Divergent responses to spring and winter warming drive community level flowering trends.

Analyses of datasets throughout the temperate midlatitude regions show a widespread tendency for species to advance their springtime phenology, consistent with warming trends over the past 20-50 y. Within these general trends toward earlier spring, however, are species that either have insignificant trends or have delayed their timing. Various explanations have been offered to explain this apparent nonresponsiveness to warming, including the influence of other abiotic cues (e.g., photoperiod) or reductions in fall/winter chilling (vernalization). Few studies, however, have explicitly attributed the historical trends of nonresponding species to any specific factor. Here, we analyzed long-term data on phenology and seasonal temperatures from 490 species on two continents and demonstrate that (i) apparent nonresponders are indeed responding to warming, but their responses to fall/winter and spring warming are opposite in sign and of similar magnitude; (ii) observed trends in first flowering date depend strongly on the magnitude of a given species' response to fall/winter vs. spring warming; and (iii) inclusion of fall/winter temperature cues strongly improves hindcast model predictions of long-term flowering trends compared with models with spring warming only. With a few notable exceptions, climate change research has focused on the overall mean trend toward phenological advance, minimizing discussion of apparently nonresponding species. Our results illuminate an understudied source of complexity in wild species responses and support the need for models incorporating diverse environmental cues to improve predictability of community level responses to anthropogenic climate change.