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

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