Habitat characteristics predict occupancy patterns of the endangered amphibian Litoria raniformis in flow‐regulated flood plain wetlands

Identification of habitat features that are strongly associated with the occurrence of threatened species is important in terms of predicting impacts of habitat change and identifying key habitats for conservation. In this paper, we apply habitat‐based statistical models to predict occupancy patterns of the endangered southern bell frog (Litoria raniformis) across inland New South Wales (Australia). Litoria raniformis previously occupied a wide range of natural and man‐made waterbodies across a large geographic range, including flood plain wetlands, oxbow lagoons, irrigation canals and rice bays. Alteration of natural flooding regimes has affected a large proportion of habitats within the historical range of Litoria raniformis, but it is not clear how these changes have influenced habitat occupancy patterns. Fifty‐two waterbodies were surveyed for presence/absence of Litoria raniformis in 2001 and 2004. Stepwise logistic regression models were generated to select a subset of variables that best predicted occupancy. Using three predictor variables, vacant and occupied habitats could be predicted with an accuracy of 90% and 70%, respectively. The predictor variables were: the interaction between wetland hydrology and complexity of aquatic vegetation, complexity of fringing vegetation and water temperature. While this study demonstrated that a range of waterbody types were occupied by Litoria raniformis, these habitats shared common hydrological conditions and vegetation characteristics. Altered flooding regimes and reductions in the complexity of aquatic and fringing vegetation are likely to increase the probability of localized extinctions of Litoria raniformis populations.     

Frog breeding in rain-fed wetlands after a period of severe drought: implications for predicting the impacts of climate change

Globally, rain-fed wetlands provide critical habitat for a wide range of amphibian species, however, information on the use of rain-fed wetlands by Australian frog species is extremely limited. This study examined the distribution of frog breeding in rain-fed wetlands following the first significant rain event after a period of severe drought (2002–2009) in order to predict how frog communities may be affected in the future by changed climate. Tadpole communities along with vegetation and water quality variables were described in 35 rain-fed wetlands across the South West Slopes and Riverina bioregions of inland south-eastern Australia. In addition, weekly tadpole surveys were conducted in a subset of these wetlands to describe temporal patterns of occupancy. Despite the protracted dry period prior to this study 50% of the rain-fed wetlands surveyed contained tadpoles. However, frog communities were species poor with only five species recorded. The majority of wetlands were dominated be a single species, Limnodynastes tasmaniensis which is also common within permanent waterbodies such as farm dams and irrigation infrastructure in both bioregions. Tadpoles of two burrowing species L. interioris and Neobatrachus sudelli were restricted to a small number of wetlands mostly in the South West Slopes. The composition of tadpole communities changed over time, and Crinia parinsignifera was the only species that continued to breed over winter. The dominance of generalist species within rain-fed wetlands indicates that characteristics such as dispersal capability, flexibility in breeding times and the ability to utilise created habitats may be more important than burrowing ability and longevity when predicting vulnerability to climate change.     

Neotropical tadpoles influence stream benthos: evidence for the ecological consequences of decline in amphibian populations

1. Few studies have assessed the role of tadpoles in tropical streams, although they are often abundant and conspicuous components of these systems. Moreover, amphibian populations are declining around the globe, particularly stream‐dwelling species in tropical uplands, and the ecological consequences of these losses are not understood. 2. We chose a stream in the central Panamanian highlands, which has an intact fauna of stream‐breeding anurans, to examine the ecological consequences of amphibian losses. This site differs dramatically from sites in nearby western Panama and Costa Rica where anuran diversity and abundance have declined greatly in the last two decades. 3. We used an underwater electric field to create tadpole exclosures in runs, so that we could evaluate their influence on sediment dynamics and the abundance and community structure of algae and aquatic insects. Tadpoles reduced total sediments and both organic and inorganic fractions on substrata. Tadpoles also reduced algal abundance on substrata by approximately 50% and decreased algal biovolume. Gut content analyses showed that tadpoles consumed algae and sediments and we could see that algae and sediments were also displaced through bioturbation. 4. Atelopus zeteki, Rana warszewitschii, and Hyla spp. were the dominant larval anurans responsible for the effects observed. Visual surveys indicated that the densities of these taxa ranged from 23 (R. warszewitschii and Hyla spp. combined) to 43 m^?2 (A. zeteki) in runs. 5. The abundance of baetid mayflies was lower in tadpole exclosures compared with controls, and this was attributed to tadpoles facilitating mayfly feeding by removing sediments and exposing underlying periphyton. 6. Tadpoles affect the abundance and diversity of basal resources and other primary consumers, and thus influence food web dynamics and energy flow in these tropical streams. Catastrophic decline in stream‐breeding anuran populations will influence the structure and function of neotropical stream ecosystems.     

