title: A synthesis of radial growth patterns preceding tree mortality.
abstract: Tree mortality is a key factor influencing forest functions and dynamics, but our understanding of the mechanisms leading to mortality and the associated changes in tree growth rates are still limited. We compiled a new pan-continental tree-ring width database from sites where both dead and living trees were sampled (2970 dead and 4224 living trees from 190 sites, including 36 species), and compared early and recent growth rates between trees that died and those that survived a given mortality event. We observed a decrease in radial growth before death in ca. 84% of the mortality events. The extent and duration of these reductions were highly variable (1-100 years in 96% of events) due to the complex interactions among study species and the source(s) of mortality. Strong and long-lasting declines were found for gymnosperms, shade- and drought-tolerant species, and trees that died from competition. Angiosperms and trees that died due to biotic attacks (especially bark-beetles) typically showed relatively small and short-term growth reductions. Our analysis did not highlight any universal trade-off between early growth and tree longevity within a species, although this result may also reflect high variability in sampling design among sites. The intersite and interspecific variability in growth patterns before mortality provides valuable information on the nature of the mortality process, which is consistent with our understanding of the physiological mechanisms leading to mortality. Abrupt changes in growth immediately before death can be associated with generalized hydraulic failure and/or bark-beetle attack, while long-term decrease in growth may be associated with a gradual decline in hydraulic performance coupled with depletion in carbon reserves. Our results imply that growth-based mortality algorithms may be a powerful tool for predicting gymnosperm mortality induced by chronic stress, but not necessarily so for angiosperms and in case of intense drought or bark-beetle outbreaks.
authors.affiliation: Forest Ecology, Department of Environmental Systems Science, Institute of Terrestrial Ecosystems, ETH Zürich, Universitätstrasse 22, 8092, Zürich, Switzerland.
authors.forename: Maxime
authors.initials: M
authors.lastname: Cailleret
authors.affiliation: Institute of Systematic Botany and Ecology, Ulm University, Albert-Einstein-Allee 11, 89081, Ulm, Germany.
authors.forename: Steven
authors.initials: S
authors.lastname: Jansen
authors.affiliation: CREAF, Campus UAB, 08193, Cerdanyola del Vallès, Spain.|Laboratory of Plant Biology and Nature Management (APNA), Vrije Universiteit Brussel, Pleinlaan 2, 1050, Brussels, Belgium.|Laboratory of Wood Biology and Xylarium, Royal Museum for Central Africa (RMCA), Leuvensesteenweg 13, 3080, Tervuren, Belgium.
authors.forename: Elisabeth M R
authors.initials: EM
authors.lastname: Robert
authors.affiliation: Department of Life Sciences, Centre for Functional Ecology, University of Coimbra, Calçada Martim de Freitas, 3000-456, Coimbra, Portugal.
authors.forename: Lucía
authors.initials: L
authors.lastname: Desoto
authors.affiliation: Department of Forest Sciences, University of Helsinki, P.O. Box 27 (Latokartanonkaari 7), 00014, Helsinki, Finland.
authors.forename: Tuomas
authors.initials: T
authors.lastname: Aakala
authors.affiliation: Department of Biology, University of Victoria, PO Box 3020, STN CSC, Victoria, BC, V8W 3N5, Canada.
authors.forename: Joseph A
authors.initials: JA
authors.lastname: Antos
authors.affiliation: Institute of Botany, University of Innsbruck, Sternwartestrasse 15, 6020, Innsbruck, Austria.
authors.forename: Barbara
authors.initials: B
authors.lastname: Beikircher
authors.affiliation: Forest Ecology, Department of Environmental Systems Science, Institute of Terrestrial Ecosystems, ETH Zürich, Universitätstrasse 22, 8092, Zürich, Switzerland.
authors.forename: Christof
authors.initials: C
authors.lastname: Bigler
authors.affiliation: Forest Ecology, Department of Environmental Systems Science, Institute of Terrestrial Ecosystems, ETH Zürich, Universitätstrasse 22, 8092, Zürich, Switzerland.
