Climate conditions and resource availability drive return elevational migrations in a single-brooded insect.

Seasonal elevational migrations have important implications for life-history evolution and ecological responses to environmental change. However, for most species, particularly invertebrates, evidence is still scarce for the existence of such migrations, as well as for the potential causes. We tested the extent to which seasonal abundance patterns in central Spain for overwintering (breeding) and summer (non-breeding) individuals of the butterfly Gonepteryx rhamni were consistent with three hypotheses explaining elevational migration: resource limitation (host plant and flower availability), physiological constraints of weather (maximum temperatures) and habitat limitation (forest cover for overwintering). For overwintering adults, abundance was positively associated with host plant density during two intensive survey seasons (2007-2008), and the elevational distribution was relatively stable over a 7-year period (2006-2012). The elevational distribution of summer adults was highly variable, apparently related both to temperature and habitat type. Sites occupied by adults in the summer were on average 3 °C cooler than their breeding sites, and abundance showed negative associations with summer temperature, and positive associations with forest cover and host plant density in 2007 and 2008. The results suggest that the extent of uphill migration in summer could be driven by different factors, depending on the year, and are mostly consistent with the physiological constraint and habitat limitation hypotheses. In contrast, the patterns for overwintering adults suggest that downhill migration can be explained by resource availability. Climate change could generate bottlenecks in the populations of elevational migrant species by constraining the area of specific seasonal habitat networks or by reducing the proximity of environments used at different times of year.