Exceptionally preserved tadpoles from the Miocene of Libros,Spain: ecomorphological reconstruction and the impact of ontogeny upon taphonomy

McNamara, M.E., Orr, P.J., Kearns, S.L., Alcalá, L., Anadón, P. & Peñalver‐Mollá, E. 2010: Exceptionally preserved tadpoles from the Miocene of Libros, Spain: ecomorphological reconstruction and the impact of ontogeny upon taphonomy. Lethaia, Vol. 43, pp. 290–306. The Libros exceptional biota from the Upper Miocene of NE Spain includes abundant frog tadpoles (Rana pueyoi) preserved in finely laminated lacustrine mudstones. The tadpoles exhibit a depressed body, short tail, low tail fins, dorso‐laterally directed eyes and jaw sheaths; these features identify the Libros tadpoles as members of the benthic lentic ecomorphological guild. This, the first ecomorphological reconstruction of a fossil tadpole, supports phylogenetic evidence that this ecology is a conserved ranid feature. The soft‐tissue features of the Libros tadpoles are characterized by several modes of preservation. The space occupied previously by the brain is defined by calcium carbonate, the nerve cord is defined by calcium phosphate, and jaw sheaths and bone marrow are preserved as organic remains. Gut contents (and coprolites adjacent to specimens) comprise ingested fine‐grained sedimentary detritus and epiphyton. The body outline and the eyespots, nares, abdominal cavity, notochord, caudal myotomes and fins are defined by a carbonaceous bacterial biofilm. A similar biofilm in adult specimens of R. pueyoi from Libros defines only the body outline, not any internal anatomical features. In the adult frogs, but not in the tadpoles, calcium phosphate and calcium sulphate precipitated in association with integumentary tissues. These differences in the mode of preservation between the adult frogs and tadpoles reflect ontogenetic factors. □Anuran, ecology, soft‐tissue, tadpoles, taphonomy.     

Multiple predator effects on size-dependent behavior and mortality of two species of anuran larvae

This study examined the effects of multiple predators on size‐specific behavior and mortality of two species of anuran larvae. Particularly, we focused on how trait changes in predators and prey may be transmitted to other species in the food web. In laboratory experiments, we examined the effects of bluegill sunfish, Lepomis macrochirus, and the odonate larva Anax junius on behavior and mortality of tadpoles of the bullfrog, Rana catesbeiana, and the green frog R. clamitans. Experiments were conducted with predators alone and together to assess effects on behavior and mortality of the tadpoles. The experiments were replicated on five size classes of the tadpoles to evaluate how responses varied with body size.
Predation rates by Anax were higher on bullfrogs than on green frogs, and both bullfrogs and green frogs suffered greater mortality from Anax than from bluegill. Bluegill only consumed green frogs. Predation rates by both predators decreased with increasing tadpole size and decreased in the non‐lethal (caged) presence of the other predator. Both anuran larvae decreased activity when exposed to predators. Bullfrogs, however, decreased activity more in the presence of Anax than in the presence of bluegill, whereas green frogs decreased activity similarly in the presence of both predators. The largest size class of green frogs, but not of bullfrogs, exhibited spatial avoidance of bluegill. These responses were directly related to the risk posed by the different predators to each anuran species. Anax activity (speed and move frequency) also was higher when alone than in the non‐lethal presence of bluegill. We observed decreased predation rate of each predator in the non‐lethal presence of the other, apparently caused by two different mechanisms. Bluegill decreased Anax mortality on tadpoles by restricting the Anax activity. In contrast, Anax decreased bluegill mortality on tadpoles by reducing tadpole activity. We discuss how the activity and spatial responses of the tadpoles interact with palatability and body size to create different mortality patterns in the prey species and the implications of these results to direct and indirect interactions in this system.     

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

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Do tadpoles affect leaf decomposition in neotropical streams?

