Chronic warming and dry soils limit carbon uptake and growth despite a longer growing season in beech and oak.

Progressively warmer and drier climatic conditions impact tree phenology and carbon cycling with large consequences for forest carbon balance. However, it remains unclear how individual impacts of warming and drier soils differ from their combined effects and how species interactions modulate tree responses. Using mesocosms, we assessed the multiyear impact of continuous air warming and lower soil moisture alone or in combination on phenology, leaf-level photosynthesis, nonstructural carbohydrate concentrations, and aboveground growth of young European beech (Fagus sylvatica L.) and Downy oak (Quercus pubescens Willd.) trees. We further tested how species interactions (in monocultures and in mixtures) modulated these effects. Warming prolonged the growing season of both species but reduced growth in oak. In contrast, lower moisture did not impact phenology but reduced carbon assimilation and growth in both species. Combined impacts of warming and drier soils did not differ from their single effects. Under warmer and drier conditions, performances of both species were enhanced in mixtures compared to monocultures. Our work revealed that higher temperature and lower soil moisture have contrasting impacts on phenology vs. leaf-level assimilation and growth, with the former being driven by temperature and the latter by moisture. Furthermore, we showed a compensation in the negative impacts of chronic heat and drought by tree species interactions.

Interactions between beech and oak seedlings can modify the effects of hotter droughts and the onset of hydraulic failure.

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.

High heat tolerance, evaporative cooling, and stomatal decoupling regulate canopy temperature and their safety margins in three European oak species.

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.

Absence of canopy temperature variation despite stomatal adjustment in Pinus sylvestris under multidecadal soil moisture manipulation.

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.

Impact of warmer and drier conditions on tree photosynthetic properties and the role of species interactions.

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.

Aridity and cold temperatures drive divergent adjustments of European beech xylem anatomy, hydraulics and leaf physiological traits.

Understanding plant trait coordination and variance across climatic gradients is critical for assessing forests' adaptive potential to climate change. We measured 11 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 with 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.