Increasing tree diversity is considered a key management option to adapt forests to climate change. However, the effect of species diversity on a forest's ability to cope with extreme drought remains elusive. In this study, we assessed drought tolerance (xylem vulnerability to cavitation) and water stress (water potential), and combined them into a metric of drought-mortality risk (hydraulic safety margin) during extreme 2021 or 2022 summer droughts in five European tree diversity experiments encompassing different biomes. Overall, we found that drought-mortality risk was primarily driven by species identity (56.7% of the total variability), while tree diversity had a much lower effect (8% of the total variability). This result remained valid at the local scale (i.e within experiment) and across the studied European biomes. Tree diversity effect on drought-mortality risk was mediated by changes in water stress intensity, not by changes in xylem vulnerability to cavitation. Significant diversity effects were observed in all experiments, but those effects often varied from positive to negative across mixtures for a given species. Indeed, we found that the composition of the mixtures (i.e., the identities of the species mixed), but not the species richness of the mixture per se, is a driver of tree drought-mortality risk. This calls for a better understanding of the underlying mechanisms before tree diversity can be considered an operational adaption tool to extreme drought. Forest diversification should be considered jointly with management strategies focussed on favouring drought-tolerant species.

AgroParisTech INRAE UMR Silva Université de Lorraine Nancy France

Chair of Silviculture Faculty of Environment and Natural Resources University of Freiburg Freiburg im Breisgau Germany

CIRAD UMR Eco and Sols Montpellier France

Community Ecology Unit Swiss Federal Institute for Forest Snow and Landscape Research WSL Birmensdorf Switzerland

Department of Crop Production Ecology Swedish University of Agricultural Sciences Uppsala Sweden

Department of Forest and Soil Sciences Institute of Forest Ecology University of Natural Resources and Life Sciences Vienna Austria

Department of Forest Protection and Wildlife Management Mendel University in Brno Brno Czech Republic

Department of Forest Sciences ESALQ University of São Paulo Piracicaba Brazil

Eco and Sols Univ Montpellier CIRAD INRAE Institut Agro IRD Montpellier France

Forest and Nature Lab Department of Environment Faculty of Bioscience Engineering Ghent University Melle Gontrode Belgium

French Environment and Energy Management Agency Angers France

Geobotany Faculty of Biology University of Freiburg Freiburg im Breisgau Germany

INRAE Piaf Université Clermont Auvergne Clermont Ferrand France

INRAE UMR BIOGECO University of Bordeaux Pessac France

INRAE Univ Bordeaux Biogeco Cestas France

INRAE URFM Avignon France

Institute of BioEconomy National Research Council Sassari Italy

Instituto de Recursos Naturales y Agrobiologıa Seville Spain

National Biodiversity Future Center S C A R L Palermo Italy

Plant Ecology Research Laboratory PERL School of Architecture Civil and Environmental Engineering EPFL Lausanne Switzerland

UCLouvain Université Catholique de Louvain Earth and Life Institute Louvain La Neuve Belgium

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Multiyear Drought Strengthens Positive and Negative Functional Diversity Effects on Tree Growth Response