Interactions of an Insecticide with Larval Density and Predation in Experimental Amphibian Communities

Abstract: This study examines the effects of the short-lived insecticide carbaryl, a neurotoxin, on amphibian communities experiencing natural stresses of competition and predation. Tadpoles of three species (  Woodhouse's toad [ Bufo woodhousii ], gray treefrog [ Hyla versicolor ], and green frog [ Rana clamitans ]), representing a commonly encountered assemblage in Missouri, were reared in outdoor polyethylene pond mesocosms. We determined the effects of initial tadpole density ( low or high), predation (newts [  Notophthalmus viridescens ] absent or present), chemical exposure (0, 3.5, or 7.0 mg /L carbaryl), and their interactions on body mass, larval period, and survival to metamorphosis. Green frogs in high-density ponds did not reach metamorphosis, but metamorphs in low-density ponds and tadpoles in high-density ponds were not significantly affected by treatments or their interaction. Carbaryl reduced survival to metamorphosis in toads and treefrogs and increased mass at metamorphosis in treefrogs. Effects of carbaryl varied with predator environment and initial larval density. Interactions of carbaryl with predator and with density may result in an indirect effect of carbaryl causing increased food resources through the elimination of zooplankton populations that may compete for similar resources. Our results indicate that differences in biotic conditions influenced the potency of carbaryl and that even low concentrations induce changes that may alter community dynamics in ways not predicted from single-factor, laboratory-based studies.     

Community ecology of invasions: direct and indirect effects of multiple invasive species on aquatic communities

With many ecosystems now supporting multiple nonnative species from different trophic levels, it can be challenging to disentangle the net effects of invaders within a community context. Here, we combined wetland surveys with a mesocosm experiment to examine the individual and combined effects of nonnative fish predators and nonnative bullfrogs on aquatic communities. Among 139 wetlands, nonnative fish (bass, sunfish, and mosquitofish) negatively influenced the probability of occupancy of Pacific treefrogs (Pseudacris regilla), but neither invader correlated strongly with occupancy by California newts (Taricha torosa), western toads (Anaxyrus boreas), or California red-legged frogs (Rana draytonii). In mesocosms, mosquitofish dramatically reduced the abundance of zooplankton and palatable amphibian larvae (P. regilla and T. torosa), leading to increases in nutrient concentrations and phytoplankton (through loss of zooplankton), and rapid growth of unpalatable toad larvae (through competitive release). Bullfrog larvae reduced the growth of native anurans but had no effect on survival. Despite strong effects on natives, invaders did not negatively influence one another, and their combined effects were additive. Our results highlight how the net effects of multiple nonnative species depend on the trophic level of each invader, the form and magnitude of invader interactions, and the traits of native community members.     

Suitability of Golf Course Ponds for Amphibian Metamorphosis When Bullfrogs Are Removed

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

Multiple stressors in amphibian communities: effects of chemical contamination, bullfrogs, and fish.

A leading hypothesis of amphibian population declines is that combinations of multiple stressors contribute to declines. We examined the role that chemical contamination, competition, and predation play singly and in combination in aquatic amphibian communities. We exposed larvae of American toads (Bufo americanus), southern leopard frogs (Rana sphenocephala), and spotted salamanders (Ambystoma maculatum) to overwintered bullfrog tadpoles (R. catesbeiana), bluegill sunfish (Lepomis macrochirus), the insecticide carbaryl, and ammonium nitrate fertilizer in 1000-L mesocosms. Most significantly, our study demonstrated that the presence of multiple factors reduced survival of B. americanus and A. maculatum and lengthened larval periods of R. sphenocephala. The presence of bluegill had the largest impact on the community; it eliminated B. americanus and A. maculatum and reduced the abundance of R. sphenocephala. Chemical contaminants had the second strongest effect on the community with the insecticide, reducing A. maculatum abundance by 50% and increasing the mass of anurans (frogs and toads) at metamorphosis; the fertilizer positively influenced time and mass at metamorphosis for both anurans and A. maculatum. Presence of overwintered bullfrogs reduced mass and increased time to metamorphosis of anurans. While both bluegill and overwintered bullfrog tadpoles had negative effects on the amphibian community, they performed better in the presence of one another and in contaminated habitats. Our results indicate that predicting deleterious combinations from single-factor effects may not be straightforward. Our research supports the hypothesis that combinations of factors can negatively impact some amphibian species and could contribute to population declines.     

