Evenness mediates the global relationship between forest productivity and richness
Journal of Ecology, vol. 111, pp. 1308–1326
Abstract
Abstract 1. Biodiversity is an important component of natural ecosystems, with higher species richness often correlating with an increase in ecosystem productivity. Yet, this relationship varies substantially across environments, typically becoming less pronounced at high levels of species richness. However, species richness alone cannot reflect all important properties of a community, including community evenness, which may mediate the relationship between biodiversity and productivity. If the evenness of a community correlates negatively with richness across forests globally, then a greater number of species may not always increase overall diversity and productivity of the system. Theoretical work and local empirical studies have shown that the effect of evenness on ecosystem functioning may be especially strong at high richness levels, yet the consistency of this remains untested at a global scale. 2. Here, we used a dataset of forests from across the globe, which includes composition, biomass accumulation and net primary productivity, to explore whether productivity correlates with community evenness and richness in a way that evenness appears to buffer the effect of richness. Specifically, we evaluated whether low levels of evenness in speciose communities correlate with the attenuation of the richness–productivity relationship. 3. We found that tree species richness and evenness are negatively correlated across forests globally, with highly speciose forests typically comprising a few dominant and many rare species. Furthermore, we found that the correlation between diversity and productivity changes with evenness: at low richness, uneven communities are more productive, while at high richness, even communities are more productive. 4. Synthesis . Collectively, these results demonstrate that evenness is an integral component of the relationship between biodiversity and productivity, and that the attenuating effect of richness on forest productivity might be partly explained by low evenness in speciose communities. Productivity generally increases with species richness, until reduced evenness limits the overall increases in community diversity. Our research suggests that evenness is a fundamental component of biodiversity–ecosystem function relationships, and is of critical importance for guiding conservation and sustainable ecosystem management decisions.
Authors 100
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Iris Hordijk corresponding
ETH Zurich · Institute of Integrative Biology
Affiliation as printed
Institute of Integrative Biology, ETH Zurich (Swiss Federal Institute of Technology) Universitätsstrasse 16 8092 Zurich Switzerland
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ETH Zurich · Institute of Integrative Biology
Affiliation as printed
Institute of Integrative Biology, ETH Zurich (Swiss Federal Institute of Technology) Universitätsstrasse 16 8092 Zurich Switzerland
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Affiliation as printed
School of Biological Sciences University of Queensland Brisbane Australia
School of Biological Sciences, University of Queensland, Brisbane, Australia
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ETH Zurich · Institute of Integrative Biology
Affiliation as printed
Institute of Integrative Biology, ETH Zurich (Swiss Federal Institute of Technology) Universitätsstrasse 16 8092 Zurich Switzerland
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Hans ter Steege Aachen Naturalis Biodiversity Center Leiden The Netherlands Naturalis Biodiversity Center
Naturalis Biodiversity Center · Vrije Universiteit Amsterdam
Affiliation as printed
Naturalis Biodiversity Centre Leiden The Netherlands
Systems Ecology Free University Amsterdam Amsterdam The Netherlands
Naturalis Biodiversity Centre, Leiden, The Netherlands
Systems Ecology, Free University Amsterdam, Amsterdam, The Netherlands
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Purdue University West Lafayette
Affiliation as printed
Department of Forestry and Natural Resources Purdue University West Lafayette Indiana USA
Department of Forestry and Natural Resources, Purdue University, West Lafayette, Indiana, USA
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Universitat de Lleida · Forest Science and Technology Centre of Catalonia · Agrotecnio - Centre for Food and Agriculture Research
Affiliation as printed
Department of Crop and Forest Sciences University of Lleida Lleida Spain
Joint Research Unit CTFC—AGROTECNIO—CERCA Solsona Spain
Department of Crop and Forest Sciences, University of Lleida, Lleida, Spain
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Wageningen University & Research
Affiliation as printed
Wageningen University and Research Wageningen The Netherlands
Wageningen University and Research, Wageningen, The Netherlands
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University of Minnesota · Western Sydney University
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Department of Forest Resources University of Minnesota St Paul Minnesota USA
Hawkesbury Institute for the Environment Western Sydney University Penrith New South Wales Australia
Department of Forest Resources, University of Minnesota, St Paul, Minnesota, USA