authors.forename: Harald
authors.initials: H
authors.lastname: Bugmann
authors.affiliation: Dipartimento di Bioscienze, Università degli Studi di Milano, Via Giovanni Celoria 26, 20133, Milano, Italy.
authors.forename: Marco
authors.initials: M
authors.lastname: Caccianiga
authors.affiliation: Faculty of Forestry and Wood Sciences, Czech University of Life Sciences, Kamýcká 961/129, 165 21, Praha 6-Suchdol, Czech Republic.
authors.forename: Vojtěch
authors.initials: V
authors.lastname: Čada
authors.affiliation: Instituto Pirenaico de Ecología (IPE-CSIC), Avenida Montañana 1005, 50192, Zaragoza, Spain.
authors.forename: Jesus J
authors.initials: JJ
authors.lastname: Camarero
authors.affiliation: Swiss Federal Institute for Forest, Snow and Landscape Research - WSL, Zürcherstrasse 111, 8903, Birmensdorf, Switzerland.
authors.forename: Paolo
authors.initials: P
authors.lastname: Cherubini
authors.affiliation: Unité Mixte de Recherche (UMR) 547 PIAF, Institut National de la Recherche Agronomique (INRA), Université Clermont Auvergne, 63100, Clermont-Ferrand, France.
authors.forename: Hervé
authors.initials: H
authors.lastname: Cochard
authors.affiliation: Département des sciences du bois et de la forêt, Centre for Forest Research, Faculté de foresterie, de géographie et de géomatique, Université Laval, 2405 rue de la Terrasse, Québec, QC, G1V 0A6, Canada.
authors.forename: Marie R
authors.initials: MR
authors.lastname: Coyea
authors.affiliation: Biotechnical Faculty, University of Ljubljana, Jamnikarjeva 101, 1000, Ljubljana, Slovenia.
authors.forename: Katarina
authors.initials: K
authors.lastname: Čufar
authors.affiliation: U.S. Geological Survey, Western Ecological Research Center, 47050 Generals Highway, Three Rivers, CA, 93271, USA.
authors.forename: Adrian J
authors.initials: AJ
authors.lastname: Das
authors.affiliation: Ecologie des Forest Méditerranéennes (URFM), Institut National de la Recherche Agronomique (INRA), Domaine Saint Paul, Site Agroparc, 84914, Avignon Cedex 9, France.
authors.forename: Hendrik
authors.initials: H
authors.lastname: Davi
authors.affiliation: Unité Mixte de Recherche (UMR) 1202 BIOGECO, Institut National de la Recherche Agronomique (INRA), Université de Bordeaux, 33615, Pessac, France.
authors.forename: Sylvain
authors.initials: S
authors.lastname: Delzon
authors.affiliation: Department of Geography and Environmental Development, Ben-Gurion University of the Negev, 84105, Beer-Sheva, Israel.
authors.forename: Michael
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authors.lastname: Dorman
authors.affiliation: Centro de Investigación Forestal (CIFOR), Instituto Nacional de Investigación y Tecnología Agraria y Alimentaria (INIA), Carretera La Coruña km 7.5, 28040, Madrid, Spain.
authors.forename: Guillermo
authors.initials: G
authors.lastname: Gea-Izquierdo
authors.affiliation: Institute of Forest Botany and Forest Zoology, TU Dresden, 01062, Dresden, Germany.|Fachgebiet Vegetationstechnik und Pflanzenverwendung, Institut für Landschaftsarchitektur und Umweltplanung, TU Berlin, 10623, Berlin, Germany.
authors.forename: Sten
authors.initials: S
authors.lastname: Gillner
authors.affiliation: Department of Entomology, University of Arkansas, Fayetteville, AR, 72701, USA.|Department of Ecology and Evolutionary Biology, University of Kansas, 1450 Jayhawk Boulevard, Lawrence, KS, 66045, USA.
