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Tree species diversity in forest ecosystems could reduce their vulnerability to extreme droughts through improved microclimate and below‐ground water source partitioning driven by contrasting species‐specific water use patterns. However, little is known about the seasonal dynamics of belowground water uptake that determine whether diversity positively or negatively impacts tree carbon assimilation and water exchange.Using a network of 30 permanent plots in Mediterranean forests with increasing tree species diversity (from monospecific to four‐species mixtures), we examined the seasonal patterns of in‐situ aboveground carbon and water relations and belowground water sources on 265 trees from four pine and oak species over 2 years using hydraulic and stable isotope approaches.We found that increasing species diversity in broadleaf and conifer mixtures induced strong soil water source partitioning between oak and pine species. As conditions became drier during the summer in mixed stands, oak species took up water from deeper soil sources, while pines were systematically limited to shallow ones. Despite significant belowground moisture partitioning, stronger drought‐induced reductions in photosynthesis, stomatal conductance, and leaf water potential were still observed in diverse compared with monospecific stands for pines but with some benefits for oaks.Synthesis: Our findings reveal that tree species diversity promoted belowground water source partitioning in mixed oak and pine stands, potentially reducing competition for water in more diverse ecosystems. Yet, our results show that it is insufficient to buffer the adverse impacts of severe droughts on aboveground tree carbon and water use, leading to higher water stress, especially for pines. [ABSTRACT FROM AUTHOR]

Heatwaves and soil droughts are increasing in frequency and intensity, leading many tree species to exceed their thermal thresholds, and driving wide‐scale forest mortality. Therefore, investigating heat tolerance and canopy temperature regulation mechanisms is essential to understanding and predicting tree vulnerability to hot droughts. We measured the diurnal and seasonal variation in leaf water potential (Ψ), gas exchange (photosynthesis Anet and stomatal conductance gs), canopy temperature (Tcan), and heat tolerance (leaf critical temperature Tcrit and thermal safety margins TSM, i.e., the difference between maximum Tcan and Tcrit) in three oak species in forests along a latitudinal gradient (Quercus petraea in Switzerland, Quercus ilex in France, and Quercus coccifera in Spain) throughout the growing season. Gas exchange and Ψ of all species were strongly reduced by increased air temperature (Tair) and soil drying, resulting in stomatal closure and inhibition of photosynthesis in Q. ilex and Q. coccifera when Tair surpassed 30°C and soil moisture dropped below 14%. Across all seasons, Tcan was mainly above Tair but increased strongly (up to 10°C > Tair) when Anet was null or negative. Although trees endured extreme Tair (up to 42°C), positive TSM were maintained during the growing season due to high Tcrit in all species (average Tcrit of 54.7°C) and possibly stomatal decoupling (i.e., Anet ≤0 while gs >0). Indeed, Q. ilex and Q. coccifera trees maintained low but positive gs (despite null Anet), decreasing Ψ passed embolism thresholds. This may have prevented Tcan from rising above Tcrit during extreme heat. Overall, our work highlighted that the mechanisms behind heat tolerance and leaf temperature regulation in oak trees include a combination of high evaporative cooling, large heat tolerance limits, and stomatal decoupling. These processes must be considered to accurately predict plant damages, survival, and mortality during extreme heatwaves. [ABSTRACT FROM AUTHOR]

Summary: Mixing species with contrasting resource use strategies could reduce forest vulnerability to extreme events. Yet, how species diversity affects seedling hydraulic responses to heat and drought, including mortality risk, is largely unknown.Using open‐top chambers, we assessed how, over several years, species interactions (monocultures vs mixtures) modulate heat and drought impacts on the hydraulic traits of juvenile European beech and pubescent oak. Using modeling, we estimated species interaction effects on timing to drought‐induced mortality and the underlying mechanisms driving these impacts.We show that mixtures mitigate adverse heat and drought impacts for oak (less negative leaf water potential, higher stomatal conductance, and delayed stomatal closure) but enhance them for beech (lower water potential and stomatal conductance, narrower leaf safety margins, faster tree mortality). Potential underlying mechanisms include oak's larger canopy and higher transpiration, allowing for quicker exhaustion of soil water in mixtures.Our findings highlight that diversity has the potential to alter the effects of extreme events, which would ensure that some species persist even if others remain sensitive. Among the many processes driving diversity effects, differences in canopy size and transpiration associated with the stomatal regulation strategy seem the primary mechanisms driving mortality vulnerability in mixed seedling plantations. See also the Commentary on this article by Wright, 241: 955–957. [ABSTRACT FROM AUTHOR]

Summary: Global warming and droughts push forests closer to their thermal limits, altering tree carbon uptake and growth. To prevent critical overheating, trees can adjust their thermotolerance (Tcrit), temperature and photosynthetic optima (Topt and Aopt), and canopy temperature (Tcan) to stay below damaging thresholds. However, we lack an understanding of how soil droughts affect photosynthetic thermal plasticity and Tcan regulation.In this study, we measured the effect of soil moisture on the seasonal and diurnal dynamics of net photosynthesis (A), stomatal conductance (gs), and Tcan, as well as the thermal plasticity of photosynthesis (Tcrit, Topt, and Aopt), over the course of 1 yr using a long‐term irrigation experiment in a drought‐prone Pinus sylvestris forest in Switzerland.Irrigation resulted in higher needle‐level A, gs, Topt, and Aopt compared with naturally drought‐exposed trees. No daily or seasonal differences in Tcan were observed between treatments. Trees operated below their thermal thresholds (Tcrit), independently of soil moisture content.Despite strong Tcan and Tair coupling, we provide evidence that drought reduces trees' temperature optimum due to a substantial reduction of gs during warm and dry periods of the year. These findings provide important insights regarding the effects of soil drought on the thermal tolerance of P. sylvestris. [ABSTRACT FROM AUTHOR]