A quantitative synthesis of and predictive framework for studying winter warming effects in reptiles

Increases in temperature related to global warming have important implications for organismal fitness. For ectotherms inhabiting temperate regions, 'winter warming' is likely to be a key source of the thermal variation experienced in future years. Studies focusing on the active season predict largely positive responses to warming in the reptiles; however, overlooking potentially deleterious consequences of warming during the inactive season could lead to biased assessments of climate change vulnerability. Here, we review the overwinter ecology of reptiles, and test specific predictions about the effects of warming winters, by performing a meta-analysis of all studies testing winter warming effects on reptile traits to date. We collated information from observational studies measuring responses to natural variation in temperature in more than one winter season, and experimental studies which manipulated ambient temperature during the winter season. Available evidence supports that most reptiles will advance phenologies with rising winter temperatures, which could positively affect fitness by prolonging the active season although effects of these shifts are poorly understood. Conversely, evidence for shifts in survivorship and body condition in response to warming winters was equivocal, with disruptions to biological rhythms potentially leading to unforeseen fitness ramifications. Our results suggest that the effects of warming winters on reptile species are likely to be important but highlight the need for more data and greater integration of experimental and observational approaches. To improve future understanding, we recap major knowledge gaps in the published literature of winter warming effects in reptiles and outline a framework for future research.