Dissecting impacts of phenological shifts for performance across biological scales

Although phenological shifts in response to climate are often assumed to benefit species' performance and viability, phenology's role in allowing population persistence and mediating species-level responses in the face of climate change remain unclear. Here, we develop a framework to understand when and why phenological shifts at three biological scales will influence performance: individuals, populations, and macroecological patterns. Specifically, we highlight three underexplored assumptions: (i) individual variability in phenology does not affect population fitness; (ii) population growth rates are sensitive to vital rates affected by phenology; and (iii) phenology mediates species-level responses to climate change including patterns of extinction, invasion, and range shifts. We outline promising methods for understanding how phenological shifts will influence performance within and across biological scales.