RESEARCH ARTICLE A Range-Expanding Shrub Species Alters Plant Phenological Response to Experimental Warming

Shifts in plant species phenology (the timing of life-history events such as flowering) have been observed worldwide in concert with rising global temperatures. While most species display earlier phenology with warming, there is large variation among, and even within, species in phenological sensitivity to rising temperatures. Other indirect effects of climate change, such as shifting species composition and altered species interactions, may also be contributing to shifting plant phenology. Here, we describe how experimental warming and the presence of a range-expanding species, sagebrush (Artemisia rothrockii), interact to influence the flowering phenology (day of first and peak flowering) and production (number of flowers) of an alpine cushion plant, Trifolium andersonii, in California’s White Mountains. Both first flowering and peak flowering were strongly accelerated by warming, but not when sagebrush was present. Warming significantly increased flower production of T. andersonii, but less so in the presence of sagebrush. A shading treatment delayed phenology and low-ered flower production, suggesting that shading may be the mechanism by which sage-brush presence delayed flowering of the understory species. This study demonstrates that species interactions can modify phenological responses to climate change, and suggests that indirect effects of rising temperatures arising from shifting species ranges and altered species interactions may even exceed the direct effects of rising temperatures on phenology