1. Of the relatively few studies that have examined consequences of amphibian declines on stream ecosystems, virtually all have focused on changes in algae (or algal‐based food webs) and little is known about the potential effects of tadpoles on leaf decomposition. We compared leaf litter decomposition dynamics in two neotropical streams: one with an intact community of tadpoles (with frogs) and one where tadpoles were absent (frogless) as a result of a fungal pathogen that had driven amphibians locally extinct. The stream with tadpoles contained a diverse assemblage (23 species) of larval anurans, and we identified five species of glass frog (Centrolenidae) tadpoles that were patchily distributed but commonly associated with leaf detritus and organic sediments in pools. The latter reached total densities of 0–318 tadpoles m^?2. 2. We experimentally excluded tadpoles from single‐species leaf packs incubated over a 40‐day period in streams with and without frogs. We predicted that decomposition rates would be higher in control (allowing access of tadpoles) treatments in the study stream with frogs than in the frogless stream and, in the stream with frogs, in the control than in the tadpole exclusion treatment. 3. In the stream with frogs, Centrolene prosoblepon and Cochranella albomaculata tadpoles were patchily distributed in leaf packs (0.0–33.3 m^?2). In contrast to our predictions, leaf mass loss and temperature‐corrected leaf decomposition rates in control treatments were almost identical in our stream with frogs (41.01% AFDM lost, k_{degree day} = ?0.028 day^?1) and in the frogless stream (41.81% AFDM lost, k_{degree day} = ?0.027 day^?1) and between control and tadpole exclusion treatments within each stream. Similarly, there were no significant differences in leaf pack bacterial biomass, microbial respiration rates or macroinvertebrate abundance between treatments or streams. Invertebrate assemblages on leaf packs were similar between treatments (SIMI = 0.97) and streams (SIMI = 0.95) and were dominated by larval Chironomidae, Simuliidae (Diptera) and larval Anchytarsus spp. (Coleoptera). 4. In contrast to dramatic effects of grazing tadpoles on algal communities observed previously, tadpoles had no major effects on decomposition. While centrolenid tadpoles were common in the stream with frogs, their patchy distribution in both experimental and natural leaf packs suggests that their effects on detrital dynamics and microbes are probably more localised than those of grazing tadpoles on algae.     

Phenotypic plasticity in complex environments: effects of structural complexity on predator‐ and competitor‐induced phenotypes of tadpoles of the wood frog,Rana sylvatica

Theoretical and empirical research has demonstrated that phenotypically plastic responses to one environment are dependent on other environmental attributes. Such research is critical considering the complexity of natural habitats, yet few studies have examined how multiple environments affect patterns of plasticity and the adaptiveness of the resulting phenotypes within complex habitats. The present study examines how wood frog (Rana sylvatica) tadpoles alter their behavioural and morphological phenotypes in response to predation risk from larval diving beetles (Dytiscus spp.), competition from conspecifics, and physical structural complexity. It also tests whether structure affects selection intensities by Dytiscus larvae on tadpole morphological traits. Predation risk and competition induced typical changes to tadpole behaviour and morphology. Structure did not induce changes to any phenotype, nor did it interact with predation risk or competition in affecting phenotypes. Furthermore, structure did not affect the predator selection intensities on any morphological trait. Dytiscus larvae selected for shallow, short tailfins, and large tail muscles, yet tadpoles only developed deep tail muscles when raised in the presence of predator cues. These apparently maladaptive responses may have been a result of correlations between phenotypes. The present study expands plasticity research by examining the adaptiveness of plastic responses in complex environments. Additionally, the present study demonstrates that not all environments induce plastic responses. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 853–863.     

Elliptic Fourier analysis of leaf shape in southern African Strychnos section Densiflorae (Loganiaceae)

Leaves can be a useful source of taxonomic information in plants particularly when flowers and fruits are absent during certain periods of the year. In this study, we applied an elliptic Fourier analysis (EFA)‐based morphometric technique to assess leaf morphological divergence among four species of southern African Strychnos section Densiflorae. Using leaf specimen images from field and herbarium collections, we extracted six shape variables [i.e. principal components (PCs)] from the Fourier coefficients and used these variables to describe leaf outline among the species. Our results indicate that the symmetric component of a leaf is the main source of shape differences accounting for 90.25% of total leaf shape variation and captures the more obvious range of observed shapes. PC1 of the symmetric variables describes a wide range of visually observable leaf shape among the species. MANOVA revealed significant interspecific differences except between S. innocua and S. madagascariensis, which could not be separated by outline analysis. A cross‐validated group classification suggests that S. gerrardii, with a classification rate of 88.4%, is distinct from S. madagascariensis, contrary to some taxonomic treatments. We discuss the value of geometric morphometrics at detecting subtle morphological variations and the evolutionary implications of such variations, which may be undetectable to the human eye. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 170 , 542–553.     

