Drivers of phenotypic variation and plasticity to drought in populations of a Mediterranean shrub along an environmental gradient

Assessing the factors driving intraspecific phenotypic variation is crucial to understand the evolutionary trajectories of plant populations and predict their vulnerability to climate change. Environmental gradients often lead to phenotypic divergence in functional traits and their plasticity across populations. We studied the entire environmental range of the Mediterranean gypsum endemic shrub Helianthemum squamatum to evaluate the factors underlying quantitative population differentiation and phenotypic plasticity to drought, using a common garden with 16 populations that covered the main geographic and the entire climatic range of the species. Sampling followed a hierarchical approach to assess trait genetic variation within and among four distinct geographical regions. We found high but similar plastic responses across populations, which were consistent with adaptive plasticity to drought, including advanced phenology, more sclerophyllous leaves, higher water use efficiency and larger seeds in dry conditions. Despite these generally similar plastic responses, we found significant population differentiation in quantitative traits, part of which was structured at the regional scale. Such differentiation was not associated with environmental variation, including differences in climate and soil conditions. This suggests that non-adaptive processes might have had a role on genetic differentiation in H. squamatum, likely due to the island-like configuration of gypsum habitats and the lack of effective seed dispersal of the study species. Our results emphasize the role of phenotypic plasticity in adaptive drought response and the importance of considering both adaptive and non-adaptive processes shaping intraspecific phenotypic variation, which is crucial for predicting plant population vulnerability to climate change.