authors.forename: Laurel J
authors.initials: LJ
authors.lastname: Haavik
authors.affiliation: Max-Planck Institute for Biogeochemistry, Hans Knöll Strasse 10, 07745, Jena, Germany.
authors.forename: Henrik
authors.initials: H
authors.lastname: Hartmann
authors.affiliation: CREAF, Campus UAB, 08193, Cerdanyola del Vallès, Spain.|Department of Biogeography and Global Change, National Museum of Natural History (MNCN), Consejo Superior de Investigaciones Científicas (CSIC), C/Serrano 115bis, 28006, Madrid, Spain.
authors.forename: Ana-Maria
authors.initials: AM
authors.lastname: Hereş
authors.affiliation: Department of Research, Conservation and Collections, Desert Botanical Garden, 1201 N Galvin Parkway, Phoenix, AZ, USA.
authors.forename: Kevin R
authors.initials: KR
authors.lastname: Hultine
authors.affiliation: Faculty of Forestry and Wood Sciences, Czech University of Life Sciences, Kamýcká 961/129, 165 21, Praha 6-Suchdol, Czech Republic.
authors.forename: Pavel
authors.initials: P
authors.lastname: Janda
authors.affiliation: Department of Forestry and Wildland Resources, Humboldt State University, 1 Harpst Street, Arcata, CA, 95521, USA.
authors.forename: Jeffrey M
authors.initials: JM
authors.lastname: Kane
authors.affiliation: Siberian Division of the Russian Academy of Sciences (RAS), Sukachev Institute of Forest, Krasnoyarsk, 660036, Russia.
authors.forename: Vyacheslav I
authors.initials: VI
authors.lastname: Kharuk
authors.affiliation: Department of Ecology, Universidad Nacional del Comahue, Quintral S/N, Barrio Jardín Botánico, 8400, San Carlos de Bariloche, Río Negro, Argentina.|Instituto de Investigaciones de Biodiversidad y Medio Ambiente (INIBOMA), Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET), Quintral 1250, 8400, San Carlos de Bariloche, Río Negro, Argentina.
authors.forename: Thomas
authors.initials: T
authors.lastname: Kitzberger
authors.affiliation: Institute of Soil, Water, and Environmental Sciences, Volcani Center, Agricultural Research Organization (ARO), PO Box 6, 50250, Beit Dagan, Israel.
authors.forename: Tamir
authors.initials: T
authors.lastname: Klein
authors.affiliation: Alterra - Green World Research, Wageningen University, Droevendaalse steeg 1, 6700AA, Wageningen, The Netherlands.
authors.forename: Koen
authors.initials: K
authors.lastname: Kramer
authors.affiliation: Naturalis Biodiversity Center, Leiden University, PO Box 9517, 2300RA, Leiden, The Netherlands.
authors.forename: Frederic
authors.initials: F
authors.lastname: Lens
authors.affiliation: Department of Yield and Silviculture, Slovenian Forestry Institute, Večna pot 2, 1000, Ljubljana, Slovenia.
authors.forename: Tom
authors.initials: T
authors.lastname: Levanic
authors.affiliation: Department of Physical, Chemical and Natural Systems, Pablo de Olavide University, Carretera de Utrera km 1, 41013, Seville, Spain.
authors.forename: Juan C
authors.initials: JC
authors.lastname: Linares Calderon
authors.affiliation: CREAF, Campus UAB, 08193, Cerdanyola del Vallès, Spain.|Universitat Autònoma de Barcelona, 08193, Cerdanyola del Vallès, Spain.
authors.forename: Francisco
authors.initials: F
authors.lastname: Lloret
authors.affiliation: Forest Research Centre, School of Agriculture, University of Lisbon, Tapada da Ajuda, 1349-017, Lisboa, Portugal.
authors.forename: Raquel
authors.initials: R
authors.lastname: Lobo-Do-Vale
authors.affiliation: Department of Agricultural Science, Mediterranean University of Reggio Calabria, loc. Feo di Vito, 89060, Reggio Calabria, Italy.
