Different drought-tolerance strategies of tree species to cope with increased water stress under climate change in a mixed forest

Trees' functional strategies to cope with extreme drought are essential under climate change. In a mixed Mediterranean forest, we analyzed the functional strategy in response to drought of four co-occurring species (Pinus pinea, Pinus pinaster, Juniperus oxycedrus, and Quercus ilex) during two years. Specifically, we assessed functional traits related to tree water status, leaf water relations, and gas exchange. Different trait-syndrome metrics and the functional strategies under water stress observed suggested a species drought-tolerance differentiation, with the more anysohidric Q. ilex and J. oxycedrus showing a much higher drought tolerance than the more isohydric P. pinea and P. pinaster. All species recovered from negative leaf turgor reached during peak water stress in summer. Q. ilex and J. oxycedrus kept lower leaf osmotic potentials and lower sensitivity of leaf gas exchange and leaf photochemistry to water stress. In contrast, the pine species exhibited more drought-avoidant and water-conservative strategies, yet this behavior was less effective in mitigating water stress's impact on their physiology. The pine species were the most affected by drought, with prolonged near-zero net photosynthesis during summer. P. pinaster was more isohydric than P. pinea and exhibited a lower capacity to maintain leaf turgor. Physiological processes regulating leaf turgor under drought constitute a key functional strategy involved in the carbon and water-related mechanisms, ultimately inducing mortality under hot drought. The currently observed mortality dynamics for P. pinaster, and to a lower extent in P. pinea, may be exacerbated by loss of functional homeostasis.