Embryonic and larval defensive responses of agile frog (Rana dalmatina) to alien crayfish

Red swamp crayfish Procambarus clarkii, a widespread invasive alien crayfish, represents a serious threat for several freshwater species, including amphibians, which are declining at a global scale. As a shared coevolutionary history is the main factor determining the emergence of antipredator responses, Anuran tadpoles may not be able to cope effectively with this introduced predator. We performed two experiments to assess agile frog's (Rana dalmatina) defensive responses to both P. clarkii and native dragonfly larvae (Anax imperator). First, we conditioned embryos (collected from two ponds 30 km away from each other) with predators’ chemical cues to explore possible variation in hatching time caused by predation risk. In the second experiment, to evaluate how predators’ diet affects tadpole behavior, we conditioned tadpoles for a 5‐week period with cues from tadpole‐fed and gammarid‐fed predators and recorded behavioral and morphological responses. Embryos did not alter hatching time in the presence of any predator cue, while tadpoles from both populations strongly reduced activity and visibility when raised in the presence of tadpole‐fed dragonfly larvae. Morphological changes were less straightforward and were induced only in one population, for which broader tails and a slight increase in body size of tadpoles exposed to tadpole‐fed predators were observed. The lack of defensive responses in crayfish‐exposed tadpoles suggests that the spreading of this invasive species in agricultural lowlands of northern Italy may represent a further threat to their conservation.     

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.     

Vulnerability of an Australian anuran tadpole assemblage to the toxic eggs of the invasive cane toad (Bufo marinus)

The invasion of cane toads (Bufo marinus) across tropical Australia has fatally poisoned many native predators; the most frequent victims may be tadpoles of native frogs, which die when they consume the toxic eggs of the toads. Field studies have documented high and species‐specific mortality of tadpoles following toad spawning. To clarify the determinants of tadpole vulnerability, we conducted 1593 laboratory trials in which single tadpoles were exposed to 10 toad eggs, either with or without an alternative food source (lettuce). At least some tadpoles within all 15 species tested consumed toad eggs. Interspecific variance in survival rates (from 0 to >70%) was driven by feeding responses not by physiological tolerance to toxins: almost all native tadpoles that consumed eggs died rapidly. Tadpole mortality was decreased by the presence of an alternative food source in four species, increased in two species, and not affected in seven species. In three of four taxa where we tested both small (early‐stage) and large (late‐stage) tadpoles, both mean survival rates and the effects of alternative food on survival shifted with tadpole body size. Trials with one species (Limnodynastes convexiusculus) showed no significant inter‐clutch variation in feeding responses or tolerance to toxins. Overall, our data show that cane toad eggs are highly toxic to native anuran tadpoles, but that whether or not a tadpole is killed by encountering toad eggs depends upon a complex interaction between the native anuran's species, its body size, and whether or not alternative food was present. In nature, larval vulnerability also depends upon the seasonal timing and location of spawning events, and habitat selection and foraging patterns of the tadpoles. Our results highlight the complexity of vulnerability determinants, and identify ecological factors (rather than physiology or feeding behaviour) as the primary determinants of cane toad impact on native tadpoles.     

Interactions between freshwater snails and tadpoles: competition and facilitation

Summary Freshwater snails and anuran tadpoles have been suggested to have their highest population densities in ponds of intermediate size where abiotic disturbance (e.g. desiccation) is low and large predators absent. Both snails and tadpoles feed on periphytic algae and, thus, there should be a large potential for competitive interactions to occur between these two distantly related taxa. In a field experiment we examined the relative strength of competition between two closely related snail species, Lymnaea stagnalis and L. peregra, and between L. stagnalis and tadpoles of the common frog, Rana temporaria. Snail growth and egg production and tadpole size at and time to metamorphosis were determined. Effects on the common food source, periphyton, were monitored with the aid of artificial substrates. Periphyton dry weight was dramatically reduced in the presence of snails and/or tadpoles. There were no competitive effects on growth or egg production of the two snail species when they were coexisting. Mortality of L. peregra was high (95%) after reproduction, but independent of treatment. Growth of L. stagnalis was reduced only at the highest tadpole densities, whereas egg production was reduced both by intraspecific competition and by competition with tadpoles. Differences in egg production were retained after tadpole metamorphosis. Tadpole larval period increased, weight of metamorphosing frogs decreased and growth rate was reduced as a function of increasing tadpole density. However, contrary to expectation, snails had a positive effect on tadpole larval period, weight and growth rate. Further, in experimental containers without snails there was a dense growth of the filamentous green alga Cladophora sp. We suggest that the facilitative effects of snails on tadpoles are due to an indirect mutualistic mechanism, involving competition between food sources of different quality (microalgae and Cladophora sp.) and tadpoles being competitively dominant over snails for the preferred food source (microalgae). In the presence of tadpoles snails will be forced to feed on low-quality Cladophora, increasing nutrient turnover rates, which results in enhanced productivity of microalgae, increasing tadpole food resources. Thus, tadpoles have a negative effect on snails through resource depression, while snails facilitate tadpole growth through an indirect enhancement of food availability.     