authors.forename: Fabio
authors.initials: F
authors.lastname: Lombardi
authors.affiliation: Forest Genetics and Physiology Research Group, Technical University of Madrid, Calle Ramiro de Maeztu 7, 28040, Madrid, Spain.|Hawkesbury Institute for the Environment, University of Western Sydney, Science Road, Richmond, NSW, 2753, Australia.
authors.forename: Rosana
authors.initials: R
authors.lastname: López Rodríguez
authors.affiliation: Natural Resources Institute Finland (Luke), Viikinkaari 4, 00790, Helsinki, Finland.
authors.forename: Harri
authors.initials: H
authors.lastname: Mäkinen
authors.affiliation: Institute of Botany, University of Innsbruck, Sternwartestrasse 15, 6020, Innsbruck, Austria.
authors.forename: Stefan
authors.initials: S
authors.lastname: Mayr
authors.affiliation: Department of Botany, Faculty of Science and Technology, University of Debrecen, Egyetem tér 1, 4032, Debrecen, Hungary.
authors.forename: Ilona
authors.initials: I
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authors.affiliation: Northern Forestry Centre, Canadian Forest Service, Natural Resources Canada, 5320-122nd Street, Edmonton, AB, T6H 3S5, Canada.
authors.forename: Juha M
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authors.affiliation: Department of Forest Sciences, University of Helsinki, P.O. Box 27 (Latokartanonkaari 7), 00014, Helsinki, Finland.
authors.forename: Francesco
authors.initials: F
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authors.affiliation: Institute of Botany, University of Innsbruck, Sternwartestrasse 15, 6020, Innsbruck, Austria.
authors.forename: Walter
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authors.affiliation: Department of Forestry and Natural Environment Management, Technological Educational Institute (TEI) of Stereas Elladas, Ag Georgiou 1, 36100, Karpenissi, Greece.
authors.forename: Andreas
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authors.affiliation: Natural Resources Institute Finland (Luke), PO Box 18 (Jokiniemenkuja 1), 01301, Vantaa, Finland.
authors.forename: Mikko
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authors.affiliation: Swiss Federal Institute for Forest, Snow and Landscape Research - WSL, Zürcherstrasse 111, 8903, Birmensdorf, Switzerland.|National Institute for Research-Development in Forestry ''Marin Dracea'', Eroilor 128, 077190, Voluntari, Romania.
authors.forename: Any M
authors.initials: AM
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authors.affiliation: Forest Ecology, Department of Environmental Systems Science, Institute of Terrestrial Ecosystems, ETH Zürich, Universitätstrasse 22, 8092, Zürich, Switzerland.|Swiss Federal Institute for Forest, Snow and Landscape Research - WSL, Zürcherstrasse 111, 8903, Birmensdorf, Switzerland.
authors.forename: Brigitte
authors.initials: B
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authors.affiliation: Instituto Pirenaico de Ecología (IPE-CSIC), Avenida Montañana 1005, 50192, Zaragoza, Spain.
authors.forename: Gabriel
authors.initials: G
authors.lastname: Sangüesa-Barreda
authors.affiliation: Faculty of Pure and Applied Sciences, Open University of Cyprus, Latsia, 2252, Nicosia, Cyprus.|Department of Biological Sciences, University of Cyprus, PO Box 20537, 1678, Nicosia, Cyprus.|Division of Plant Biology, Department of Biology, University of Patras, 26500, Patras, Greece.
authors.forename: Dimitrios
authors.initials: D
authors.lastname: Sarris
authors.affiliation: Department of Geography, University of Colorado, Boulder, CO, 80309-0260, USA.
authors.forename: Jeremy M
authors.initials: JM
authors.lastname: Smith
authors.affiliation: Department of Geography, Planning and Recreation, Northern Arizona University, PO Box 15016, Flagstaff, AZ, 86011, USA.
