No abstract

Summary Phenology is a major component of an organism's fitness. While individual phenological events affect fitness, there is growing evidence to suggest that the relationship between events could be equally or more important. This could explain why temperate deciduous woody plants exhibit considerable variation in the order of reproductive and vegetative events, or flower–leaf sequences (FLSs). There is evidence to suggest that FLSs may be adaptive, with several competing hypotheses to explain their function. Here, we advance existing hypotheses with a new framework that accounts for quantitative FLS variation at multiple taxonomic scales using case studies from temperate forests. Our inquiry provides several major insights towards a better understanding of FLS variation. First, we show that support for FLS hypotheses is sensitive to how FLSs are defined, with quantitative definitions being the most useful for robust hypothesis testing. Second, we demonstrate that concurrent support for multiple hypotheses should be the starting point for future FLS analyses. Finally, we highlight how adopting a quantitative, intraspecific approach generates new avenues for evaluating fitness consequences of FLS variation and provides cascading benefits to improving predictions of how climate change will alter FLSs and thereby reshape plant communities and ecosystems.

Recently, multiple studies have reported declining phenological sensitivities (∆ days per ℃) with higher temperatures. Such observations have been used to suggest climate change is reshaping biological processes, with major implications for forecasts of future change. Here, we show that these results may simply be the outcome of using linear models to estimate nonlinear temperature responses, specifically for events that occur after a cumulative thermal threshold is met-a common model for many biological events. Corrections for the nonlinearity of temperature responses consistently remove the apparent decline. Our results show that rising temperatures combined with linear estimates based on calendar time produce the observations of declining sensitivity-without any shift in the underlying biology. Current methods may thus undermine efforts to identify when and how warming will reshape biological processes.Climate change has reshaped biological processes around the globe, with shifts in the timing of major life history events (phenology), carbon dynamics, and other ecosystem processes. With

1. The relative timing of growth and reproduction is an important driver of plant fitness. For deciduous woody species in temperate regions, leaves and flowers both | 2923