Temporal Hierarchies of Variation and the Maintenance of Diversity

Abstract A general model shows how the long‐term growth rate of a population can be partitioned into components representing various mechanisms of maintenance of species diversity. One component summarises the effects of fluctuation‐independent mechanisms, which include classical resource partitioning and frequency‐dependent herbivory. Two other components represent fluctuation‐dependent mechanisms, the storage effect and relative nonlinearity of competition. The general model shows how a community will track an equilibrium set by fluctuation‐independent mechanisms and the environmental state when community dynamics are faster than the rate of environmental change. Fluctuation‐dependent mechanisms can be important for diversity maintenance with or without such tracking, but on long timescales their effects are indistinguishable from those of fluctuation‐independent mechanisms. These considerations lead to a hierarchical view of mechanisms of diversity maintenance where the effects of different timescales are partitioned or merged depending on the timescale of observation. These issues are illustrated with model examples involving various combinations of resource partitioning, fluctuations in recruitment rates, variation in the timing of germination, and seasonality. The very long timescales associated with climate change contain many complexities but nevertheless many ideas applicable to shorter timescales may be useful in a modified form.