Density is more important than predation risk for predicting growth and developmental outcomes in tadpoles of spotted tree frog,Litoria spenceri (Dubois 1984)

The diverse benefits of group living include protection against predators through dilution effects and greater group vigilance. However, intraspecific aggregation can decrease developmental rates and survival in prey species. We investigated the impact on tadpole development and behaviour of the interaction between population density and predation risk. Spotted tree frog (Litoria spenceri: Hylidae, Dubois 1984) tadpoles were kept at one of three different densities (two tadpoles per litre, five tadpoles per litre or 10 tadpoles per litre) until metamorphosis in the presence or absence of predatory cues. We aimed to determine the influence of population density, predation and the interaction of both factors in determining growth rates in tadpoles. Tadpoles were measured weekly to assess growth and development and filmed to quantify differences in activity and feeding frequency between groups. Generally, tadpoles housed without predators had longer developmental periods when housed with a predator, but there was no effect on tail length or total length. There was no effect of either predation cues or density on percentage of individuals feeding or moving. Although the effects of the presence of predators alone may appear to be less than the effects of the presence of competitors, the prioritisation of competitiveness over predator avoidance may increase vulnerability of tadpoles to the lethal threat of predators. This is particularly important in species such as L. spenceri, which is at risk from introduced fish predators.     

Female-offspring communication in a Taiwanese tree frog, Chirixalus eiffingeri (Anura: Rhacophoridae)

We assessed the roles of visual and olfactory cues in female-tadpole communication inChirixalus eiffingeri . The mean cumulative time that at least one tadpole was active or begged for food was significantly longer when a female C. eiffingeri was present than when a plastic frog was introduced and when no frog was present. Tadpoles did not respond visually to a female frog physically separated from them by transparent Plexiglas. However, tadpoles were more active in water conditioned by female frogs than in unconditioned water. Tadpole activity was further elevated by water conditioned by a female frog and tadpoles. Tadpoles were more active in water conditioned by male frogs than in unconditioned water, but water conditioned by a male frog and tadpoles did not further elevate tadpole activity. Thus, water conditioned by adults of either sex contains substances that increase tadpole activity, but only females show a synergistic effect with conditioning by tadpoles. Copyright 2002 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour     

Contrasting effects of functionally distinct tadpole species on nutrient cycling and litter breakdown in a tropical rainforest stream

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Competing tadpoles: Australian native frogs affect invasive cane toads (Rhinella marina) in natural waterbodies

The cane toad (Rhinella marina) is one of the most successful invasive species worldwide, and has caused significant negative impacts on Australian fauna. Experimental work in the laboratory and in mesocosms has shown that tadpoles of native frogs can affect survival, size at metamorphosis and duration of larval period of cane toad tadpoles. To test if these effects occur in nature, we conducted a field experiment using two temporary ponds where we set up enclosures with tadpoles of native green tree frogs (Litoria caerulea) and cane toads in treatments with a range of densities and combinations. The presence of green tree frog tadpoles significantly decreased the growth rate of toad tadpoles and increased the duration of their larval period in both ponds; in one pond, frog tadpoles also significantly reduced the body length and mass of metamorph toads. Toad tadpoles did not have any significant negative effects on green tree frog tadpoles, but there was strong intraspecific competition within the latter species: increased frog tadpole density resulted in increased larval period and reduced survival, growth rate and size at metamorphosis for frogs at one or both ponds. Our results are encouraging for the possibility of using native frogs as one component of an integrated approach to the biological control of cane toads.     