Visual cues contribute to predator detection in anuran larvae

The ability of prey to detect predators directly affects their probability of survival. Chemical cues are known to be important for predator detection in aquatic environments, but the role of other potential cues is controversial. We tested for changes in behaviour of Rana temporaria tadpoles in response to chemical, visual, acoustic, and hydraulic cues originating from dragonfly larvae (Aeshna cyanea) and fish (Gasterosteus aculeatus). The greatest reduction in tadpole activity occurred when all cues were available, but activity was also significantly reduced by visual cues only. We did not find evidence for tadpoles lowering their activity in response to acoustic and hydraulic cues. There was no spatial avoidance of predators in our small experimental containers. The results show that anuran larvae indeed use vision for predator detection, while acoustic and hydraulic cues may be less important. Future studies of predator‐induced responses of tadpoles should not only concentrate on chemical cues but also consider visual stimuli. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ??, ??–??.     

用功能生态学方法揭示峨眉髭蟾蝌蚪和棘腹蛙蟾蝌蚪共存的原因

采用功能生态学的研究方法,选取了3个指标(功能丰富度、功能定位和功能生态位的重叠程度)对峨眉山清音阁黑龙江中峨眉髭蟾蝌蚪和棘腹蛙蝌蚪共存的原因进行了解析。结果表明,峨眉髭蟾蝌蚪功能丰富度的实测值远远小于棘腹蛙蝌蚪。采用自助法随机抽样15和51个个体时,峨眉髭蟾蝌蚪的功能丰富度值显著小于棘腹蛙蝌蚪。同时,非参数多元方差分析的结果显示二者的功能定位存在显著差异。此外,二者功能生态位重叠程度的实测值与采用自助法计算所得值均极低。以上结果表明二者的功能生态位存在明显分化,表现为峨眉髭蟾蝌蚪个体较大,开口较小,眼间距较大,尾部发达且多采用背腹弯曲的游泳模式;而棘腹蛙蝌蚪开口较大,眼睛间距较小,尾部较为短小且多采用脊椎弯曲的游泳模式。鉴于功能性状可以反映生物体对生长环境的响应,推测两种蝌蚪主要栖息地的微生境也存在分化。     

Factors influencing tadpole deposition site choice in a frog with male parental care: An experimental field study

Parents have evolved a variety of strategies to minimize risks to their offspring, including complex choices regarding suitable rearing sites, based on abiotic and biotic factors, which differentially affect offspring survival. Because availability and quality of these sites are variable, parents may have to choose between immediately available lower‐quality rearing sites or extended search time. In some frog species with larval transport, parents are known to select bodies of water that are free of predators, cannibalistic tadpoles, or intraspecific competitors for larval deposition and rearing sites. We tested whether abiotic factors and the presence of predators and conspecific tadpoles affect tadpole deposition behavior in a population of smooth guardian frog, Limnonectes palavanensis, on the island of Borneo. Females lay eggs on land and males guard them until they hatch; after hatching, tadpoles climb onto the male's back and are subsequently transported to small pools of water on the forest floor, which are scarce and patchily distributed. We estimated the abundance of natural tadpole rearing sites in our study area and conducted experiments using artificial pools to test whether abiotic characteristics of these pools affect the probability of larval deposition. We also performed choice experiments to test whether males of L. palavanensis avoid pools with conspecific tadpoles or predators. Lastly, we tested whether the tadpoles of this species exhibit cannibalism. The abundance of natural deposition sites was low, and males readily used artificial pools for tadpole deposition. Males were less likely to deposit tadpoles in artificial pools located in steep areas, and males did not avoid depositing tadpoles in pools with conspecifics or with experimentally introduced predators. Males exhibited clutch‐partitioning behavior, dividing tadpoles between adjacent artificial pools. Pool availability, rather than the presence of potential competitors or predators in a pool, affects tadpole deposition decisions in this species.     

