The storage effect is not about bet-hedging or population stage-structure

The storage effect is a well-known explanation for coexistence in temporally varying environments. Like many complex ecological theories, the storage effect is often used as an explanation for observed coexistence on the basis of heuristic understanding, rather than careful application of a detailed model. One interpretation states that species coexist by specializing on specific environmental states, and therefore must have a robust life-stage (e.g., long-lived adults, a seed-bank) in order to "wait it out" for favorable conditions. Here we show that this widely employed interpretation can be misleading. Multiple models show that stage-structure, long lifespans, and overlapping generations are neither necessary nor sufficient for the storage effect. In models where a robust life-stage does engender a storage effect, it does not do so by preventing stochastic extinction or by improving relative bet-hedging. A robust life-stage is best understood as one of many ways to fulfill an abstract condition for the storage effect: an interaction effect of environment and competition on per capita growth rates. Using a dataset of annual plants from a Mediterranean grassland in Spain, we show that such interaction effects occur between water availability and the number of germinant competitors, leading to storage in the absence of a persistent seed bank. Empiricists hoping to uncover the storage effect should look for interaction effects between environmental conditions and competition -- easily identifiable with multiple regression -- at all stages of a species' life-cycle.