Experimental Test of the Combined Effects of Water Availability and Flowering Time on Pollinator Visitation and Seed Set

Climate change is likely to alter both flowering phenology and water availability for plants. Either of these changes alone can affect pollinator visitation and plant reproductive success. The relative impacts of phenology and water, and whether they interact in their impacts on plant reproductive success remain, however, largely unexplored. We manipulated flowering phenology and soil moisture in a factorial experiment with the subalpine perennial Mertensia ciliata (Boraginaceae). We examined responses of floral traits, floral abundance, pollinator visitation, and composition of visits by bumblebees vs. other pollinators. To determine the net effects on plant reproductive success, we also measured seed production and seed mass. Reduced water led to shorter, narrower flowers that produced less nectar. Late flowering plants produced fewer and shorter flowers. Both flowering phenology and water availability influenced pollination and reproductive success. Differences in flowering phenology had greater effects on pollinator visitation than did changes in water availability, but the reverse was true for seed production and mass, which were enhanced by greater water availability. The probability of receiving a flower visit declined over the season, coinciding with a decline in floral abundance in the arrays. Among plants receiving visits, both the visitation rate and percent of non-bumblebee visitors declined after the first week and remained low until the final week. We detected interactions of phenology and water on pollinator visitor composition, in which plants subject to drought were the only group to experience a late-season resurgence in visits by solitary bees and flies. Despite that interaction, net reproductive success measured as seed production responded additively to the two manipulations of water and phenology. Commonly observed declines in flower size and reward due to drought or shifts in phenology may not necessarily result in reduced plant reproductive success, which in M. ciliata responded more directly to water availability. The results highlight the need to go beyond studying single responses to climate changes, such as either phenology of a single species or how it experiences an abiotic factor, in order to understand how climate change may affect plant reproductive success.