Pollen resistance to water in 80 angiosperm species: flower structures protect rain-susceptible pollen

Flowers exhibit adaptive responses to biotic and abiotic factors. It remains unclear whether pollen susceptibility to rain damage plays a role in the evolution of floral form. We investigated flower performance in rain and compared pollen longevity in dry conditions, pure water and solutions with different sucrose concentrations in 80 flowering species from 46 families with diverse floral shapes and pollination modes. A pollen viability test showed that pollen longevity in all studied species was greatly reduced by wetting. We found that pollen of species with complete protection by flower structures was susceptible to water damage and a high proportion of resistant pollen occurred in unprotected species. Flowers whose structures expose pollen to rain may also reduce rain damage through temporal patterns of pollen presentation. This prediction was supported by our direct measurement of pollen presentation duration on rainy days. Our observations showed that variation in pollen performance in water was associated with differences in floral forms. Water-resistant pollen and extended pollen presentation duration were favored by selection via rain contact in species in which pollen was not protected from rain. These findings support the functional hypothesis that flower structures protect susceptible pollen from rain, demonstrating that rain acts as a force shaping floral form.