Stability and structure of a fish community on a coral patch reef in Hawaii

Two complete collections of the fishes residing on an isolated coral patch reef ( 1500 m²) at Oahu, Hawaii, were made 11 years apart. Of the 112 species of fishes in both collections combined, only 40% were in common, but these made up more than 85% of the wet biomass in each collection. The two assemblages of fishes were similar in trophic structure and standing crop. Many coral reef fish communities are dominated by carnivorous forms. In the present study, planktivorous fishes were the most important trophic group in the community; this was related to abundant zooplankton resources. Following the second collection in 1977, recolonization by fishes was followed for 1 year. Recolonization proceeded rapidly and was primarily by juvenile fishes well beyond larval metamorphosis. Within 6 months of the second collection, the trophic structure had been re-established. The MacArthur-Wilson model of insular colonization described the recolonization process and predicted an equilibrium situation in less than 2 years. The recolonization data suggested that chance factors may explain the colonization process on a small scale, but a relatively deterministic pattern emerged when considering the entire reef. Thus, at the community level the fishes are a persistent and predictable entity.     

Effects of the Interaction between Genetic Diversity and UV-B Radiation on Wood Frog Fitness

Abstract:  Genetic diversity may buffer amphibian populations against environmental vicissitudes. We hypothesized that wood frogs (  Rana sylvatica ) from populations with lower genetic diversity are more susceptible to ultraviolet-B (UV-B) radiation than those from populations with higher diversity. We used RAPD markers to obtain genetic diversity estimates for 12 wood frog populations. We reared larval wood frogs from these populations and exposed experimental groups of eggs and larvae to one of three treatments: unfiltered sunlight, sunlight filtered through a UV-B-blocking filter (Mylar), and sunlight filtered through a UV-B-transmitting filter (acetate). In groups exposed to UV-B, larval mortality and deformity rates increased significantly, but egg mortality did not. We found a significant negative relationship between genetic diversity and egg mortality, larval mortality, and deformity rates. Furthermore, the interaction between UV-B treatment and genetic diversity significantly affected larval mortality. Populations with low genetic diversity experienced higher larval mortality rates when exposed to UV-B than did populations with high genetic diversity. This is the first time an interaction between genetic diversity and an environmental stressor has been documented in amphibians. Differences in genetic diversity among populations, coupled with environmental stressors, may help explain patterns of amphibian decline.     

The role of environmental and spatial processes in structuring lake communities from bacteria to fish

We assessed the relative roles of local environmental conditions and dispersal on community structure in a landscape of lakes for the major trophic groups. We use taxonomic presence-absence and abundance data for bacteria, phytoplankton, zooplankton, and fish from 18 lakes in southern Quebec, Canada. The question of interest was whether communities composed of organisms with more limited dispersal abilities, because of size and life history (zooplankton and fish) would show a different effect of lake distribution than communities composed of good dispersers (bacteria and phytoplankton). We examine the variation in structure attributable to local environmental (i.e., lake chemical and physical variables) vs. dispersal predictors (i.e., overland and watercourse distances between lakes) using variation partitioning techniques. Overall, we show that less motile species (crustacean zooplankton and fish) are better predicted by spatial factors than by local environmental ones. Furthermore, we show that for zooplankton abundances, both overland and watercourse dispersal pathways are equally strong, though they may select for different components of the community, while for fish, only watercourses are relevant dispersal pathways. These results suggest that crustacean zooplankton and fish are more constrained by dispersal and therefore more likely to operate as a metacommunity than are bacteria and phytoplankton within this studied landscape.     

Amphibian Breeding Distribution in an Urbanized Landscape

Abstract:  Amphibians commonly use wetlands for breeding habitat, and given the concern about their ongoing global declines, the effects of urbanization on the breeding distribution of amphibians need to be quantified. Thus, we conducted a survey of the larval amphibian community in central Pennsylvania (U.S.A.) wetlands along an urbanization gradient. Wetlands in urban areas had less surrounding forest and wetlands and greater road density than rural wetlands. Urbanization was also associated with increases in hydroperiod (i.e., wetland permanency) and the presence of fish predators. Moreover, urban wetlands had lower larval amphibian species richness than rural wetlands. This decrease in richness was attributable to a decrease in occurrence of wood frogs ( Rana sylvatica ) and ambystomatid salamanders ( Ambystoma maculatum and A. jeffersonianum ) in urban sites. Wood frogs and ambystomatid salamanders were positively associated with the amount of forest surrounding sites and negatively associated with hydroperiod. As a result, we hypothesize that these species are sensitive to the effects of urban development. The remaining species in this study appear fairly resilient to the effects of urbanization. These data demonstrate the importance of quantifying both local and landscape attributes when describing the factors that limit the breeding distribution of amphibians. We recommend that to preserve amphibian biodiversity in urbanized landscapes, it is best to focus on regional diversity, which protects a variety of sites that encompass various hydroperiods, have adequate buffer habitat, and are connected by dispersal routes.     

