Cyclical environments drive variation in life-history strategies: a general theory of cyclical phenology.

Cycles, such as seasons or tides, characterize many systems in nature. Overwhelming evidence shows that climate change-driven alterations to environmental cycles-such as longer seasons-are associated with phenological shifts around the world, suggesting a deep link between environmental cycles and life cycles. However, general mechanisms of life-history evolution in cyclical environments are still not well understood. Here, I build a demographic framework and ask how life-history strategies optimize fitness when the environment perturbs a structured population cyclically and how strategies should change as cyclicality changes. I show that cycle periodicity alters optimality predictions of classic life-history theory because repeated cycles have rippling selective consequences over time and generations. Notably, fitness landscapes that relate environmental cyclicality and life-history optimality vary dramatically depending on which trade-offs govern a given species. The model tuned with known life-history trade-offs in a marine intertidal copepod Tigriopus californicus successfully predicted the shape of life-history variation across natural populations spanning a gradient of tidal periodicities. This framework shows how environmental cycles can drive life-history variation-without complex assumptions of individual responses to cues such as temperature-thus expanding the range of life-history diversity explained by theory and providing a basis for adaptive phenology.