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Multiple temperature effects on phenology and body size in wild butterflies predict a complex response to climate change.

Temperature-induced alterations in phenology and body size are the cumulative outcome of sequential effects impacting development and are universal responses to climate change. Most studies have so far focused on phenological responses to warming in multiple taxa across space and time, or the ontogenetic effects of temperature in the laboratory. I here complement this work by investigating shifts in phenology and body size (wing length) attributable to temperature changes operating over the entire lifespan of the univoltine orange-tip butterfly Anthocharis cardamines in a single wild population over 14 generations. Phenology was affected by temperatures during three discrete periods in the year prior to emergence, corresponding to late larval/early pupal life, the onset of the chilling period required to break pupal diapause, and postdiapause pupal development prior to eclosion. Higher temperatures during late larval/early pupal life and postdiapause pupal development advanced the subsequent emergence of the butterflies, whereas higher temperatures at the onset of the chilling period retarded it. The synchronization of the butterflies' emergence schedule increased when pupae were exposed to milder midwinter temperatures. Wing length increased with warmer temperatures at distinct points in the early and midpupal periods; such direct effects of temperature on body size could complement season length effects in explaining the reversal of the temperature-size rule in univoltine insects. The periods during which temperature affected the phenology of the butterfly only partially overlapped those affecting the first flowering date of its host plants lady's smock (Cardamine pratensis) and garlic mustard (Alliaria petiolata). Observed thermal effects on flowering time, emergence timing, and emergence synchronization indicate that phenological convergence as well as phenological mismatching could affect host-plant availability and diet breadth; thermal effects on body size imply that important population-level processes could be impacted through correlated changes in fecundity and dispersal rate. In general, the combined effects of phenological and ontogenetic responses to temperature changes across the whole lifespan will likely be important in modeling the demographic responses of interacting species to climate change.