Zooplankton biodiversity and lake trophic state: explanations invoking resource abundance and distribution

While empirical studies linking biodiversity to local environmental gradients have emphasized the importance of lake trophic status (related to primary productivity), theoretical studies have implicated resource spatial heterogeneity and resource relative ratios as mechanisms behind these biodiversity patterns. To test the feasibility of these mechanisms in natural aquatic systems, the biodiversity of crustacean zooplankton communities along gradients of total phosphorus (TP) as well as the vertical heterogeneity and relative abundance of their phytoplankton resources were assessed in 18 lakes in Quebec, Canada. Zooplankton community richness was regressed against TP, the spatial distribution of phytoplankton spectral groups, and the relative biomass of spectral groups. Since species richness does not adequately capture ecological function and life history of different taxa, features which are important for mechanistic theories, relationships between zooplankton functional diversity (FD) and resource conditions were examined. Zooplankton species richness showed the previously established tendency to a unimodal relationship with TP, but functional diversity declined linearly over the same gradient. Changes in zooplankton functional diversity could be attributed to changes in both the spatial distribution and type of phytoplankton resource. In the studied lakes, spatial heterogeneity of phytoplankton groups declined with TP, even while biomass of all groups increased. Zooplankton functional diversity was positively related to increased heterogeneity in cyanobacteria spatial distribution. However, a smaller amount of variation in functional diversity was also positively related to the ratio of biomass in diatoms/chrysophytes to cyanobacteria. In all observed relationships, a greater variation of functional diversity than species richness measures was explained by measured factors, suggesting that functional measures of zooplankton communities will benefit ecological research attempting to identify mechanisms behind environmental gradients affecting diversity.     

Effects of the herbicide Roundup on freshwater microbial communities: a mesocosm study

The impact of the widely used herbicide glyphosate has been mainly studied in terrestrial weed control, laboratory bioassays, and field studies focusing on invertebrates, amphibians, and fishes. Despite the importance of phytoplankton and periphyton communities at the base of the aquatic food webs, fewer studies have investigated the effects of glyphosate on freshwater microbial assemblages. We assessed the effect of the commercial formulation Roundup using artificial earthen mesocosms. The herbicide was added at three doses: a control (without Roundup) and two treatments of 6 and 12 mg/L of the active ingredient (glyphosate). Estimates of the dissipation rate (k) were similar in the two treatments (half-lives of 5.77 and 7.37 d, respectively). The only two physicochemical parameters showing statistically significant differences between treatments and controls were the downward vertical spectral attenuation coefficient kd(lambda), where lambda is wavelength, and total phosphorus concentration (TP). At the end of the experiment, the treated mesocosms showed a significant increase in the ratio kd(490 nm)/k(d)(550 nm) and an eightfold increase in TP. Roundup affected the structure of phytoplankton and periphyton assemblages. Total micro- and nano-phytoplankton decreased in abundance in treated mesocosms. In contrast, the abundance of picocyanobacteria increased by a factor of about 40. Primary production also increased in treated mesocosms (roughly by a factor of two). Similar patterns were observed in the periphytic assemblages, which showed an increased proportion of dead: live individuals and increased abundances of cyanobacteria (about 4.5-fold). Interestingly, the observed changes in the microbial assemblages were captured by the analysis of the pigment composition of the phytoplankton, the phytoplankton absorption spectra, and the analysis of the optical properties of the water. The observed changes in the structure of the microbial assemblages are more consistent with a direct toxicological effect of glyphosate rather than an indirect effect mediated by phosphorus enrichment.     