Foraging responses of predators to novel toxic prey: effects of predator learning and relative prey abundance

Learning to avoid toxic prey items may aid native predators to survive the invasion of highly toxic species, such as cane toads Bufo marinus in tropical Australia. If the predators’ initial aversion is generalized, native prey that resemble the toxic invader may receive a benefit through accidental mimicry. What ecological factors influence the acquisition of learned avoidance (and hence, the impact of invasion on both predators and native prey)? We conducted laboratory experiments to evaluate how the relative abundance of toad tadpoles compared to palatable native tadpoles (Litoria caerulea and L. rubella) affected the ability of native aquatic predators to discriminate between these two prey types. Both fish (northern trout gudgeon, Mogurnda mogurnda) and frogs (Dahl's aquatic frog, Litoria dahlii) learned to discriminate between toads and frogs within an eight‐day period. Higher abundance of toad tadpoles relative to frog tadpoles enhanced rates of predator learning, and thus reduced predation on toads and increased predation on native tadpoles. In the field, spatial and temporal variation in the relative abundance of cane toads compared to native frogs may influence the rates at which these novel toxic items are deleted from predator diets, and the duration of predator protection afforded to natives that resemble the invader.     

Habitat specialization and adaptive phenotypic divergence of anuran populations

We tested for adaptive population structure in the frog Rana temporaria by rearing tadpoles from 23 populations in a common garden experiment, with and without larval dragonfly predators. The goal was to compare tadpole phenotypes with the habitats of their source ponds. The choice of traits and habitat variables was guided by prior information about phenotypic function. There were large differences among populations in life history, behaviour, morphological shape, and the predator-induced plasticities in most of these. Body size and behaviour were correlated with predation risk in the source pond, in agreement with adaptive population divergence. Tadpoles from large sunny ponds were morphologically distinct from those inhabiting small woodland ponds, although here an adaptive explanation was unclear. There was no evidence that plasticity evolves in populations exposed to more variable environments. Much among-population variation in phenotype and plasticity was not associated with habitat, perhaps reflecting rapid changes in wetland habitats.     

Amphibian communities associated with billabong habitats on the Murrumbidgee floodplain,Australia

Abstract A study of frog species richness and abundance at four permanent billabongs located in and around the city of Wagga Wagga, southern New South Wales, was carried out from the spring of 1991 to the autumn of 1992. During the sampling period a total of 404 individual adult frogs from six species were recorded. No egg masses or tadpoles were detected during a total of 32 survey hours. Four species of the Family Myobatrachidae represented 81% of the total number of adult individuals sampled, the remaining species belonging to the Family Hylidae. There were significant differences in the mean species richness and mean total number of adult individuals among billabongs. Significant positive correlations were recorded between total adult individuals, total frog species richness and the number of riparian plant species recorded at all billabongs. There were no significant correlations between adult frog numbers and meteorological and water quality parameters. The adults of the four most abundant frog species exhibited significant association with particular microhabitats surrounding billabongs; Crinia parinsignifera was associated with the creeping grass Paspalum distichum, Limnodynastes fletcheri with the rush Eleocharis sphacelata, L. tasmaniensis with sedges Cyperus spp. and the grass P. distichum and Litoria peronii with red gum Eucalyptus camaldulensis and introduced tree species. Differences in adult frog abundance between billabongs appears to be related to the disturbance of riparian vegetation by domestic stock, while the absence of eggs and tadpoles from these permanent billabongs may be explained by the presence of abundant exotic fish species in the billabongs.     

