibian communities suggest that losses of this magnitude are likely to have strong effects at the ecosystem level. To improve understanding of the implications of their loss we used stable isotope analysis to examine trophic relationships in an ecosystem in which amphibians are dominant in a second-order forest stream at 750 m asl in Parque Nacional Omar Torrijos Herrera, Panama. We analysed δ 13 C, δ 15 N and C:N ratios of major biotic components (basal resources, invertebrates, amphibians, fish and reptiles) in the stream and of the adjacent riparian food web. Tadpoles (mean δ 15 N = 4.49%o) and adult amphibians (mean δ 15 N = 5.45‰) were intermediate links in the aquatic and terrestrial food web respectively. High δ 15 N signatures identified fish as top predators in the aquatic food web and snakes and the toad Bufo as top predators in the terrestrial food web. Isotopic signatures clearly distinguished between trophic groups of tadpoles: microbial feeders (Centrolenidae, δ 15 N range = 1).91-3.05‰), herbivores (Rana and Hyla, δ 15 N range = 4.74-5.15‰) and neuston feeders (Colostethus, δ 15 N range = 5.31-6.40%o). Dependence on autotrophic production was indicated by enriched signatures of carbon isotopes in pool dwellers versus those that reside in faster-flowing sections of the stream. High nitrogen concentrations in detrital matter (average 0.8%, C:N = 10.3) suggested that grazing tadpoles enhanced nitrogen fluxes and improved the quality of organic matter available to detritivores.

Occurrence of Oral Deformities in Larval Anurans

Abstract We quantified deformities in the marginal papillae, tooth rows, and jaw sheaths of tadpoles from 13 population samples representing three families and 11 sites in the southeastern United States. Oral deformities were observed in all samples and in 13.5–98% of the specimens per sample. Batrachochytrium dendrobatidis (chytrid) infections were detected in three samples. There was high variability among samples in the pattern and number of discovered deformities. Pairwise associations between oral structures containing deformities were nonrandom for several populations, especially those with B. dendrobatidis infections or high total numbers of deformities. Comparisons of deformities among samples using multivariate analyses revealed that tadpole samples grouped together by family. Analyses of ordination indicated that three variables, the number of deformities, the number of significant associations among deformity types within populations, and whether populations were infected with B. dendrobatidis, were significantly correlated with the pattern of deformities. Our data indicate that the incidence of oral deformities can be high in natural populations and that phylogeny and B. dendrobatidis infection exert a strong influence on the occurrence and type of oral deformities in tadpoles.

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.

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.

Predation of Eggs and Recently Hatched Larvae of Endemic Ringed Salamanders (Ambystoma annulatum) by Native and Introduced Aquatic Predators

Abstract:  Predation is a key determinant of pond community structure, yet not all predators are equally effective and not all life stages of potential prey are similarly susceptible. Understanding the effects of native and introduced species is essential to informing management strategies, especially for endemic and species of conservation concern. We examined the effects of five common predators (three native: Central Newts [Notophthalmus viridescens louisianensis], aeshnid dragonfly naiads [Aeshnidae], and Southern Leopard Frog tadpoles [Lithobates sphenocephalus]; and two introduced: Fathead Minnows [Pimephales promelas] and Mosquitofish [Gambusia affinis]) on survival of eggs and recently hatched larvae of Ringed Salamanders (Ambystoma annulatum). We also examined the effect of supplemental food or cover availability on survival at each stage. Predators primarily showed a binary response to eggs, consuming all or none of them. Supplemental food did not influence whether eggs or larvae were consumed. Larvae were consumed by all predator species although the effect varied. The presence of cover did not reduce the impacts of the other predators on larval survival. Overall, the two introduced fish species had a greater impact on survival of the early stages of Ringed Salamanders than did the native predators. Further inquiries into the susceptibility of different life stages and survival will improve conservation strategies for rare and endemic species such as Ringed Salamanders.

