The effect of urbanization on plant phenology depends on regional temperature

Plant and animal phenology is shifting in response to urbanization, with most hypotheses focusing on the 'urban heat island' (UHI) effect as the driver. However, generalities regarding the direction and magnitude of phenological response to urbanization have not yet emerged because most studies have focused on remote-sensed vegetative phenologies or at local scales with relatively few species. Furthermore, how urbanization interacts with broad-scale climate gradients remains an unknown but important component of anthropogenically driven phenological change. Here, we used a database with >22 million in situ plant phenological observations from the United States and Europe to study the joint influence of varying human population density, which serves as an urbanization measure, and of regional temperature on median flowering and leaf-out dates across a wide plant phylogenetic spectrum. Separately, increasing population density and warmer regional temperature both advanced plant flowering and leaf-out. However, the influence of human population density on plant flowering and leaf-out depends on the regional temperature: high population density advanced plant phenology in cold areas but this effect disappeared or even reversed in warm areas. UHI effects (as measured by daily land surface temperature) alone cannot explain the overall influence of urbanization on plant phenology, suggesting that urbanization also affects plant phenology via other mechanisms. Shorter plants with large specific leaf areas and early flower or leaf-out dates were most affected by urbanization and temperature changes. Our study provides strong empirical evidence that the influence of urbanization on plant phenology varies with regional temperature. Therefore, robust understanding and accurate prediction of phenological changes must take this interaction into account.