Consequences of dam‐altered thermal regimes for a riverine herbivore's digestive efficiency,growth and vulnerability to predation

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Predator-prey relationships among larval dragonflies,salamanders, and frogs

Summary Tadpoles of the barking tree frog, Hyla gratiosa, are abundant in spring and summer in some ponds and Carolina bays on the Savannah River Plant near Aiken, South Carolina. To determine how these tadpoles survive in the presence of predaceous salamander larvae, Ambystoma talpoideum, and larvae of an aeshnid dragonfly, Anax junius, we determined fields densities and sizes of the predators and the prey and conducted predation experiments in the laboratory. Tadpoles rapidly grow to a size not captured by Ambystoma, although Anax larvae can capture slightly larger tadpoles. Differing habitat preferences among the tadpoles and the two predator species probably aid in reducing predation pressure. Preliminary work indicates that the tadpoles may have an immobility response to an attack by a predator. In addition, the smallest, most vulnerable tadpoles have a distinctive color pattern which may function to disrupt the body outline and make them indiscernable to predators.     

The role of biotic and abiotic cues in stimulating aggregation by larval cane toads (Rhinella marina)

Tadpoles of the cane toad (Rhinella marina) form dense aggregations in the field, but the proximate cues eliciting this behavior are not well understood. We sampled water‐bodies in the Northern Territory of Australia, finding that the density of cane toad tadpoles increased with increasing temperature. Furthermore, we conducted laboratory experiments to explore the roles of biotic factors (attraction to conspecifics; chemical cues from an injured conspecific; food) and spatially heterogeneous abiotic factors (light levels, water depth, physical structure) to identify the cues that induce tadpole aggregation. Annulus and binary choice trials demonstrated weak but significant attraction between conspecifics. Tadpoles decreased swimming speeds, but did not increase grouping in response to cues from an injured conspecific. Larvae aggregated in response to abiotic cues (high levels of illumination and proximity to physical structures) and were strongly attracted to feeding conspecifics. Overall, aggregation by cane toad tadpoles is likely driven by weak social attraction coupled with a shared preference for specific abiotic features, creating loose aggregations that are then reinforced by movement toward feeding conspecifics.     

Risk of predation affects aggregation size: a study with tadpoles of Phrynomantis microps (Anura: Microhylidae)

Tadpoles of the microhylid frog Phrynomantis microps form swarms near the water surface of savannah ponds during the day. Observations of tadpole distribution at night showed that these aggregations disappear and that tadpoles instead swim in a random distribution near the water surface. The clarity of pond water had significant effects on the distribution of tadpoles during the day. The size of aggregations, and the densities of tadpoles within them, both increased with increasing water clarity. Since tadpoles of this species are known to aggregate in response to visually hunting aquatic predators, these results suggest that tadpoles respond to high water clarity as an indicator of increased predation risk. This supports the hypothesis that, under conditions of increasing predation risk, animals should form larger and more dense groups up to a critical group size. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.      

Divergent responses of exposed and naive Pacific tree frog tadpoles to invasive predatory crayfish

Invasive predators can devastate native species and ecosystems. However, native species may be able to coexist with invasive predators through a variety of mechanisms, such as changes in morphology or behavior due to a plastic response or selection on fixed anti-predator traits. We examined whether exposed and naive populations of Pacific tree frog tadpoles (Pseudacris regilla) display divergent morphological and behavioral traits in response to the invasive predatory red swamp crayfish (Procambarus clarkii). Tadpoles were collected from three study streams with and three without crayfish, in the Santa Monica Mountains of Southern California. We analyzed tadpole morphology and tested anti-predator behavior and survival in the laboratory. Tadpoles from streams with crayfish had shallower, narrower tails than tadpoles from streams without crayfish. Tadpoles from streams with and without crayfish were less active after exposure to crayfish chemical cues. The divergent morphology of naive and exposed tadpoles is consistent with tadpoles exhibiting a plastic response to crayfish or undergoing selection from crayfish predation. In laboratory predation experiments, we found no difference in survival between tadpoles from streams with and without crayfish but tadpoles that survived predation had deeper tail muscles than those that were killed or injured. Our results suggest that deeper tails are advantageous in the presence of crayfish, yet tadpoles from crayfish streams had shallower tails than those from crayfish-free streams. Shallower tails may have an alternative unmeasured advantage or there may be a physiological constraint to developing deeper tails in the wild. These results highlight the ability of a native frog to respond to an invasive predatory crayfish, potentially allowing for coexistence.     

