Competition for water and species coexistence in phenologically structured annual plant communities

Both competition for water and phenological variation are important determinants of plant community structure, but ecologists lack a synthetic theory for how they affect coexistence outcomes. We developed an analytically tractable model of water competition for Mediterranean annual communities and demonstrate that variation in phenology alone can maintain high diversity in spatially homogenous assemblages of water-limited plants. We modeled a system where all water arrives early in the season and species vary in their ability to grow under drying conditions. As a consequence, species differ in growing season length, and compete by shortening the growing season of their competitors. This model replicates and offers mechanistic explanations for qualitative patterns observed in prior empirical studies of how phenology influences coexistence among Mediterranean annuals. Additionally, we found that a decreasing, concave-up tradeoff between growth rate and access to water can theoretically maintain infinite diversity under simple but realistic assumptions. High diversity is possible because: 1) later plants escape competition after their earlier-season competitors have gone to seed and 2) early-season species are more than compensated for their shortened growing season by a growth-rate advantage. Together, these mechanisms provide an explanation for how annual plant species might coexist when competing only for water.