Determining the drivers of non-native plant invasions is critical for managing native ecosystems and limiting the spread of invasive species1,2. Tree invasions in particular have been relatively overlooked, even though they have the potential to transform ecosystems and economies3,4. Here, leveraging global tree databases5-7, we explore how the phylogenetic and functional diversity of native tree communities, human pressure and the environment influence the establishment of non-native tree species and the subsequent invasion severity. We find that anthropogenic factors are key to predicting whether a location is invaded, but that invasion severity is underpinned by native diversity, with higher diversity predicting lower invasion severity. Temperature and precipitation emerge as strong predictors of invasion strategy, with non-native species invading successfully when they are similar to the native community in cold or dry extremes. Yet, despite the influence of these ecological forces in determining invasion strategy, we find evidence that these patterns can be obscured by human activity, with lower ecological signal in areas with higher proximity to shipping ports. Our global perspective of non-native tree invasion highlights that human drivers influence non-native tree presence, and that native phylogenetic and functional diversity have a critical role in the establishment and spread of subsequent invasions.

5 N Sukachev Institute of Forest FRC KSC Siberian Branch of the Russian Academy of Sciences Krasnoyarsk Russia

AgroParisTech UMR AMAP Cirad CNRS INRA IRD Université de Montpellier Montpellier France

AMAP University of Montpellier Montpellier France

Andes to Amazon Biodiversity Program Madre de Dios Peru

Bavarian State Institute of Forestry Freising Germany

Biodiversity and Natural Resources Program International Institute for Applied Systems Analysis Laxenburg Austria

Biology Centre of the Czech Academy of Sciences Institute of Entomology Ceske Budejovice Czech Republic

Biology Department Centre for Structural and Functional Genomics Concordia University Montreal Quebec Canada

Botanical Garden of Ural Branch of Russian Academy of Sciences Ural State Forest Engineering University Yekaterinburg Russia

CAVElab Computational and Applied Vegetation Ecology Department of Environment Ghent University Ghent Belgium

Center for Ecological Dynamics in a Novel Biosphere Department of Biology Aarhus University Aarhus Denmark

Center for Forest Ecology and Productivity Russian Academy of Sciences Moscow Russia

Center for Tropical Research Institute of the Environment and Sustainability UCLA Los Angeles CA USA

Centre for Agricultural Research in Suriname Paramaribo Suriname

Centre for Conservation Science The Royal Society for the Protection of Birds Sandy UK

Centre for Forest Research Université du Québec à Montréal Montreal Quebec Canada

Centre for Invasion Biology Department of Mathematical Sciences Stellenbosch University Stellenbosch South Africa

Centre for the Research and Technology of Agro Environmental and Biological Sciences CITAB University of Trás os Montes and Alto Douro UTAD Viseu Portugal

Centro Agricoltura Alimenti Ambiente University of Trento San Michele All'adige Italy

Centro de Ciências Biológicas e da Natureza Universidade Federal do Acre Rio Branco Acre Brazil

Centro de Modelación y Monitoreo de Ecosistemas Universidad Mayor Santiago Chile

Centro Multidisciplinar Universidade Federal do Acre Rio Branco Brazil

Chair for Forest Growth and Yield Science TUM School for Life Sciences Technical University of Munich Munich Germany

CIRAD CNRS INRAE IRD Montpellier France

Cirad UMR EcoFoG Université des Antilles Université de la Guyane Campus Agronomique Kourou France

Cirad UPR Forêts et Sociétés University of Montpellier Montpellier France

Climate Fire and Carbon Cycle Sciences USDA Forest Service Durham NC USA

Colegio de Profesionales Forestales de Cochabamba Cochabamba Bolivia

Compensation International S A Ci Progress GreenLife Bogotá Colombia

CTFS ForestGEO Smithsonian Tropical Research Institute Balboa Panama

Datascientist ch Wallisellen Switzerland

Departamento de Biología Universidad de la Serena La Serena Chile

Departamento de Ciências Biológicas Universidade do Estado de Mato Grosso Nova Xavantina Brazil

Departamento de Ecologia Universidade Federal do Rio Grande do Norte Natal Rio Grande do Norte Brazil

Departamento de Ecología y Recursos Naturales Facultad de Ciencias Universidad Nacional Autónoma de México Mexico City Mexico

Department of Agricultural Food Environmental and Animal Sciences University of Udine Udine Italy

Department of Agriculture Food Environment and Forest University of Firenze Florence Italy