Hawkesbury Institute for the Environment, Western Sydney University, Penrith, New South Wales, Australia
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Swiss Federal Institute for Forest, Snow and Landscape Research
Affiliation as printed
Swiss Federal Institute for Forest, Snow and Landscape Research, WSL Birmensdorf Switzerland
Swiss Federal Institute for Forest, Snow and Landscape Research, WSL, Birmensdorf, Switzerland
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Université Félix Houphouët-Boigny
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UFR Biosciences University Félix Houphouët‐Boigny Abidjan Côte d'Ivoire
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University of Udine · Free University of Bozen-Bolzano
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Department of Agricultural, Food, Environmental and Animal Sciences University of Udine Udine Italy
Faculty of Science and Technology Free University of Bolzano Bolzano Italy
Department of Agricultural, Food, Environmental and Animal Sciences, University of Udine, Udine, Italy
Faculty of Science and Technology, Free University of Bolzano, Bolzano, Italy
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Affiliation as printed
Spatial Ecology and Conservation Laboratory, Center for Latin American Studies University of Florida Gainesville Florida 32611 USA
Spatial Ecology and Conservation Laboratory, Center for Latin American Studies, University of Florida, Gainesville, Florida, 32611 USA
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Instituto Tecnológico de Costa Rica
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Forestry School, Tecnológico de Costa Rica TEC Cartago Costa Rica
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Fundación Con Vida · Universidad Nacional Abierta y a Distancia
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Fundacion ConVida Universidad Nacional Abierta y a Distancia, UNAD Medellin Colombia
Fundacion ConVida, Universidad Nacional Abierta y a Distancia, UNAD, Medellin, Colombia
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Field Museum of Natural History
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Field Museum of Natural History Chicago Illinois USA
Field Museum of Natural History, Chicago, Illinois, USA
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University of California, Los Angeles
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Center for Tropical Research, Institute of the Environment and Sustainability, UCLA Los Angeles California USA
Center for Tropical Research, Institute of the Environment and Sustainability, UCLA, Los Angeles, California, USA
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Affiliation as printed
Silviculture and Forest Ecology of the Temperate Zones University of Göttingen Göttingen Germany
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Norwegian Institute of Bioeconomy Research
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Division of Forest and Forest Resources Norwegian Institute of Bioeconomy Research (NIBIO) Ås Norway
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Museo de Historia Natural Noel Kempff Mercado
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Museo de Historia natural Noel kempff Mercado Santa Cruz Bolivia
Museo de Historia natural Noel kempff Mercado, Santa Cruz, Bolivia
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Museo de Historia Natural Noel Kempff Mercado
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Museo de Historia natural Noel kempff Mercado Santa Cruz Bolivia
Museo de Historia natural Noel kempff Mercado, Santa Cruz, Bolivia
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European Commission · Joint Research Centre
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European Commission Joint Research Centre Ispra Italy
European Commission, Joint Research Centre, Ispra, Italy
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Universidad Nacional Experimental de los Llanos Occidentales Ezequiel Zamora
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UNELLEZ‐Guanare, Programa de Ciencias del Agro y el Mar Herbario Universitario (PORT), Portuguesa, Venezuela Compensation International S. A. Ci Progress‐GreenLife Bogotá Distrito Capital Colombia
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School of Geography University of Leeds Leeds UK
School of Geography, University of Leeds, Leeds, UK
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Department of Geomatics Forest Research Institute Raszyn Poland
Department of Geomatics, Forest Research Institute, Raszyn, Poland
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Systems Ecology Free University Amsterdam Amsterdam The Netherlands
Systems Ecology, Free University Amsterdam, Amsterdam, The Netherlands
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Centro Multidisciplinar Universidade Federal do Acre Rio Branco Brazil
Centro Multidisciplinar, Universidade Federal do Acre, Rio Branco, Brazil
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National Academy of Sciences · Duke University
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Department of Evolutionary Anthropology Duke University Durham North Carolina 27708 USA
Proceedings of the National Academy of Sciences Washington DC USA
Department of Evolutionary Anthropology, Duke University, Durham, North Carolina, 27708 USA
Proceedings of the National Academy of Sciences, Washington, DC, USA