Effects of chytrid fungus and a glyphosate-based herbicide on survival and growth of wood frogs (Lithobates sylvaticus)

Anthropogenic-derived stressors in the environment, such as contaminants, are increasingly considered important cofactors that may decrease the immune response of amphibians to pathogens. Few studies, however, have integrated amphibian disease and contaminants to test this multiple-stressor hypothesis for amphibian declines. We examined whether exposure to sublethal concentrations of a glyphosate-based herbicide and two strains of the pathogenic chytrid fungus, Batrachochrytrium dendrobatidis (Bd) could: (1) sublethally affect wood frogs (Lithobates sylvaticus) by altering the time to and size at metamorphosis, and (2) directly affect survivability of wood frogs after metamorphosis. Neither Bd strain nor herbicide exposure alone significantly altered growth or time to metamorphosis. The two Bd strains did not differ in their pathogenicity, and both caused mortality in post-metamorphic wood frogs. There was no evidence of an interaction between treatments, indicating a lack of herbicide-induced susceptibility to Bd. However, the trends in our data suggest that exposure of wood frogs to a high concentration of glyphosate-based herbicide may reduce Bd-caused mortality compared to animals exposed to Bd alone. These results exemplify the complexities inherent when populations are coping with multiple stressors. In this case, the perceived stressor, glyphosate-based herbicide, appeared to affect the pathogen more than the host's immune system, relieving the host from disease-caused effects. This suggests caution when invoking multiple stressors as a cause for increased disease susceptibility and indicates that the effects of multiple stressors on disease outcome depend on the interrelationships of stressors to both the pathogen and the host.     

Intraspecific variation in a predator affects community structure and cascading trophic interactions

Intraspecific phenotypic variation in ecologically important traits is widespread and important for evolutionary processes, but its effects on community and ecosystem processes are poorly understood. We use life history differences among populations of alewives, Alosa pseudoharengus, to test the effects of intraspecific phenotypic variation in a predator on pelagic zooplankton community structure and the strength of cascading trophic interactions. We focus on the effects of differences in (1) the duration of residence in fresh water (either seasonal or year-round) and (2) differences in foraging morphology, both of which may strongly influence interactions between alewives and their prey. We measured zooplankton community structure, algal biomass, and spring total phosphorus in lakes that contained landlocked, anadromous, or no alewives. Both the duration of residence and the intraspecific variation in foraging morphology strongly influenced zooplankton community structure. Lakes with landlocked alewives had small-bodied zooplankton year-round, and lakes with no alewives had large-bodied zooplankton year-round. In contrast, zooplankton communities in lakes with anadromous alewives cycled between large-bodied zooplankton in the winter and spring and small-bodied zooplankton in the summer. In summer, differences in feeding morphology of alewives caused zooplankton biomass to be lower and body size to be smaller in lakes with anadromous alewives than in lakes with landlocked alewives. Furthermore, intraspecific variation altered the strength of the trophic cascade caused by alewives. Our results demonstrate that intraspecific phenotypic variation of predators can regulate community structure and ecosystem processes by modifying the form and strength of complex trophic interactions.     

Independent gradients of producer, consumer, and microbial diversity in lake plankton

Interactions between trophic levels during food web assembly can drive positive correlations in diversity between producers, consumers, and decomposers. However, the contribution of trophic interactions relative to local environmental factors in promoting species diversity is poorly understood, with many studies only considering two trophic levels. Here we examine correlations in diversity among zooplankton, phytoplankton, and bacteria in the pelagic zone of 31 lakes in British Columbia, Canada. We sampled species diversity of zooplankton and phytoplankton through morphological identification, and bacterial genetic diversity was estimated by denaturing gradient gel electrophoresis (DGGE) of 16S rDNA polymorphisms. We looked for correlations in diversity that were independent of the abiotic environment by statistically controlling for 18 limnological variables. No significant correlations were found between the diversity of zooplankton, phytoplankton, and bacteria. In addition, the physical factors that were associated with species composition in one trophic level were independent of those that were important for another. Our results provide no support for the importance of direct feedbacks between producers, consumers, and decomposers in maintaining diversity. Zooplankton, phytoplankton, and bacterial diversity and composition are regulated independently from one another and respond to different environmental variables. These results suggest that species of lake plankton show loose trophic associations with one another due to broad diets in consumers and decomposers.     