Microgeographic Adaptation of Wood Frog Tadpoles to an Apex Predator

Rapid adaptation of defenses can alter ecological dynamics following introduction of a new predator. We tested for local adaptation in Wood Frog (Rana sylvatica) populations that face varying selection from an apex predator, the Marbled Salamander (Ambystoma opacum), which is expanding its distribution in the study region. We performed a reciprocal transplant experiment with Wood Frog eggs and tadpoles and tested survival of tadpoles when exposed to Marbled Salamander larvae in experimental predation trials. We also evaluated life history, behavioral, and morphological trait variation with respect to origin and transplant environments. We found that tadpoles from populations exposed to high risk from Marbled Salamanders survived better when raised in high-risk environments than tadpoles from low-risk populations. However, tadpoles from high-risk environments experienced lower survival than those from low-risk environments when raised in low-risk environments. Development rate, activity, and morphology differed among populations and environments. Faster development of high-risk populations in high-risk environments and activity patterns best explained observed survival differences. These results suggest that tadpoles have evolved adaptive plasticity at microgeographic scales in response to a mosaic of varying predation risk. Fine-scaled evolution of prey survival and local gene flow could enhance the resilience of Wood Frogs to this predator expansion. As warming winters allow Marbled Salamanders to increase in abundance and distribution, the rapid and fine-scaled evolution of their prey could mediate predicted changes to temporary pond communities and ecosystems. Rapid prey evolution might often promote ecological resilience to predator introductions.

Differences in Larval Allometry among Three Ambystomatid Salamanders

Abstract Ecological and evolutionary processes commonly result in morphological variation among larval amphibians. Variation in head shape plays a critical role in both food capture and predation risk in gape-limited salamanders, yet in situ studies of head shape variability are rare outside of cannibal morph assessments. We examined allometry differences in larval head width (HW) and snout–vent length (SVL) among three sympatric species of ambystomatid salamanders from 166 ponds in Missouri, USA: Ringed Salamander (Ambystoma annulatum), Marbled Salamander (A. opacum), and Spotted Salamander (A. maculatum). We also tested whether several abiotic and biotic factors would predict HW after accounting for SVL. We found that larval HW and SVL were strongly correlated for all species but that the strength of this relationship varied among species. For early-stage larvae, Marbled Salamanders showed isometric scaling relationships, whereas both Spotted Salamanders and Ringed Salamanders were allometric. For late-stage larvae, all three species showed allometric patterns. At a small SVL, HW of small Ringed Salamanders was greater than the other species. As larvae increased in SVL, Marbled Salamander HW increased most rapidly and eventually exceeded both Ringed Salamanders and Spotted Salamanders of a similar size. We also found that both abiotic and biotic factors predicted significant differences in HW corrected for SVL among species, including predator density, competitor density, and hydroperiod. Overall, variability in scaling relationships may provide ecological advantages to each species at different points in ontogeny and different biotic and abiotic factors may induce such variation in asymmetric ways among species.

Pond-Breeding Amphibian Community Composition in Missouri

Abstract We examined pond-breeding amphibian community composition at 210 ponds in Missouri between 2002 and 2012 using drift fence, dipnet, and funnel trap data. We encountered a total of 20 pond-breeding amphibian species in the combined surveys. We also examined whether the presence of American Bullfrogs, Lithobates catesbeianus, and fish influenced these patterns of diversity. Our results indicate the presence of American Bullfrogs, fish, and their interaction influenced the community composition of amphibians at these sites but in opposite patterns. American Bullfrogs often had a positive relationship with the total number of species, total caudate species, and total anuran species, whereas fish presence was negatively associated overall with species diversity, and the presence of both American Bullfrogs and fish was negatively associated with anuran species diversity. It is important to have baseline community species composition data from wide geographical ranges so spatiotemporal changes in community structure can be noted and assessed.

Sampling of Terrestrial Salamanders Reveals Previously Unreported Atypical Color Morphs in the Southern Red-backed Salamander Plethodon serratus

Abstract Atypical color morphs have been described in many species of salamander, but few descriptions are quantitative, and none address those found in the Southern Red-backed Salamander, Plethodon serratus. In 2010 and 2011, we conducted 5 weekly rounds of diurnal leaf litter and natural cover object searches per season (Spring and Fall) in the Ozark Mountains of South central Missouri, U.S.A., with a total of 800 individual 3 × 3 m plots searched. We found 1876 Plethodon serratus (Southern Red-backed Salamander), 20 P. albagula (Western Slimy Salamander), and four Eurycea longicauda melanopleura (Dark-sided Salamander). Of the 1876 P. serratus encountered, all but six exhibited the red-backed phase. Atypical color morphs included the lead-backed phase (n  =  2, including a gravid female), the silver-backed (grey stripe) phase (n  =  1), a ghost-backed (white stripe) phase (n  =  1) and the hypomelanistic phase (n  =  2). The majority of salamanders were found in leaf litter, followed by woody cover and rocks. Two adults of P. serratus were encountered in burrows of cicada nymphs. We encountered only 3 of the 4 color morphs described for P. cinereus, the sister species of P. serratus, but added two color morphs to the list known for P. serratus. We expect that additional surveys for P. serratus in other parts of their range will help determine whether color morph frequency varies between populations and habitat as it does for P. cinereus.