Density regulation in toad populations (Epidalea calamita, Bufotes viridis) by differential winter survival of juveniles

Abstract

The size of amphibian populations varies considerably between years, so that systematic trends in dynamics are difficult to detect. Informed conservation management of presumably declining populations requires the identification of the most sensitive life stage. In temperate-zone anurans there is growing evidence that juveniles hibernating for the first time suffer from substantial winter losses. In two syntopic toads (Epidalea calamita, Bufotes viridis) we monitored survival of such juveniles during four consecutive winters in the natural habitat and in four temperature treatments (3°, 5°C, 10°/15°C or 20°C, natural light-dark cycle) in temperature-controlled chambers during winter. Specifically, we tested the hypotheses that (1) winter mortality of juvenile toads which hibernate for the first time in their life is an important component of population dynamics, and that (2) mortality rates differed between the two species. Parameters quantified were size-dependent winter mortality and body condition of pre- and post-hibernating juveniles. Field data provided evidence for the important role of winter mortality of first-hibernators in population dynamics. Choice of hibernacula differed in E. calamita between small and medium-sized individuals and also between the two species suggesting distinct mortality risks. The inability of small E. calamita to reach frost-proof hibernacula by burrowing, and the exposure of small B. viridis to predators are the most probable causes of size-assortative winter mortality. In conclusion, E. calamita juveniles may benefit from rising average winter temperatures in the future by decreased risk of freezing to death, whereas predator-caused winter mortality of B. viridis juveniles will also depend on the effects of climate warming on predator phenology.