Shift in nutrient and plankton community in eutrophic lake following introduction of a freshwater bivalve

The impact of the freshwater bivalve Corbicula leana on plankton community dynamics was examined during a cyanobacterial bloom period. Nutrient and chlorophyll concentrations, primary productivity, and phytoplankton and zooplankton communities in the experimental enclosures were measured at 2-3 day intervals. The introduction of mussels reduced net primary productivity and phytoplankton and chlorophyll. Chlorophyll decreased immediately following addition of 100 mussels and then increased over time. After 600 mussels were added, chlorophyll decreased continuously from 87to 25 microg l(-1), approaching that in the mussel-free enclosure. Simultaneously, water transparency increased and concentrations of suspended solids and total phosphorus decreased. Mussel addition caused short-term increases in nutrient concentrations, especially following high-density treatment: phytoplankton density decreased, while cell density in the mussel-free enclosure increased. Zooplankton densities in the two enclosures were similar; however, carbon biomass in the mussel enclosure increased, associated with an increase in large zooplankton. The trophic relationship between phytoplankton and zooplankton was positive in the mussel-free enclosure and negative in the mussel-treatment enclosure, possibly reflecting effects of mussels on both consumer and resource control in the plankton community. Thus, filter feeding by Corbicula affects nutrient recycling and plankton community structure in a freshwater ecosystem through direct feeding and competition for food resources.     

Interspecific Variation in Susceptibility of Frog Tadpoles to the Pathogenic Fungus Batrachochytrium dendrobatidis

Abstract:  As part of an overall biodiversity crisis many amphibian populations are in decline throughout the world. Numerous causes have been invoked to explain these declines. These include habitat destruction, climate change, increasing levels of ultraviolet radiation, environmental contamination, and the introduction of non-native species and diseases. Several types of pathogens have been implicated in contributing to amphibian population declines: viruses, bacteria, oomycetes, and fungi. One particular fungus, the chytridiomycete Batrachochytrium dendrobatidis may have caused amphibian population declines in several regions. This pathogen causes chytridiomycosis, which is fatal to newly metamorphic and adult amphibians of certain species. We present experimental evidence that larval stages may also be susceptible to exposure to Batrachochytrium . There was, however, differential sensitivity to B. dendrobatidis in larvae we examined. In laboratory experiments, larval western toads (  Bufo boreas ) exposed to B. dendrobatidis experienced increased mortality and behaviors that suggested they were affected by exposure compared with unexposed control tadpoles. Larvae of Cascades frogs (  Rana cascadae ), bullfrogs ( R. catesbeiana ), and Pacific treefrogs ( Hyla regilla ) did not die after exposure to Batrachochytrium and appeared to behave normally. R. cascadae larvae exposed to B. dendrobatidis , however, showed an increase incidence in mouthpart abnormalities, a characteristic effect of chytridiomycosis, compared with unexposed controls. These results show that Batrachochytrium can negatively affect some species of amphibians at the larval stage and not others. The implications of interspecific variation in susceptibility to fungal infection are broad.     

Prevalence of malformed frogs in Kaoping and Tungkang river basins of southern Taiwan

In this study we found many amphibians with bizarre appearances, known as malformations in Pingtung County southern Taiwan. For this investigation we collected frogs inhabiting the Kaoping and Tungkang river watersheds between February 2006 and June 2007. Among the total number of 10,909 normal frogs (i.e., anurans) collected during the investigation period, the Indian rice frogs (Rana limnocharis) account for the greatest number next is the Chinese bullfrog (Rana rugulosa). Of all the 244 captured malformed frogs, the Indian rice frog account for the greatest proportion. These malformed frogs have their main distribution in upstream areas of these two rivers. Our result indicates that the appearance rate of malformed frogs is 1.8% in the upstream reaches of the Kaoping River and 2.6%, and 0.8%, respectively in the upstream and midstream reaches of the Tungkang river. The most-commonly-found malformation is the lack of palms, followed by the lack of appendages, exostosis, and a malformed appendicular. It is, therefore, reasonable to speculate that the causes for the malformation may be related to the increased organic pollutants and agricultural chemicals used in the upstream reaches of these two rivers.     

Food limitation stimulates metamorphosis of competent larvae and alters postmetamorphic growth rate in the marine prosobranch gastropodCrepidula fornicata