authors.forename: Amanda B
authors.initials: AB
authors.lastname: Stan
authors.affiliation: Forest Ecology and Forest Management Group, Wageningen University, Droevendaalsesteeg 3a, 6708 PB, Wageningen, The Netherlands.
authors.forename: Frank
authors.initials: F
authors.lastname: Sterck
authors.affiliation: Institute of Lowland Forestry and Environment, University of Novi Sad, Antona Cehova 13, PO Box 117, 21000, Novi Sad, Serbia.
authors.forename: Dejan B
authors.initials: DB
authors.lastname: Stojanović
authors.affiliation: Instituto de Investigaciones de Biodiversidad y Medio Ambiente (INIBOMA), Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET), Quintral 1250, 8400, San Carlos de Bariloche, Río Negro, Argentina.
authors.forename: Maria L
authors.initials: ML
authors.lastname: Suarez
authors.affiliation: Faculty of Forestry and Wood Sciences, Czech University of Life Sciences, Kamýcká 961/129, 165 21, Praha 6-Suchdol, Czech Republic.
authors.forename: Miroslav
authors.initials: M
authors.lastname: Svoboda
authors.affiliation: Dipartimenti di Bioscienze e Territorio, Università del Molise, C. da Fonte Lappone, 86090, Pesche, Italy.|European Forest Institute (EFI) Project Centre on Mountain Forests (MOUNTFOR), Via E. Mach 1, 38010, San Michele all'Adige, Italy.
authors.forename: Roberto
authors.initials: R
authors.lastname: Tognetti
authors.affiliation: Unité Mixte de Recherche (UMR) 1202 BIOGECO, Institut National de la Recherche Agronomique (INRA), Université de Bordeaux, 33615, Pessac, France.
authors.forename: José M
authors.initials: JM
authors.lastname: Torres-Ruiz
authors.affiliation: Faculty of Forestry and Wood Sciences, Czech University of Life Sciences, Kamýcká 961/129, 165 21, Praha 6-Suchdol, Czech Republic.
authors.forename: Volodymyr
authors.initials: V
authors.lastname: Trotsiuk
authors.affiliation: Laboratorio de Dendrocronología e Historia Ambiental, Instituto Argentino de Nivología, Glaciología y Ciencias Ambientales (IANIGLA), CCT CONICET Mendoza, Av. Ruiz Leal s/n, Parque General San Martín, Mendoza, CP 5500, Argentina.
authors.forename: Ricardo
authors.initials: R
authors.lastname: Villalba
authors.affiliation: Institute of Forestry and Rural Engineering, Estonian University of Life Sciences, Kreutzwaldi 5, 51014, Tartu, Estonia.
authors.forename: Floor
authors.initials: F
authors.lastname: Vodde
authors.affiliation: Boreal Avian Modelling Project, Department of Renewable Resources, University of Alberta, 751 General Services Building, Edmonton, AB, T6G 2H1, Canada.
authors.forename: Alana R
authors.initials: AR
authors.lastname: Westwood
authors.affiliation: University of Minnesota, 600 East 4th Street, Morris, MN, 56267, USA.
authors.forename: Peter H
authors.initials: PH
authors.lastname: Wyckoff
authors.affiliation: University of Forestry, Kliment Ohridski Street 10, 1756, Sofia, Bulgaria.
authors.forename: Nikolay
authors.initials: N
authors.lastname: Zafirov
authors.affiliation: CREAF, Campus UAB, 08193, Cerdanyola del Vallès, Spain.|Universitat Autònoma de Barcelona, 08193, Cerdanyola del Vallès, Spain.
authors.forename: Jordi
authors.initials: J
authors.lastname: Martínez-Vilalta
mesh_terms: D000818:Animals; D002244:Carbon; D001517:Coleoptera; D055864:Droughts; D013312:Stress, Physiological; D014197:Trees / Q000254:growth & development*
chemical_list: D002244:Carbon
keywords: angiosperms; death; drought; growth; gymnosperms; pathogens; ring-width; tree mortality