Abstract
At ontogenetic transitions, animals often exhibit plastic variation in development, behavior and physiology in response to environmental conditions. Most terrestrial-breeding frogs have aquatic larval periods. Some species can extend their initial terrestrial period, as either a plastic embryonic response to balance trade-offs across environments or an enforced wait for rain that allows larvae to access aquatic habitats. Terrestrial larvae of the foam-nesting frog, Leptodactylus fragilis, can arrest development, make their own nest foam to prevent dehydration, and synthesize urea to avoid ammonia toxicity. These plastic responses enable survival during unpredictably long periods in underground nest chambers, waiting for floods to enable exit and continued development in water. However, such physiological and behavioral responses may have immediate and long-term carry-over effects across subsequent ecological and developmental transitions. We examined effects of prolonged terrestriality and larval foam-making activity on larval physiology, development, and metamorphosis in L. fragilis. We tested for changes in foam-making ability by measuring the nests larvae produced following complete removal of parental foam at different ages. We measured ammonia and urea levels in larval foam nests to assess nitrogen excretion patterns, testing for effects of larval age, soil hydration around parental nests, and repeated nest construction. We also assessed immediate and long-term effects of larval foam-making and prolonged terrestriality on larval morphology at water entry and development to metamorphosis. We found that larvae arrested development during prolonged time on land and even young larvae were able to effectively produce multiple foam nests. We found high ammonia concentrations in larval nests, very high urea excretion by developmentally arrested older larvae, and faster growth of larvae in water than while constructing nests. Nonetheless, sibling larvae had a similar aquatic larval period and size at metamorphosis, regardless of their nest-making activity and timing of water entry. Sibship size increased the size of larval foam nests, but reduced per-capita foam production and increased size at metamorphosis, suggesting maternal effects in cooperative groups. Metamorph size also decreased with aquatic larval period. Our results highlight the extent of larval ability to maintain and construct a suitable developmental environment and excrete N-waste as urea, which are both crucial for survival during enforced extensions of terrestriality. Our results suggest that the energetic reserves in large eggs are sufficient to meet metabolic costs of urea synthesis and foam production during developmental arrest over an extended period on land, with no apparent carry-over effects on fitness-relevant traits at metamorphosis.