Towards a heuristic understanding of the storage effect

The storage effect is a general explanation for coexistence in a variable environment. Unfortunately, the storage effect is poorly understood, in part because the generality of the storage effect precludes an interpretation that is simultaneously simple, intuitive and correct. Here, we explicate the storage effect by dividing one of its key conditions-covariance between environment and competition-into two pieces, namely that there must be a strong causal relationship between environment and competition, and that the effects of the environment do not change too quickly. This finer-grained definition can explain a number of previous results, including (1) that the storage effect promotes annual plant coexistence when the germination rate fluctuates, but not when the seed yield fluctuates, (2) that the storage effect is more likely to be induced by resource competition than the apparent competition, and (3) why the storage effect arises readily in models with either stage structure or environmental autocorrelation. Additionally, our expanded definition suggests two novel mechanisms by which the temporal storage effect can arise-transgenerational plasticity and causal chains of environmental variables-thus suggesting that the storage effect is a more common phenomenon than previously thought.