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University of Liège · Gembloux Agro-Bio Tech
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Gembloux Agro Bio‐Tech University of Liege Liege Belgium
Gembloux Agro Bio-Tech, University of Liege, Liege, Belgium
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United Nation Framework Convention on Climate Change Bonn Germany
United Nation Framework Convention on Climate Change, Bonn, Germany
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Centre National de la Recherche Scientifique · Université de Montpellier · Institut National de Recherche pour l'Agriculture, l'Alimentation et l'Environnement · UMR Botanique et Modélisation de l’Architecture des Plantes et des végétations · Institut de Recherche pour le Développement
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Cirad, UMR‐AMAP, CNRS, INRA, IRD Université de Montpellier Montpellier France
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Mbarara University of Science and Technology
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Institute of Tropical Forest Conservation, Mbarara University of Sciences and Technology Mbarara Uganda
Institute of Tropical Forest Conservation, Mbarara University of Sciences and Technology, Mbarara, Uganda
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Isotope Bioscience Laboratory—ISOFYS Ghent University Ghent Belgium
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Wageningen University & Research
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Wageningen University and Research Wageningen The Netherlands
Wageningen University and Research, Wageningen, The Netherlands
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Ştefan cel Mare University of Suceava
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Integrated Center for Research, Development and Innovation in Advanced Materials, Nanotechnologies, and Distributed Systems for Fabrication and Control (MANSiD) Stefan cel Mare University of Suceava Suceava Romania
Integrated Center for Research, Development and Innovation in Advanced Materials, Nanotechnologies, and Distributed Systems for Fabrication and Control (MANSiD), Stefan cel Mare University of Suceava, Suceava, Romania
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Department of Forest Sciences, Luiz de Queiroz College of Agriculture University of São Paulo Piracicaba Brazil
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Bavarian State Institute of Forestry Freising Germany
Bavarian State Institute of Forestry, Freising, Germany
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School of Geography University of Leeds Leeds UK
School of Geography, University of Leeds, Leeds, UK
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Spatial Ecology and Conservation Laboratory, School of Forest, Fisheries, and Geomatics Sciences University of Florida Gainesville Florida 32611 USA
Spatial Ecology and Conservation Laboratory, School of Forest, Fisheries, and Geomatics Sciences, University of Florida, Gainesville, Florida, 32611 USA
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German Centre for Integrative Biodiversity Research · Martin Luther University Halle-Wittenberg
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German Centre for Integrative Biodiversity Research (iDiv) Halle‐Jena‐Leipzig Leipzig Germany
Institute of Biology, Geobotany and Botanical Garden, Martin Luther University Halle‐Wittenberg Halle‐Wittenberg Germany
German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig, Leipzig, Germany
Institute of Biology, Geobotany and Botanical Garden, Martin Luther University Halle-Wittenberg, Halle-Wittenberg, Germany
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Department of Agriculture, Food, Environment and Forest (DAGRI) University of Firenze Florence Italy
Department of Agriculture, Food, Environment and Forest (DAGRI), University of Firenze, Florence, Italy
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National Research Tomsk State University
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Biological Institute, Tomsk State University Tomsk Russia
Biological Institute, Tomsk State University, Tomsk, Russia
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Department of Forest Sciences, Luiz de Queiroz College of Agriculture University of São Paulo Piracicaba Brazil
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Department of Spatial Regulation, GIS and Forest Policy Institute of Forestry Belgrade Serbia
Department of Spatial Regulation, GIS and Forest Policy, Institute of Forestry, Belgrade, Serbia
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University of Connecticut · University of the Sunshine Coast
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Department of Ecology and Evolutionary Biology University of Connecticut Storrs Connecticut USA
Tropical Forests and People Research Centre, University of the Sunshine Coast Maroochydore Queensland Australia
Department of Ecology and Evolutionary Biology, University of Connecticut, Storrs, Connecticut, USA
Tropical Forests and People Research Centre, University of the Sunshine Coast, Maroochydore, Queensland, Australia
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Faculty of Natural Resources Management Lakehead University Thunder Bay Ontario Canada
Faculty of Natural Resources Management, Lakehead University, Thunder Bay, Ontario, Canada
