Maintenance of high diversity in mechanistic forest dynamics models of competition for light

Although early theoretical work suggests that competition for light erodes successional diversity in forests, verbal models and recent numerical work with complex mechanistic forest simulators suggest that disturbance in such systems can maintain successional diversity. Nonetheless, if and how allocation trade-offs between competitors interact with disturbance to maintain high diversity in successional systems remains poorly understood. Here, using mechanistic and analytically tractable models, we show that a theoretically unlimited number of coexisting species can be maintained by allocational trade-offs such as investing in light-harvesting organs versus height growth, investing in reproduction versus growth or survival versus growth. The models describe the successional dynamics of a forest composed of many patches subjected to random or periodic disturbance, and are consistent with physiologically mechanistic terrestrial ecosystem models, including the terrestrial components of recent Earth System Models. We show that coexistence arises in our models because species specialize in the successional time they best exploit the light environment and convert resources into seeds or contribute to advance regeneration. We also show that our results are relevant to non-forested ecosystems by demonstrating the emergence of similar dynamics in a mechanistic model of competition for light among annual plant species. Finally, we show that coexistence in our models is relatively robust to the introduction of intraspecific variability that weakens the competitive hierarchy caused by asymmetric competition for light.