Impact of rainfall seasonality on intraspecific trait variation in a shrub from a Mediterranean climate

Selection pressures along climate gradients give rise to predictable variation in plant functional traits of individual species suggestive of local adaptation. Species whose ranges include winter rainfall, Mediterranean climates, or other strongly seasonal climates, may be exposed to divergent selection pressures at different ends of seasonality gradients. Here, we evaluate how rainfall seasonality in conjunction with other key climatic variables impacts patterns of trait variation in Pelargonium scabrum, a woody shrub from the Greater Cape Floristic Region of South Africa. This biodiversity hotspot encompasses a Mediterranean climate (wet winters and hot, dry summers) and displays steep gradients in temperature and water availability. We used Bayesian regression models to evaluate leaf trait-trait and trait-climate relationships among 26 populations. Models included rainfall seasonality and its interaction with other climate variables (mean annual temperature, mean annual precipitation and potential evapotranspiration) as predictors to test for the impact of climate variation on three leaf traits: size, dissection and leaf mass per area (LMA). We evaluated model explanatory power by calculating Bayesian R-2 values, and predictive power via leave-one-out cross-validation. Trait-trait associations were modulated by rainfall seasonality, including a reversal in the relationship between leaf size and dissection depending on the proportion of rain received in winter. Trait-climate models were improved by including rainfall seasonality as a predictor for both explanatory and predictive power. For leaf dissection and LMA, we detected significant interactions between rainfall seasonality and other environmental variables, leading to reversals in the relationships between these traits and the three environmental variables depending on the proportion of winter rainfall. Differences in the timing of rainfall, coupled with strong differences in the covariation of climate variables, impose divergent selection pressures on P. scabrum populations resulting in divergence of trait values, trait integration and responses to climate gradients. These patterns are consistent with local adaptation of P. scabrum populations mediated by the interactions between temperature and the amount and timing of rainfall. Species arrayed along broad climate gradients represent an excellent opportunity for investigating patterns of trait variation and abundances and distributions of species in relation to future changes in climate. A free Plain Language Summary can be found within the Supporting Information of this article.