Ecotypic variation and stability in growth performance of the thermophilic conifer Pinus halepensis across the Mediterranean basin

Abstract

The combined effects of local adaptation and phenotypic plasticity influencing plant performance are relevant to understand the capacity for genetic responses to climate change. Pinus halepensis is a native species of low- to mid-elevation Mediterranean forests with a high ecological value in drought-prone areas. Thus, it is of utmost importance to determine its adaptive structure for key traits such as growth or survival. Here, we analyse a highly unbalanced dataset collated from different common-garden networks that cover the distribution range of the species. A total of 82 range-wide populations were evaluated in nine Mediterranean trials located in Israel, Italy and Spain. A climate classification of populations allowed for the definition of six different groups, or ecotypes, which showed contrasting performances for tree height and survival at age 15. The effects of ecotypic differentiation and among-ecotypes genetic variation in plasticity were disentangled by fitting stability models accounting for interaction and heteroscedasticity in genotype-by-environment tables. For growth, a Finlay–Wilkinson model suggested high predictability of ecotypic plastic responses in P. halepensis, as described by different linear reaction norms. However, differences in mean height of ca. 15% among ecotypes dominated intra-specific patterns of tree growth across trials, pointing to preponderance of genotypic adaptation over differential ecotypic plasticity in this species. For survival, ecotypic differences were approximately constant across trials, suggesting lack of genotype-by-environment effects. Sub-humid cool climate populations from the eastern Mediterranean (e.g., Greek populations) showed general adaptation and high sensitivity to improved growing conditions, as opposed to populations from the driest ecological extreme of the species (e.g., south Spain and Maghreb populations), which exhibited specific adaptation to harsh environments. Altogether, our results indicate a general adaptive syndrome by which less reactive ecotypes to ameliorated conditions (e.g., non-water-limited) would be associated with high survival rates and low growth. The reported ecotypic differentiation constitutes the basis for tailoring intra-specific responses to climate and disentangling the relationship between adaptive variation and resilience towards climatic warming for this exemplary Mediterranean pine.

This study was funded by the Spanish Government (FUTURPIN project, grant number AGL2015-68274-C3-3-R MINECO/FEDER). We acknowledge the collaboration of the Spanish network of forest genetic trials (GENFORED) and especially of Maria Regina Chambel and Eduardo Notivol for data accessing and technical cooperation. GDM wishes to thank Ernesto Fusaro and Francesco Righi (CREA-PLF) for their contribution in gathering data and for establishing the Italian trials. TK wishes to thank the Merle S. Cahn Foundation and the Monroe and Marjorie Burk Fund for Alternative Energy Studies; Mr. and Mrs. Norman Reiser, together with the Weizmann Center for New Scientists; and the Edith & Nathan Goldberg Career Development Chair.