Linking inter-annual variation in environment, phenology, and abundance for a montane butterfly community.

Climate change has caused widespread shifts in species' phenology, but the consequences for population and community dynamics remain unclear because of uncertainty regarding the species-specific drivers of phenology and abundance, and the implications for synchrony among interacting species. Here, we develop a statistical model to quantify inter-annual variation in phenology and abundance over an environmental gradient, and use it to identify potential drivers of phenology and abundance in co-occurring species. We fit the model to counts of 10 butterfly species with single annual generations over a mountain elevation gradient, as an exemplar system in which temporally limited availability of biotic resources and favorable abiotic conditions impose narrow windows of seasonal activity. We estimate parameters describing changes in abundance, and the peak time and duration of the flight period, over ten years (2004-2013) and across twenty sample locations (930-2,050 m) in central Spain. We also use the model outputs to investigate relationships of phenology and abundance with temperature and rainfall. Annual shifts in phenology were remarkably consistent among species, typically showing earlier flight periods during years with warm conditions in March or May-June. In contrast, inter-annual variation in relative abundance was more variable among species, and generally less well associated with climatic conditions. Nevertheless, warmer temperatures in June were associated with increased relative population growth in three species, and five species had increased relative population growth in years with earlier flight periods. These results suggest that broadly coherent interspecific changes to phenology could help to maintain temporal synchrony in community dynamics under climate change, but that the relative composition of communities may vary due to interspecific inconsistency in population dynamic responses to climate change. However, it may still be possible to predict abundance change for species based on a robust understanding of relationships between their population dynamics and phenology, and the environmental drivers of both.