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ETH Zurich · Institute of Integrative Biology
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Institute of Integrative Biology, ETH Zurich (Swiss Federal Institute of Technology) Universitätsstrasse 16 8092 Zurich Switzerland
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Czech Academy of Sciences, Global Change Research Institute · Institute of Forest Ecosystem Research
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Global Change Research Institute CAS Brno Czech Republic
IFER—Institute of Forest Ecosystem Research Jilove u Prahy Czech Republic
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Nicholas School of the Environment Duke University Durham North Carolina USA
Nicholas School of the Environment, Duke University, Durham, North Carolina, USA
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University of Missouri–St. Louis
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Department of Biology University of Missouri‐St Louis St Louis Missouri USA
Department of Biology, University of Missouri-St Louis, St Louis, Missouri, USA
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Universidade Estadual de Campinas (UNICAMP)
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Programa de Pós‐graduação em Biologia Vegetal Instituto de Biologia, Universidade Estadual de Campinas Campinas Brazil
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Department of Plant Sciences and Conservation Research Institute University of Cambridge Cambridge UK
Department of Plant Sciences and Conservation Research Institute, University of Cambridge, Cambridge, UK
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Andes to Amazon Biodiversity Program Madre de Dios Peru
Andes to Amazon Biodiversity Program, Madre de Dios, Peru
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Universidad Juárez del Estado de Durango
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Facultad de Ciencias Forestales Universidad Juárez del Estado de Durango Durango Mexico
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College of Saint Rose · West Virginia University
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Department of Biology West Virginia University Morgantown West Virginia USA
Department of Physical and Biological Sciences The College of Saint Rose Albany New York USA
Department of Biology, West Virginia University, Morgantown, West Virginia, USA
Department of Physical and Biological Sciences, The College of Saint Rose, Albany, New York, USA
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Department of Biology West Virginia University Morgantown West Virginia USA
Department of Biology, West Virginia University, Morgantown, West Virginia, USA
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Biology Department, Centre for Structural and Functional Genomics Concordia University Montreal Quebec Canada
Biology Department, Centre for Structural and Functional Genomics, Concordia University, Montreal, Quebec, Canada
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Universidade Regional de Blumenau
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Natural Science Department Universidade Regional de Blumenau Blumenau Brazil
Natural Science Department, Universidade Regional de Blumenau, Blumenau, Brazil
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World Agroforestry Centre · Wageningen University & Research
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Wageningen University and Research Wageningen The Netherlands
World Agroforestry (ICRAF) P.O. Box 30677 00100 Nairobi Kenya
Wageningen University and Research, Wageningen, The Netherlands
World Agroforestry (ICRAF), P.O. Box 30677 00100 Nairobi, Kenya
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Centre National de la Recherche Scientifique · Institut National de Recherche pour l'Agriculture, l'Alimentation et l'Environnement · Ecologie des Forêts de Guyane · Université des Antilles et de la Guyane
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Cirad, UMR EcoFoG (AgroParistech, CNRS, INRAE, Université des Antilles, Université de la Guyane), Campus Agronomique Kourou French Guiana
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University of Maryland, College Park
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Department of Geographical Sciences University of Maryland College Park Maryland USA
Department of Geographical Sciences, University of Maryland, College Park, Maryland, USA
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Institute of Forestry Belgrade Serbia
Institute of Forestry, Belgrade, Serbia
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Instituto Nacional de Pesquisas da Amazônia
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National Institute of Amazonian Research Manaus Brazil
National Institute of Amazonian Research, Manaus, Brazil
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Centre National de la Recherche Scientifique · Institut National de Recherche pour l'Agriculture, l'Alimentation et l'Environnement · Ecologie des Forêts de Guyane · Université des Antilles et de la Guyane
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Cirad, UMR EcoFoG (AgroParistech, CNRS, INRAE, Université des Antilles, Université de la Guyane), Campus Agronomique Kourou French Guiana
