1. Warming and drought alter plant phenology, photosynthesis and growth with important consequences for the global carbon cycle and the earth's climate. Yet, few studies have attempted to tease apart their effects on tree phenology, particularly leaf senescence, and on source and sink activity.2. We experimentally assessed the single and combined effects of warming and reduced soil moisture on the phenology (leaf-out and senescence date, growing season length) and above-ground sink (height and diameter growth, leaf area and Huber values) and source activity (net photosynthesis, photosynthetic efficiency, chlorophyll concentration and total carbon [C] uptake) of two tree species with distinct strategies to deal with drought: European beech and pubescent oak.3. Warming advanced leaf-out, irrespective of soil moisture levels, particularly in oak and to a lower extent in beech, leading to a prolonged growing season in oak but not beech. No impacts of warming on senescence timing were found for both species. Reduced moisture had little impact on the phenology of both species. Warming-induced advances in phenology and higher photosynthetic efficiency increased the annual C uptake for oak and compensated for the reduced photosynthetic activity in the presence of reduced moisture. Conversely, for beech, source activity, including yearly C uptake, was lower in all treatments than the control, indicating no compensation of the C budget by phenological shifts. Synthesis.Our results demonstrate that a warming-driven earlier activity and higher photosynthetic efficiency compensates for reduced photosynthesis during hot and dry periods, but only for pubescent oak, which is a rather droughttolerant species. Current predictions of warming-induced mitigation effects through extended C uptake seem incorrect for beech.

Increased temperature and prolonged soil moisture reduction have distinct impacts on tree photosynthetic properties. Yet, our knowledge of their combined effect is limited. Moreover, how species interactions alter photosynthetic responses to warming and drought remains unclear.Using mesocosms, we studied how photosynthetic properties of European beech and downy oak were impacted by multi-year warming and soil moisture reduction alone or combined, and how species interactions (intra-vs inter-specific interactions) modulated these effects.Warming of +5°C enhanced photosynthetic properties in oak but not beech, while moisture reduction decreased them in both species. Combined warming and moisture reduction reduced photosynthetic properties for both species, but no exacerbated effects were observed. Oak was less impacted by combined warming and limited moisture when interacting with beech than in intra-specific stands. For beech, species interactions had no impact on the photosynthetic responses to warming and moisture reduction, alone or combined.Warming had either no or beneficial effects on the photosynthetic properties, while moisture reduction and their combined effects strongly reduced photosynthetic responses. However, inter-specific interactions mitigated the adverse impacts of combined warming and drought in oak, thereby highlighting the need to deepen our understanding of the role of species interactions under climate change.

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This article has been accepted for publication and undergone full peer review but has not been through the copyediting, typesetting, pagination and proofreading process, which may lead to differences between this version and the Version of Record. Please cite this article as

Understanding plant trait coordination and variance across climatic gradients is critical for assessing forests’ adaptive potential to climate change. We measured eleven hydraulic, anatomical and leaf-level physiological traits in European beech (Fagus sylvatica L.) along a moisture and temperature gradient in the French Alps. We assessed how traits covaried, and how their population-level variances shifted along the gradient. The intrapopulation variances of vessel size and xylem-specific conductivity reduced in colder locations as narrow vessels were observed in response to low temperature. This decreased individual-level water transport capacity compared to the warmer and more xeric sites. Conversely, the maximum stomatal conductance and Huber value variances were constrained in the arid and warm locations, where trees showed restricted gas exchange and higher xylem-specific conductivity. The populations growing under drier and warmer conditions presented wide variance for the xylem anatomical and hydraulic traits. Our results suggest that short-term physiological acclimation to raising aridity and heat in southern beech populations may occur mainly at the leaf level. Furthermore, the wide variance of the xylem anatomical and hydraulic traits at these sites may be advantageous since more heterogeneous hydraulic conductivity could imply populations’ greater tree-tree complementarity and resilience against climatic variability. Our study highlights that both intrapopulation trait variance and trait network analysis are key approaches for understanding species adaptation and the acclimation potential to a shifting environment.