Tree mortality is a key driver of forest dynamics and its occurrence is projected to increase in the future due to climate change. Despite recent advances in our understanding of the physiological mechanisms leading to death, we still lack robust indicators of mortality risk that could be applied at the individual tree scale. Here, we build on a previous contribution exploring the differences in growth level between trees that died and survived a given mortality event to assess whether changes in temporal autocorrelation, variance, and synchrony in time-series of annual radial growth data can be used as early warning signals of mortality risk. Taking advantage of a unique global ring-width database of 3065 dead trees and 4389 living trees growing together at 198 sites (belonging to 36 gymnosperm and angiosperm species), we analyzed temporal changes in autocorrelation, variance, and synchrony before tree death (diachronic analysis), and also compared these metrics between trees that died and trees that survived a given mortality event (synchronic analysis). Changes in autocorrelation were a poor indicator of mortality risk. However, we found a gradual increase in inter-annual growth variability and a decrease in growth synchrony in the last ∼20 years before mortality of gymnosperms, irrespective of the cause of mortality. These changes could be associated with drought-induced alterations in carbon economy and allocation patterns. In angiosperms, we did not find any consistent changes in any metric. Such lack of any signal might be explained by the relatively high capacity of angiosperms to recover after a stress-induced growth decline. Our analysis provides a robust method for estimating early-warning signals of tree mortality based on annual growth data. In addition to the frequently reported decrease in growth rates, an increase in inter-annual growth variability and a decrease in growth synchrony may be powerful predictors of gymnosperm mortality risk, but not necessarily so for angiosperms.

BC3 Basque Centre for Climate Change Leioa Spain

Biotechnical Faculty University of Ljubljana Ljubljana Slovenia

Boreal Avian Modelling Project Department of Renewable Resources University of Alberta Edmonton AB Canada

Centre for Forest Research Département des Sciences du Bois et de la Forêt Faculté de Foresterie de Géographie et de Géomatique Université Laval Québec QC Canada

Centro de Investigación Forestal Instituto Nacional de Investigación y Tecnología Agraria y Alimentaria Madrid Spain

CNRS IRD EPHE ISEM Université de Montpellier Montpellier France

Consejo Nacional de Investigaciones Científicas y Técnicas CCT Patagonia Norte Río Negro Argentina

CREAF Cerdanyola del Vallès Catalonia Spain

Departament de Biologia Animal de Biologia Vegetal i d'Ecologia Universitat Autònoma de Barcelona Cerdanyola del Vallès Spain

Departamento de Ciencias Agroforestales EiFAB iuFOR University of Valladolid Soria Spain

Department of Agricultural Science Mediterranean University of Reggio Calabria Reggio Calabria Italy

Department of Biogeochemical Processes Max Planck Institute for Biogeochemistry Jena Germany

Department of Biology University of Minnesota Morris Morris MN United States

Department of Biology University of Victoria Victoria BC Canada

Department of Botany Faculty of Science and Technology University of Debrecen Debrecen Hungary

Department of Botany University of Innsbruck Innsbruck Austria

Department of Ecology Universidad Nacional del Comahue Río Negro Argentina

Department of Entomology University of Arkansas Fayetteville AR United States

Department of Environmental Systems Science Forest Ecology Institute of Terrestrial Ecosystems ETH Zürich Zurich Switzerland

Department of Environmental Systems Science Institute of Agricultural Sciences ETH Zürich Zurich Switzerland

Department of Forest Sciences Transilvania University of Brasov Brașov Romania

Department of Forest Sciences University of Helsinki Helsinki Finland

Department of Forestry and Natural Environment Management Technological Educational Institute of Stereas Elladas Karpenisi Greece

Department of Forestry and Wildland Resources Humboldt State University Arcata CA United States

Department of Geography Planning and Recreation Northern Arizona University Flagstaff AZ United States

Department of Geography University of Colorado Boulder CO United States

Department of Physical Chemical and Natural Systems Pablo de Olavide University Seville Spain

Department of Plant and Environmental Sciences Weizmann Institute of Science Rehovot Israel

Department of Research Conservation and Collections Desert Botanical Garden Phoenix AZ United States

Department of Yield and Silviculture Slovenian Forestry Institute Ljubljana Slovenia

Dipartimento di Bioscienze Università degli Studi di Milano Milan Italy

Ecologie des Forêts Méditerranéennes Institut National de la Recherche Agronomique Avignon France

Ecology and Biodiversity Vrije Universiteit Brussel Brussels Belgium

Faculty of Forestry and Wood Sciences Czech University of Life Sciences Prague Czechia

Grupo Ecología Forestal CONICET INTA EEA Bariloche Bariloche Argentina

Institute of Forest Botany and Forest Zoology TU Dresden Dresden Germany

Institute of Lowland Forestry and Environment University of Novi Sad Novi Sad Serbia

Institute of Systematic Botany and Ecology Ulm University Ulm Germany

Instituto de Investigaciones en Biodiversidad y Medioambiente Consejo Nacional de Investigaciones Científicas y Técnicas Río Negro Argentina

Instituto de Investigaciones en Recursos Naturales Agroecología y Desarrollo Rural Sede Andina Universidad Nacional de Río Negro Río Negro Argentina

Instituto Pirenaico de Ecología Zaragoza Spain

Laboratorio de Dendrocronología e Historia Ambiental Instituto Argentino de Nivología Glaciología y Ciencias Ambientales CCT CONICET Mendoza Mendoza Argentina

Laboratory of Wood Biology and Xylarium Royal Museum for Central Africa Tervuren Belgium

National Institute for Research and Development in Forestry Marin Dracea Voluntari Romania

Natural Resources Institute Finland Espoo Finland

Northern Forestry Centre Canadian Forest Service Natural Resources Canada Edmonton AB Canada

Siberian Federal University Krasnoyarsk Russia

Sukachev Institute of Forest Siberian Division of the Russian Academy of Sciences Krasnoyarsk Russia

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

United States Geological Survey Western Ecological Research Center Sequoia and Kings Canyon Field Station Three Rivers CA United States

USDA Forest Service Forest Health Protection Saint Paul MN United States

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