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IRET, Herbier National du Gabon (CENAREST) Libreville Gabon
IRET, Herbier National du Gabon (CENAREST), Libreville, Gabon
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University of Arizona · Queensland Department of Environment and Science
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Department of Ecology and Evolutionary Biology University of Arizona Tucson Arizona USA
Queensland Herbarium, Department of Environment and Science Toowong Queensland Australia
Department of Ecology and Evolutionary Biology, University of Arizona, Tucson, Arizona, USA
Queensland Herbarium, Department of Environment and Science, Toowong, Queensland, Australia
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Queensland Department of Environment and Science
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Queensland Herbarium, Department of Environment and Science Toowong Queensland Australia
Queensland Herbarium, Department of Environment and Science, Toowong, Queensland, Australia
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Université d'Agriculture de Kétou · Université Nationale d'Agriculture
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Ecole de Foresterie et Ingénierie du Bois, Université Nationale d'Agriculture Ketou Benin
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Universiti of Malaysia Sabah · Institute of Entomology · Czech Academy of Sciences, Biology Centre
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Biology Centre of the Czech Academy of Sciences, Institute of Entomology Ceske Budejovice, Czech Republic and Institute for Tropical Biology and Conservation, Universiti Malaysia Sabah Kota Kinabalu Sabah Malaysia
Biology Centre of the Czech Academy of Sciences, Institute of Entomology, Ceske Budejovice, Czech Republic and Institute for Tropical Biology and Conservation, Universiti Malaysia Sabah, Kota Kinabalu, Sabah, Malaysia
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Museu Paraense Emílio Goeldi. Coordenação de Ciências da Terra e Ecologia Belém Pará Brazil
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Geography, College of Life and Environmental Sciences University of Exeter Exeter UK
Geography, College of Life and Environmental Sciences, University of Exeter, Exeter, UK
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Natural Resources Institute Finland
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Natural Resources Institute Finland (Luke) Joensuu Finland
Natural Resources Institute Finland (Luke), Joensuu, Finland
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Affiliation as printed
Institute of Plant Sciences, University of Bern Bern Switzerland
Institute of Plant Sciences, University of Bern, Bern, Switzerland
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Forest Research Institute Malaysia
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Forest Research Institute Malaysia Kuala Lumpur Malaysia
Forest Research Institute Malaysia, Kuala Lumpur, Malaysia
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Affiliation as printed
Department of Sustainable Agro‐Ecosystems and Bioresources, Research and Innovation Center Fondazione Edmund Mach San Michele all'Adige Italy
Department of Sustainable Agro-Ecosystems and Bioresources, Research and Innovation Center, Fondazione Edmund Mach, San Michele all'Adige, Italy
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Food and Agriculture Organization of the United Nations
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Forestry Division, Food and Agriculture Organization of the United Nations Rome Italy
Forestry Division, Food and Agriculture Organization of the United Nations, Rome, Italy
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Affiliation as printed
Department of Sustainable Agro‐Ecosystems and Bioresources, Research and Innovation Center Fondazione Edmund Mach San Michele all'Adige Italy
Department of Sustainable Agro-Ecosystems and Bioresources, Research and Innovation Center, Fondazione Edmund Mach, San Michele all'Adige, Italy
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Affiliation as printed
School of Forestry and Environmental Studies Yale University New Haven Connecticut USA
School of Forestry and Environmental Studies, Yale University, New Haven, Connecticut, USA
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Royal Botanic Garden Edinburgh
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Royal Botanic Garden Edinburgh Edinburgh UK
Royal Botanic Garden Edinburgh, Edinburgh, UK
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Affiliation as printed
Department of Plant Sciences University of Oxford Oxford UK
Department of Plant Sciences, University of Oxford, Oxford, UK
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Affiliation as printed
Department of Plant Systematics University of Bayreuth Bayreuth Germany
Department of Plant Systematics, University of Bayreuth, Bayreuth, Germany
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Wageningen University & Research
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Wageningen University and Research Wageningen The Netherlands
Wageningen University and Research, Wageningen, The Netherlands
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Université de Montpellier · Forests and Societies · Institut National Polytechnique Félix Houphouët-Boigny