Ecological immunogenetics of life-history traits in a model amphibian

Major histocompatibility complex (MHC) genes determine immune repertoires and social preferences of vertebrates. Immunological regulation of microbial assemblages associated with individuals influences their sociality, and should also affect their life-history traits. We exposed Xenopus laevis tadpoles to water conditioned by adult conspecifics. Then, we analysed tadpole growth, development and survivorship as a function of MHC class I and class II peptide-binding region amino acid sequence similarities between tadpoles and frogs that conditioned the water to which they were exposed. Tadpoles approached metamorphosis earlier and suffered greater mortality when exposed to immunogenetically dissimilar frogs. The results suggest that developmental regulatory cues, microbial assemblages or both are specific to MHC genotypes. Tadpoles may associate with conspecifics with which they share microbiota to which their genotypes are well adapted.     

Mechanisms of competition between tadpoles of Australian frogs (Litoria spp.) and invasive cane toads (Rhinella marina)

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Cannibalistic Interactions Resulting from Indiscriminate Predatory Behavior in Tadpoles of Poison Frogs (Anura: Dendrobatidae)1

Poison frogs in the genus Dendrobates have very small clutch sizes (2–6 eggs among species for which there are data) and typically transport their tadpoles singly to small phytotelmata, such as bromeliad tanks, leaf axils, fallen fruit capsules, and treeholes. Tadpoles of many species are predaceous, consuming larvae of insects that use the same microhabitat for breeding, such as giant damselflies and mosquitoes. Previous studies and observations on the behavior of poison frog tadpoles led us to question whether tadpoles might be cannibalistic. We studied a population of Dendrobates castaneoticus in lowland rainforest in Pará, Brazil; additional data were collected on Dendrobates auratus in Nicaragua. At the study site in Brazil, we established a grid of 40 Brazil nut capsules, the microhabitat used by D. castaneoticus for tadpole deposition. Of 42 tadpoles deposited during the 55 days of the study, 20 were killed or died; 16 of these were presumably killed by conspecific tadpoles. Growth rate and time to metamorphosis was higher among tadpoles that consumed three or more tadpoles or relatively large larvae of the mosquito Trichoprosopon digitatum, a colonist of newly opened Brazil nut capsules. We propose that selection has favored the development of predatory behavior in poison frog tadpoles primarily as a mechanism to eliminate predators from the small phytotelmata in which they develop and that cannibalism is a secondary outcome of this behavior. Predatory behavior also provides tadpoles with a source of food, which is frequently limited in these microhabitats. Additional studies of the biology of tadpoles of other species of Dendrobates are needed to determine the evolution of predatory and cannibalistic behavior in the clade.     

Trophic roles of tadpoles in tropical Australian streams

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Evaluating monitoring methods to guide adaptive management of a threatened amphibian (Litoria aurea)

Prompt detection of declines in abundance or distribution of populations is critical when managing threatened species that have high population turnover. Population monitoring programs provide the tools necessary to identify and detect decreases in abundance that will threaten the persistence of key populations and should occur in an adaptive management framework which designs monitoring to maximize detection and minimize effort. We monitored a population of Litoria aurea at Sydney Olympic Park over 5 years using mark–recapture, capture encounter, noncapture encounter, auditory, tadpole trapping, and dip‐net surveys. The methods differed in the cost, time, and ability to detect changes in the population. Only capture encounter surveys were able to simultaneously detect a decline in the occupancy, relative abundance, and recruitment of frogs during the surveys. The relative abundance of L. aurea during encounter surveys correlated with the population size obtained from mark–recapture surveys, and the methods were therefore useful for detecting a change in the population. Tadpole trapping and auditory surveys did not predict overall abundance and were therefore not useful in detecting declines. Monitoring regimes should determine optimal survey times to identify periods where populations have the highest detectability. Once this has been achieved, capture encounter surveys provide a cost‐effective method of effectively monitoring trends in occupancy, changes in relative abundance, and detecting recruitment in populations.