Optimal germination timing in unpredictable environments: the importance of dormancy for both among- and within-season variation.

For organisms living in unpredictable environments, timing important life-history events is challenging. One way to deal with uncertainty is to spread the emergence of offspring across multiple years via dormancy. However, timing of emergence is not only important among years, but also within each growing season. Here, we study the evolutionary interactions between germination strategies that deal with among- and within-season uncertainty. We use a modelling approach that considers among-season dormancy and within-season germination phenology of annual plants as potentially independent traits and study their separate and joint evolution in a variable environment. We find that higher among-season dormancy selects for earlier germination within the growing season. Furthermore, our results indicate that more unpredictable natural environments can counter-intuitively select for less risk-spreading within the season. Furthermore, strong priority effects select for earlier within-season germination phenology which in turn increases the need for bet hedging through among-season dormancy.