The effects of food limitation on growth rates and survival of marine invertebrate larvae have been studied for many years. Far less is known about how food limitation during the larval stage influences length of larval life or postmetamorphic performance. This paper documents the effects of food limitation during larval development (1) on how long the larvae ofCrepidula fornicata (L.) can delay metamorphosis in the laboratory after they have become competent to metamorphose and (2) on postmetamorphic growth rate. To assess the magnitude of nutritional stress imposed by different food concentrations, we measured growth rates (as changes in shell length and ash-free dry weight) for larvae reared in either 0.45-m filtered seawater or at phytoplankton concentrations (Isoehrysis galbana, clone T-ISO) of 1 × l0³, 1 × 10⁴, or 1.8 × 10⁵ cells ml^–1. Larvae increased both shell length and biomass at 1 × 10⁴ cells ml^–1, although significantly more slowly than at the highest food concentration. Larvae did not significantly increase (p > 0.10) mean shell length in filtered seawater or at a phytoplankton concentration of only 1 × 10³ cells ml^–1, and in fact lost weight under these conditions. To assess the influence of food limitation on the ability of competent individuals to postpone metamorphosis, larvae were first reared to metamorphic competence on a high food concentration ofI. galbana (1.8 × 10⁵ cells ml^–1). When at least 80% of subsampled larvae were competent to metamorphose, as assessed by the numbers of indlviduals metamorphosing in response to elevated K⁺ concentration in seawater, remaining larvae were transferred either to 0.45-m filtered seawater or to suspensions of reduced phytoplankton concentration (1 × 10³, 1 × 10⁴, or 5 × 10⁴ cells ml^–1), or were maintained at 1.8 × 10⁵ cells ml^–1. All larvae were monitored daily for metamorphosis. Individuals that metamorphosed in each food treatment were transferred to high ration conditions (1.8 × 10⁵ tells ml^–1) for four additional days to monitor postmetamorphic growth. Competent larvae responded to all food-limiting conditions by metamorphosing precociously, typically 1 wk or more before larvae metamorphosed when maintained at the highest food ration. Surprisingly, juveniles reared at full ration grew more slowly if they had spent 2 or 3 d under food-limiting conditions as competent larvae. The data show that a rapid decline in phytoplankton concentration during the larval development ofC. fornicata stimulates metamorphosis, foreshortening the larval dispersal period, and may also reduce the ability of postmetamorphic individuals to grow rapidly even when food concentrations increase.     

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

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

Carryover aquatic effects on survival of metamorphic frogs during pond emigration.

In organisms with complex life cycles, physiological stressors during early life stages may have fitness-level impacts that are delayed into later stages or habitats. We tested the hypothesis that body size and date of metamorphosis, which are highly responsive to aquatic stressors, influence post-metamorphic survival and movement patterns in the terrestrial phase of an ephemeral pond-breeding frog by examining these traits in two populations of northern red-legged frogs (Rana aurora aurora). To increase variation of body size at metamorphosis, we manipulated food availability for 314 of 1045 uniquely marked tadpoles and estimated the probability that frogs survived and emigrated using concentric rings of drift fencing surrounding ponds and Bayesian capture-recapture modeling. The odds of surviving and emigrating from the ponds to the innermost drift fences, approximately 12 m, increased by factors of 2.20 (95% credibility intervals 1.39-4.23) and 2.54 (0.94-4.91) with each millimeter increase in snout-vent length and decreased by factors of 0.91 (0.85-0.96) and 0.89 (0.80-1.00) with each day's delay in metamorphosis for the two ponds. The odds of surviving and moving to the next ring of fencing, 12 m to approximately 40 m from the ponds, increased by a factor of 1.20 (0.45-4.06) with each millimeter increase in size. Our results demonstrated that body size and timing of metamorphosis relate strongly to the performance of newly metamorphosed frogs during their initial transition into terrestrial habitat. Carryover effects of aquatic stressors that reduce size and delay metamorphosis may have population-level impacts that are not expressed until terrestrial stages. Since changes in both aquatic and terrestrial systems are implicated in many amphibian declines, quantifying both immediate and delayed effects of stressors on demographic rates is critical to sound management.     

Stable isotopes identify an ontogenetic niche expansion in Antarctic krill (Euphausia superba) from the South Shetland Islands, Antarctica

Antarctic krill (Euphausia superba) occupy a key position in the Southern Ocean linking primary production to secondary consumers. While krill is a dominant grazer of phytoplankton, it also consumes heterotrophic prey and the relative importance of these two resources may differ with ontogeny. We used stable isotope analyses to evaluate body size-dependent trophic and habitat shifts in krill during the austral summer around the South Shetland Islands, Antarctica. We found evidence for an asymmetric, ontogenetic niche expansion with adults of both sexes having higher and more variable δ¹⁵N values but consistent δ¹³C values in comparison with juveniles. This result suggests that while phytoplankton likely remains an important life-long resource, krill in our study area expand their dietary niche to include higher trophic food sources as body size increases. The broader dietary niches observed in adults may help buffer them from recent climate-driven shifts in phytoplankton communities that negatively affect larval or juvenile krill that rely predominately on autotrophic resources.