Snow melt timing acts independently and in conjunction with temperature accumulation to drive subalpine plant phenology.

Organisms use environmental cues to align their phenology-the timing of life events-with sets of abiotic and biotic conditions that favor the successful completion of their life cycle. Climate change has altered the environmental cues organisms use to track climate, leading to shifts in phenology with the potential to affect a variety of ecological processes. Understanding the drivers of phenological shifts is critical to predicting future responses, but disentangling the effects of temperature from precipitation on phenology is often challenging because they tend to covary. We addressed this knowledge gap in a high-elevation environment where phenological shifts are associated with both the timing of spring snow melt and temperature. We factorially crossed early snow melt and passive warming treatments to (1) disentangle the effects of snow melt timing and warming on the phenology of flowering and fruiting and reproductive success in three subalpine plant species (Delphinium nuttallianum, Valeriana edulis, and Potentilla pulcherrima); and (2) assess whether snow melt acts via temperature accumulation or some other aspect of the environment (e.g., soil moisture) to affect phenological events. Both the timing and duration of flowering and fruiting responded to the climate treatments, but the effect of snow melt timing and warming varied among species and phenological stages. The combined effects of the treatments on phenology were always additive, and the snow melt treatment often affected phenology even when the warming treatment did not. Despite marked responses of phenology to climate manipulations, the species showed little change in reproductive success, with only one species producing fewer seeds in response to warming (Delphinium, -56%). We also found that snow melt timing can act both through temperature accumulation and as a distinct cue for phenology, and these effects are not mutually exclusive. Our results show that one environmental cue, here snow melt timing, may act through multiple mechanisms to shift phenology.