Changes to butterfly phenology versus elevation range after four decades of warming in the mountains of central Spain

Shifts over time (phenology) and space (latitude and elevation range) represent common ecological responses to climate change. However, the factors determining how changes in phenology and distribution interact, and the consequences for conservation, remain uncertain. Here, we assess how phenology responded to temperature over four decades of warming across the elevation ranges of 18 univoltine butterfly species in four mountain regions of Spain. Using count data from intermittent surveys in 166 sites between 1985 and 2022, we tested for (1) effects of monthly temperature and elevation on mean annual flight date; (2) changes to flight dates between 1985–2005 and 2017–2022; and (3) whether shifts in flight date were related to shifts in the average elevation occupied. Mean flight dates were later in years with cooler springs, and at higher sites, with a mean delay of nearly twenty days per km elevation increase. As conditions warmed over time, average flight date advanced for two thirds of species, especially those whose average elevation was stable over time. Species with stable flight dates showed greater indication of upward range shifts, although only one species showed a significant shift in average elevation. Implications for insect conservation: We show that spring temperatures influence mountain butterfly phenology, and that shifts in phenology and elevation range could compensate for each other in determining population exposure and responses to climate change. Monitoring these changes over time, including by employing evidence from historical surveys and scientific collections, can help to understand constraints on species adaptive capacity to climate change.