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     

Temporal variation in behavioral responses to dietary cues from a gape‐limited predator in tadpole prey: A test of the phylogenetic relatedness hypothesis

The ability of prey to recognize and adequately respond to predators determines their survival. Predator‐borne, post‐digestion dietary cues represent essential information for prey about the identity and the level of risk posed by predators. The phylogenetic relatedness hypothesis posits that prey should respond strongly to dietary cues from closely related heterospecifics but respond weakly to such cues from distantly related prey, following a hierarchical pattern. While such responses have mostly been observed in prey at their first encounter with predators, whether prey maintain such hierarchical levels of investment through time remains unclear. We investigated this question by exposing Rhacophorus arboreus tadpoles to the non‐consumptive effect of gape‐limited newt predators Cynops pyrrhogaster that were fed one of five prey diets across a gradient of phylogenetic relatedness: frog tadpoles (Rhacophorus arboreus, Rhacophorus schlegelii, Pelophylax nigromaculatus, and Hyla japonica) and medaka fish (Oryzias latipes). Predators’ diet, time, and their interaction significantly influenced tadpole activity level. We found support for the phylogenetic relatedness hypothesis: Investments in defense were stronger to cues from tadpole diets than to cues from fish diet. However, such a hierarchical response was recorded only in the first four days following predator exposure, then gradually disappear by day 8 on which the tadpoles exhibited similar activity level across all predator treatments. The findings suggest that, at least under the threat of gape‐limited predators, prey use phylogenetic information to evaluate risk and appropriately invest in defense during early encounters with predators; however, energy requirements may prevent prey from maintaining a high level of defense over long exposure to predation risk.     

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.     

Canopy Closure,Competition, and the Endangered Dusky Gopher Frog

Abstract: A major challenge facing wildlife biologists is understanding why some species go extinct while others persist in the same habitat. To address this question, we investigated whether tree canopy closure over ponds affects growth and survival of rare and common tadpoles within ponds and mediates competitive interactions among species. We conducted 2 experiments to test whether canopy closure and competition may have contributed to the decline of the endangered dusky gopher frog (Rana sevosa), but allowed the persistence of the southern leopard frog (R. sphenocephala). We explored the response of both species to canopy closure in single-species and mixed- (1:1) species treatments of identical total tadpole density. An experiment using aquatic enclosures in temporary ponds showed that canopy closure reduced tadpole growth approximately 20% for both species. Survival of dusky gopher frog tadpoles was higher in mixed-species enclosures than in single-species enclosures. In a complementary experiment using artificial ponds, dusky gopher frogs had lower survival to metamorphosis, reduced size at metamorphosis, and produced a lower total biomass of metamorphosed juveniles in shaded ponds. Southern leopard frogs exhibited reduced body size at metamorphosis only when shaded. These studies suggest that pond canopy closure, not larval competition, may be contributing to the decline of the dusky gopher frog. The different responses to canopy closure suggest a potential mechanism for the loss of dusky gopher frogs and the persistence of southern leopard frogs. Removal of trees from historically open-canopy ponds may help facilitate the recovery of dusky gopher frogs and benefit similar species.     

Life history consequences of feeding versus non-feeding in a facultatively non-feeding toad larva

Summary Bufo periglenes, a toad endemic to montane Costa Rica, produces an unusually small clutch of large, yolk-rich eggs. The toads breed in small ephemeral pools that are unpredictable in duration and may be low in food availability. Two congeners, Bufo coniferus and Bufo marinus, occur nearby, breed in more permanent bodies of water that offer more food, and exhibit the typical toad pattern of large clutches of small eggs. Tadpoles of all three species feed on detritus and suspended organic material. By raising tadpoles of the three species individually with and without food I investigated the relationship between egg size (yolk provision) and tadpole survival. All of the unfed B. coniferus and B. marinus tadpoles grew little and died soon after developing to the hindlimb bud stage. On the other hand, all of the unfed B. periglenes tadpoles metamorphosed successfully, demonstrating that the tadpoles are facultatively non-feeding; developmental time from hatching to metamorphosis was significantly shorter for unfed tadpoles than for fed tadpoles, but fed individuals were significantly larger at transformation. Faster developmental rate and larger body size at transformation are both advantageous for frogs and toads, but cannot be attained simultaneously. Large egg size may afford flexibility in unpredictable environments. In pools where food is available, tadpoles presumably eat, take longer to metamorphose, but are larger at transformation than tadpoles developing in nutrient-poor sites. Small body size at transformation (a consequence of not eating) has potential costs, but the large quantity of yolk provided by a large egg enhances the probability of metamorphosis in food-limited environments.