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Abstract PremiseSeed germination involves risk; post‐germination conditions might not allow survival and reproduction. Variable, stressful environments favor seeds with germination that avoids risk (e.g., germination in conditions predicting success), spreads risk (e.g., dormancy), or escapes risk (e.g., rapid germination). Germination studies often investigate trait correlations with climate features linked to variation in post‐germination reproductive success. Rarely are long‐term records of population reproductive success available. MethodsSupported by demographic and climate monitoring, we analyzed germination in the California winter‐annualClarkia xantianasubsp.xantiana. Sowing seeds of 10 populations across controlled levels of water potential and temperature, we estimated temperature‐specific base water potential for 20% germination, germination time weighted by water potential above base (hydrotime), and a dormancy index (frequency of viable, ungerminated seeds). Mixed‐effects models analyzed responses to (1) temperature, (2) discrete variation in reproductive success (presence or absence of years with zero seed production by a population), and (3) climate covariates, mean winter precipitation and coefficient of variation (CV) of spring precipitation. For six populations, records enabled analysis with a continuous metric of variable reproduction, the CV of per‐capita reproductive success. ResultsPopulations with more variable reproductive success had higher base water potential and dormancy. Higher base water potential and faster germination occurred at warmer experimental temperatures and in seeds of populations with wetter winters. ConclusionsGeographic variation in seed germination in this species suggests local adaptation to demographic risk and rainfall. High base water potential and dormancy may concentrate germination in years likely to allow reproduction, while spreading risk among years. 

Bet hedging consists of life history strategies that buffer against environmental variability by trading off immediate and long-term fitness. Delayed germination in annual plants is a classic example of bet hedging and is often invoked to explain low germination fractions. We examined whether bet hedging explains low and variable germination fractions among 20 populations of the winter annual plant Clarkia xantiana ssp. xantiana that experience substantial variation in reproductive success among years. Leveraging 15 years of demographic monitoring and 3 years of field germination experiments, we assessed the fitness consequences of seed banks and compared optimal germination fractions from a density-independent bet-hedging model to observed germination fractions. We did not find consistent evidence of bet hedging or the expected trade-off between arithmetic and geometric mean fitness, although delayed germination increased long-term fitness in 7 of 20 populations. Optimal germination fractions were two to five times higher than observed germination fractions, and among-population variation in germination fractions was not correlated with risks across the life cycle. Our comprehensive test suggests that bet hedging is not sufficient to explain the observed germination patterns. Understanding variation in germination strategies will likely require integrating bet hedging with complementary forces shaping the evolution of delayed germination.