Department of Agriculture Forestry and Bioresources Seoul National University Seoul South Korea

Department of Biological Geological and Environmental Sciences University of Bologna Bologna Italy

Department of Biological Sciences Boise State University Boise ID USA

Department of Biology Stanford University Stanford CA USA

Department of Biology University of Florence Florence Italy

Department of Biology University of Missouri St Louis St Louis MO USA

Department of Biology West Virginia University Morgantown WV USA

Department of Botany Dr Harisingh Gour Vishwavidyalaya Sagar India

Department of Botany Faculty of Science University of South Bohemia České Budějovice Czech Republic

Department of Crop and Forest Sciences University of Lleida Lleida Spain

Department of Ecology and Environmental Sciences Pondicherry University Puducherry India

Department of Ecology and Evolutionary Biology University of Arizona Tucson AZ USA

Department of Ecology and Evolutionary Biology University of Connecticut Storrs CT USA

Department of Ecology and Sustainable Agriculture Agricultural High School Polytechnic Institute of Viseu Viseu Portugal

Department of Environment and Development Studies United International University Dhaka Bangladesh

Department of Environment and Geography University of York York UK

Department of Environmental Sciences Central University of Jharkhand Ranchi Jharkhand India

Department of Evolutionary Anthropology Duke University Durham NC USA

Department of Evolutionary Biology and Environmental Studies University of Zürich Zurich Switzerland

Department of Forest and Wood Science University of Stellenbosch Stellenbosch South Africa

Department of Forest Engineering Universidade Regional de Blumenau Blumenau Brazil

Department of Forest Resources University of Minnesota St Paul MN USA

Department of Forest Science Tokyo University of Agriculture Tokyo Japan

Department of Forest Sciences Luiz de Queiroz College of Agriculture University of São Paulo Piracicaba Brazil

Department of Forestry and Environment National Polytechnic Institute Yamoussoukro Côte d'Ivoire

Department of Forestry and Natural Resources Purdue University West Lafayette IN USA

Department of Genetics Evolution and Environment University College London London UK

Department of Geographical Sciences University of Maryland College Park MD USA

Department of Geography University College London London UK

Department of Geomatics Forest Research Institute Raszyn Poland

Department of Geosciences and Natural Resource Management University of Copenhagen Copenhagen Denmark

Department of Physical and Biological Sciences The College of Saint Rose Albany NY USA

Department of Physical and Environmental Sciences Colorado Mesa University Grand Junction CO USA

Department of Plant Biology Institute of Biology University of Campinas UNICAMP Campinas Brazil

Department of Plant Sciences and Conservation Research Institute University of Cambridge Cambridge UK

Department of Plant Sciences University of Oxford Oxford UK

Department of Plant Systematics University of Bayreuth Bayreuth Germany

Department of Spatial Regulation GIS and Forest Policy Institute of Forestry Belgrade Serbia

Department of Wetland Ecology Institute for Geography and Geoecology Karlsruhe Institute for Technology Karlsruhe Germany

Department of Wildlife Management College of African Wildlife Management Mweka Tanzania

Department of Zoology University of Oxford Oxford UK

Depto de Silvicultura y Conservacion de la Naturaleza Universidad de Chile Temuco Chile

Division of Forest and Forest Resources Norwegian Institute of Bioeconomy Research Ås Norway

Division of Forest Resources Information Korea Forest Promotion Institute Seoul South Korea

Division of Forestry and Natural Resources West Virginia University Morgantown WV USA

Ecole de Foresterie et Ingénierie du Bois Université Nationale d'Agriculture Kétou Benin

Environmental and Life Sciences Faculty of Science Universiti Brunei Darussalam Bandar Seri Begawan Brunei

Environmental Change Institute School of Geography and the Environment University of Oxford Oxford UK

Environmental Studies and Research Center University of Campinas UNICAMP Campinas Brazil

European Commission Joint Research Center Ispra Italy

Facultad de Ciencias Forestales y Ambientales Universidad Juárez del Estado de Durango Durango Mexico

Faculty of Biology Białowieża Geobotanical Station University of Warsaw Białowieża Poland

Faculty of Environmental Sciences and Natural Resource Management Norwegian University of Life Sciences Ås Norway

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

Faculty of Forestry Qingdao Agricultural University Qingdao China

Faculty of Natural Resources Management Lakehead University Thunder Bay Ontario Canada

