Weather anomalies more important than climate means in driving insect phenology.

Studies of long-term trends in phenology often rely on climatic averages or accumulated heat, overlooking climate variability. Here we test the hypothesis that unusual weather conditions are critical in driving adult insect phenology. First, we generate phenological estimates for Lepidoptera (moths and butterflies) across the Eastern USA, and over a 70 year period, using natural history collections data. Next, we assemble a set of predictors, including the number of unusually warm and cold days prior to, and during, the adult flight period. We then use phylogenetically informed linear mixed effects models to evaluate effects of unusual weather events, climate context, species traits, and their interactions on flight onset, offset and duration. We find increasing numbers of both warm and cold days were strong effects, dramatically increasing flight duration. This strong effect on duration is likely driven by differential onset and termination dynamics. For flight onset, impact of unusual climate conditions is dependent on climatic context, but for flight cessation, more unusually cold days always lead to later termination particularly for multivoltine species. These results show that understanding phenological responses under global change must account for unusual weather events, especially given they are predicted to increase in frequency and severity.