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Cirad, UPR Forêts et Sociétés University of Montpellier Montpellier France
Department of Forestry and Environment National Polytechnic Institute (INP‐HB) Yamoussoukro Côte d'Ivoire
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University of the Sunshine Coast
Affiliation as printed
Tropical Forests and People Research Centre, University of the Sunshine Coast Maroochydore Queensland Australia
Tropical Forests and People Research Centre, University of the Sunshine Coast, Maroochydore, Queensland, Australia
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Royal Society for the Protection of Birds · Wild Chimpanzee Foundation
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Centre for Conservation Science, The Royal Society for the Protection of Birds Sandy UK
Wild Chimpanzee Foundation, Liberia Office Monrovia Liberia
Centre for Conservation Science, The Royal Society for the Protection of Birds, Sandy, UK
Wild Chimpanzee Foundation, Liberia Office, Monrovia, Liberia
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Instituto de Investigaciones de la Amazonía Peruana
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Instituto de Investigaciones de la Amazonía Peruana Iquitos Peru
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African Institute for Mathematical Sciences · Stellenbosch University
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Department of Mathematical Sciences, Centre for Invasion Biology Stellenbosch University Stellenbosch South Africa
Theoretical Ecology Unit, African Institute for Mathematical Sciences Cape Town South Africa
Department of Mathematical Sciences, Centre for Invasion Biology, Stellenbosch University, Stellenbosch, South Africa
Theoretical Ecology Unit, African Institute for Mathematical Sciences, Cape Town, South Africa
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Korea Forestry Promotion Institute
Affiliation as printed
Division of Forest Resources Information Korea Forest Promotion Institute Seoul South Korea
Division of Forest Resources Information, Korea Forest Promotion Institute, Seoul, South Korea
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Affiliation as printed
Institut Agronomique néo‐Calédonien (IAC), Equipe Sol & Végétation (SolVeg) Nouméa New Caledonia
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Korea Forestry Promotion Institute
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Division of Forest Resources Information Korea Forest Promotion Institute Seoul South Korea
Division of Forest Resources Information, Korea Forest Promotion Institute, Seoul, South Korea
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Tokyo University of Agriculture
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Department of Forest Science Tokyo University of Agriculture Tokyo Japan
Department of Forest Science, Tokyo University of Agriculture, Tokyo, Japan
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Institute of Dendrology · Poznań University of Life Sciences · Polish Academy of Sciences
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Department of Game Management and Forest Protection, Poznań University of Life Sciences Poznań Poland
Institute of Dendrology, Polish Academy of Sciences Kórnik Poland
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Affiliation as printed
Faculty of Biology, Białowieża Geobotanical Station University of Warsaw Białowieża Poland
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Affiliation as printed
Department of Geosciences and Natural Resource Management University of Copenhagen Copenhagen Denmark
Department of Geosciences and Natural Resource Management, University of Copenhagen, Copenhagen, Denmark
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Universidade Estadual de Campinas (UNICAMP)
Affiliation as printed
Department of Plant Biology Institute of Biology, University of Campinas, UNICAMP Campinas Brazil
Department of Plant Biology, Institute of Biology, University of Campinas, UNICAMP, Campinas, Brazil
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Affiliation as printed
School of Biological Sciences University of Bristol Bristol UK
School of Biological Sciences, University of Bristol, Bristol, UK
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Bauman Moscow State Technical University
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Forestry Faculty, Bauman Moscow State Technical University Mytischi Russia
Forestry Faculty, Bauman Moscow State Technical University, Mytischi, Russia
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Field Museum of Natural History
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Field Museum of Natural History Chicago Illinois USA
Field Museum of Natural History, Chicago, Illinois, USA
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Affiliation as printed
CAVElab‐Computational and Applied Vegetation Ecology, Department of Environment Ghent University Ghent Belgium
CAVElab-Computational and Applied Vegetation Ecology, Department of Environment, Ghent University, Ghent, Belgium
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Smithsonian Tropical Research Institute · ForestGEO
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CTFS‐ForestGEO, Smithsonian Tropical Research Institute Balboa Panama
CTFS-ForestGEO, Smithsonian Tropical Research Institute, Balboa, Panama
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