Coevolution of phenological traits shapes plant-pollinator coexistence

Previous research has revealed how species traits determine species interactions, and how species interactions influence species coexistence. However, this hierarchical view ignores the coevolutionary feedback from species interactions to species traits and its consequences for species coexistence. Here, we developed a theoretical model of quantitative genetics to explore how the coevolution of morphological and phenological traits shapes the structure and stability of mutualistic interaction networks. We found that, in comparison to morphological traits, phenological traits led to distinctive species evolutionary trajectories, resulting in different emergent properties at the community level. This is because phenological traits promoted facilitation over competition. While morphological coevolution gave rise to modular and specialized interaction networks with a negative diversity-stability trade-off, phenological coevolution was required for the emergence of nested interaction networks that exhibited a positive relationship between diversity and structural stability. Empirical observations from 17 pollination networks were consistent with the theoretical results: we found many phenological motifs promoting facilitation over competition, suggesting an important role of phenological coevolution in community assembly. The seasonal organization of empirical interactions enhanced the community stability and dampened the diversity-stability trade-off that was observed when aggregating interactions across time. Our results highlight the importance of phenological coevolution in the emergence of diverse and stable communities.