Intra- and interspecific variation in the responses of insect phenology to climate.

Phenological change is the most widely documented biological impact of climate change, but shows marked variation in magnitude among populations and species. Thus, quantifying the environmental factors and organismal differences driving this intra- and interspecific variability in phenology is vital to understand and forecast the ecological consequences of climate change. Here, we test intra- and interspecific differences for a set of butterfly species in the organismal sensitivity of flight phenology and its dependence on environmental factors, using as our model system an elevation gradient in a Mediterranean mountain range where temperature and relative humidity vary substantially over space and time. We use field-collected meteorological data, and butterfly counts for 20 univoltine species over 14 years, to test the relative effects on phenology of temperature and relative humidity, the sensitivity of phenology to spatial and temporal variation in temperature and whether ecological traits account for inter-specific variation in sensitivity. For all species, temperature in the months immediately preceding adult emergence had the strongest relationship with phenology. All species appeared earlier in warmer years, with those flying earlier in the season showing the greatest sensitivity to annual (temporal) variation in temperature. However, only a minority of species showed evidence of plastic, space-for-time responses to temperature. Instead, most species showed strong evidence that phenology was more sensitive to temporal than spatial variation in temperature. Our results support the dominant influence of temperature on phenology, even in Mediterranean environments suffering summer drought. They also suggest that accurate forecasts of species' phenological shifts could require the isolation of spatial from temporal components of temperature variation, because the sensitivity of populations and species may differ across these two dimensions. The factors driving synchronisation of phenology over space merit particular research in the context of climate change, given their potential to expose populations simultaneously to environmental extremes.