Faculty of Science and Technology Free University of Bolzano Bolzano Italy

Field Museum of Natural Histiory Chicago IL USA

Field Museum of Natural History Chicago IL USA

Flamingo Land Malton UK

Forest Ecology and Forest Management Group Wageningen University and Research Wageningen The Netherlands

Forest Ecology and Forest Management Wageningen University and Research Wageningen The Netherlands

Forest Research Institute Malaysia Kuala Lumpur Malaysia

Forest Research Institute University of the Sunshine Coast Sippy Downs Queensland Australia

Forestry Department Food and Agriculture Organization of the United Nations Rome Italy

Forestry Division Food and Agriculture Organization of the United Nations Rome Italy

Forestry Faculty Bauman Moscow State Technical University Mytischi Russia

Forestry School Tecnológico de Costa Rica TEC Cartago Costa Rica

Fundacion ConVida Universidad Nacional Abierta y a Distancia UNAD Medellin Colombia

Geobotany Faculty of Biology University of Freiburg Freiburg im Breisgau Germany

Geography College of Life and Environmental Sciences University of Exeter Exeter UK

German Centre for Integrative Biodiversity Research Halle Jena Leipzig Leipzig Germany

Glick Designs LLC Hadley MA USA

Global Change Research Institute CAS Brno Czech Republic

Graduate School of Agriculture Kyoto University Kyoto Japan

Guyana Forestry Commission Georgetown France

Hawkesbury Institute for the Environment Western Sydney University Penrith New South Wales Australia

IFER Institute of Forest Ecosystem Research Jilove u Prahy Czech Republic

Independent Researcher Bad Aussee Austria

Info Flora Geneva Switzerland

Institut Agronomique néo Calédonien Nouméa New Caledonia

Institute for Global Change Biology and School for Environment and Sustainability University of Michigan Ann Arbor MI USA

Institute for World Forestry University of Hamburg Hamburg Germany

Institute of Biology Geobotany and Botanical Garden Martin Luther University Halle Wittenberg Halle Wittenberg Germany

Institute of Botany The Czech Academy of Sciences Třeboň Czech Republic

Institute of Dendrology Polish Academy of Sciences Kórnik Poland

Institute of Forestry and Engineering Estonian University of Life Sciences Tartu Estonia

Institute of Forestry Belgrade Serbia

Institute of Integrative Biology ETH Zurich Zurich Switzerland

Institute of Plant Sciences University of Bern Bern Switzerland

Institute of Tropical Forest Conservation Mbarara University of Sciences and Technology Mbarara Uganda

Instituto de Investigaciones de la Amazonía Peruana Iquitos Peru

Instituto Nacional de Pesquisas da Amazônia Manaus Brazil

Instituto Nacional de Tecnología Agropecuaria Río Gallegos Argentina

Integrated Center for Research Development and Innovation in Advanced Materials Nanotechnologies and Distributed Systems for Fabrication and Control Stefan cel Mare University of Suceava Suceava Romania

Interdisciplinary Program in Agricultural and Forest Meteorology Seoul National University Seoul South Korea

IRET Herbier National du Gabon Libreville Gabon

Isotope Bioscience Laboratory ISOFYS Ghent University Ghent Belgium

Iwokrama International Centre for Rainforest Conservation and Development Georgetown Guyana

Jardín Botánico de Medellín Medellin Colombia

Jardín Botánico de Missouri Pasco Peru

Joint Research Unit CTFC AGROTECNIO CERCA Solsona Spain

Key Laboratory of Tropical Biological Resources Ministry of Education School of Life and Pharmaceutical Sciences Hainan University Haikou China

Laboratório de Dendrologia e Silvicultura Tropical Centro de Formação em Ciências Agroflorestais Universidade Federal do Sul da Bahia Itabuna Brazil

Laboratorio de geomática Instituto de Silvicultura e Industria de la Madera Universidad Juárez del Estado de Durango Durango Mexico

LINCGlobal Museo Nacional de Ciencias Naturales CSIC Madrid Spain

Manaaki Whenua Landcare Research Lincoln New Zealand

Museo de Historia Natural Noel kempff Mercado Santa Cruz Bolivia

Museu Paraense Emílio Goeldi Coordenação de Ciências da Terra e Ecologia Belém Pará Brazil

National Center for Agro Meteorology Seoul South Korea

National Forest Centre Forest Research Institute Zvolen Zvolen Slovakia

National Institute of Amazonian Research Manaus Brazil

Natural Science Department Universidade Regional de Blumenau Blumenau Brazil

Naturalis Biodiversity Center Leiden The Netherlands

Nicholas School of the Environment Duke University Durham NC USA

Plant Ecology and Nature Conservation Group Wageningen University Wageningen The Netherlands

