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We integrate climatic niche models and dated phylogenies to characterize the evolution of climatic niches in Oenothera sections Anogra and Kleinia (Onagraceae), and from that we make inferences on diversification in relation to climate. The evolution of climatic tolerances in Anogra + Kleinia has been heterogeneous, across phylogenetic groups and across different dimensions of climate. All the extant taxa occur in semiarid to arid conditions (annual precipitation of 10.1-49.1 cm and high temperatures in the warmest month of 28.5 degrees-40.1 degrees C), but there is striking variation among taxa in their climatic tolerances, especially temperature (minimum temperatures in the coldest month of -14.0 degrees to 5.3 degrees C) and summer versus winter precipitation (precipitation in the warmest quarter of 0.6-19.4 cm). Climatic disparity is especially pronounced in two subclades (californica, deltoides) that radiated in the southwestern United States and California, apparently including both divergent and convergent evolution of climatic tolerances. This niche evolution is remarkable, given the probable timescale of the radiation (approximately 1 million years). We suggest that the spatiotemporal climatic heterogeneity of western North America has served as a driver of diversification. Our data are also consistent with Axelrod's hypothesis that the spread of arid conditions in western North America stimulated diversification of arid-adapted lineages.

Bet hedging is one solution to the problem of an unpredictably variable environment: fitness in the average environment is sacrificed in favor of lower variation in fitness if this leads to higher long-run stochastic mean fitness. While bet hedging is an important concept in evolutionary ecology, empirical evidence that it occurs is scant. Here we evaluate whether bet hedging occurs via seed banking in natural populations of two species of desert evening primroses (Oenothera, Onagraceae), one annual and one perennial. Four years of data on plants and 3 years of data on seeds yielded two transitions for the entire life cycle. One year was exceptionally dry, leading to reproductive failure in the sample areas, and the other was above average in precipitation, leading to reproductive success in four of five populations. Stochastic simulations of population growth revealed patterns indicative of bet hedging via seed banking, particularly in the annual populations: variance in fitness and fitness in the average environment were lower with seed banking than without, whereas long-run stochastic mean fitness was higher with seed banking than without across a wide range of probabilities of the wet year. This represents a novel, unusually rigorous demonstration of bet hedging from field data.