Temperature and soil moisture manipulation yields evidence of drought-induced pollen limitation in bee-pollinated squash.

Climate change alters environmental conditions in ways that directly and indirectly affect plants. Flowering plants, for example, modify reproductive allocation in response to heat and drought stress, and such changes can in turn affect pollinator visitation and, ultimately, plant reproduction. Although the individual effects of warming and drought on plant reproductive allocation are well known, these factors may interact to influence reproduction. Here, we conducted a fully crossed temperature by irrigation manipulation in squash () to test how temperature and soil moisture variation affect pollinator-mediated reproduction. To tease apart the direct and indirect effects of temperature and soil moisture, we compared hand-pollinated plants to bee-pollinated plants and restricted bee foraging (i.e., pollen transfer) to one experimental group per day. Temperature and soil-moisture limitation acted independently of one another: warming decreased flower size and increased pollen production, whereas the effects of soil-moisture limitation were uniformly inhibitory. While treatments did not change squash bee ( spp.) behavior, floral visitation by the honey bee () increased with temperature in male flowers and decreased with soil moisture in female flowers. Pollen deposition by bees was independent of plant soil moisture, yet reducing soil moisture increased pollen limitation. This result stemmed at least in part from the effects of soil-moisture limitation on pollen viability; seed set declined with increasing deposition of fluorescent pigment (a proxy for pollen) from plants experiencing decreased soil moisture. These findings suggest that the transfer of lower-quality pollen from plants experiencing soil-moisture limitation led to drought-induced pollen limitation. Similar effects may occur in a wide variety of flowering plant species as climate warming and drought increasingly impact animal-pollinated systems.