High juvenile mortality in amphibians during overwintering related to fungal pathogen exposure

The result of pathogen exposures may depend upon trade-offs in energetic demands for immune responses against host growth and survival. Environmental conditions may influence these trade-offs by affecting host size, or trade-offs may change across seasons, altering impacts of pathogens. We exposed northern leopard frog Lithobates pipiens tadpoles to different larval environments (low leaf litter, high density of conspecifics, atrazine, caged fish, or controls) that influenced size at metamorphosis. Subsequently, we exposed metamorphs to Batrachochytrium dendrobatidis (Bd), a fungal pathogen, just after metamorphosis and/or prior to overwintering 12 wk later. Bd exposure dramatically reduced survival during overwintering, with the strongest effects when hosts were exposed at both time points. Larval environments resulted in differences in host size. Those exposed to caged fish were 2.5 times larger than the smallest (those exposed to high density of conspecifics), but larval environment did not influence Bd effects on growth and survival. The largest frogs exposed to caged fish had greater survival through overwintering, but in the absence of Bd. We built stage-structured models to evaluate if overwinter mortality from Bd is capable of having effects on host populations. Our models suggest that Bd exposure after metamorphosis or before overwintering can reduce population growth rates. Our study demonstrates that hosts suffer little effects of Bd exposures following metamorphosis and that small body size did not hamper growth and survival. Instead, we provide evidence that winter mortality from Bd exposure is capable of reducing population sizes, providing a plausible mechanism for amphibian declines in temperate regions.