Plant Systematic and Ecology Laboratory Department of Biology Higher Teachers' Training College University of Yaoundé 1 Yaoundé Cameroon

Polish State Forests Coordination Center for Environmental Projects Warsaw Poland

Pontificia Universidad Católica del Ecuador Quito Ecuador

Poznań University of Life Sciences Department of Game Management and Forest Protection Poznań Poland

Proceedings of the National Academy of Sciences Washington DC USA

Programa de doctorado en Ingeniería para el desarrollo rural y civil Escuela de Doctorado Internacional de la Universidad de Santiago de Compostela Santiago de Compostela Spain

Programa de Pós graduação em Biologia Vegetal Instituto de Biologia Universidade Estadual de Campinas Campinas Brazil

Quantitative Biodiversity Dynamics Betafaculty Utrecht University Utrecht The Netherlands

Queensland Herbarium Department of Environment and Science Toowong Queensland Australia

Research and Innovation Center Fondazione Edmund Mach San Michele All'adige Italy

Research Center of Forest Management Engineering of State Forestry and Grassland Administration Beijing Forestry University Beijing China

Research Institute for Agriculture and Life Sciences Seoul National University Seoul South Korea

Royal Botanic Garden Edinburgh Edinburgh UK

School of Biological and Behavioural Sciences Queen Mary University of London London UK

School of Biological Sciences University of Bristol Bristol UK

School of Forestry and Environmental Studies Yale University New Haven CT USA

School of Geography University of Leeds Leeds UK

School of Social Sciences Western Sydney University Penrith New South Wales Australia

Section for Ecoinformatics and Biodiversity Department of Biology Aarhus University Aarhus Denmark

Servicios Ecosistémicos y Cambio Climático Fundación Con Vida and Corporación COL TREE Medellín Colombia

Siberian Federal University Krasnoyarsk Russian Federation Krasnoyarsk Russia

Silviculture and Forest Ecology of the Temperate Zones University of Göttingen Göttingen Germany

Silviculture Research Institute Vietnamese Academy of Forest Sciences Hanoi Vietnam

Spatial Ecology and Conservation Laboratory Department of Tourism Recreation and Sport Management University of Florida Gainesville FL USA

Spatial Ecology and Conservation Laboratory School of Forest Resources and Conservation University of Florida Gainesville FL USA

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

TERRA Teach and Research Centre Gembloux Agro Bio Tech University of Liege Liege Belgium

The Forest School at The Yale School of the Environment Yale University New Haven CT USA

The Santa Fe Institute Santa Fe NM USA

Theoretical Ecology Unit African Institute for Mathematical Sciences Cape Town South Africa

Tropenbos International Wageningen The Netherlands

Tropical Biodiversity MUSE Museo delle Scienze Trento Italy

Tropical Forests and People Research Centre University of the Sunshine Coast Maroochydore Queensland Australia

UFR Biosciences University Félix Houphouët Boigny Abidjan Côte d'Ivoire

UNELLEZ Guanare Programa de Ciencias del Agro y el Mar Herbario Universitario Portuguesa Venezuela

UniSA STEM and Future Industries Institute University of South Australia Adelaide South Australia Australia

United Nation Framework Convention on Climate Change Bonn Germany

Universidad del Tolima Ibagué Colombia

Universidad Estatal Amazónica Puyo Pastaza Ecuador

Universidad Nacional de la Amazonía Peruana Iquitos Peru

Universidad Nacional de San Antonio Abad del Cusco Cusco Peru

Université de Lorraine AgroParisTech INRAE Silva Nancy France

Vicerrectoria de Investigacion y Postgrado Universidad de La Frontera Temuco Chile

Wageningen University and Research Wageningen The Netherlands

Warsaw University of Life Sciences Warsaw Poland

Wild Chimpanzee Foundation Liberia Office Monrovia Liberia

Wildlife Conservation Society Vientiane Laos

World Agroforestry Nairobi Kenya

Erratum v

Nature. 2023 Oct;622(7982):E2. doi: 10.1038/s41586-023-06654-9. PubMed

Zobrazit více v PubMed

Pyšek P, et al. Scientists’ warning on invasive alien species. Biol. Rev. 2020;95:1511–1534. PubMed PMC

Pejchar L, Mooney HA. Invasive species, ecosystem services and human well-being. Trends Ecol. Evol. 2009;24:497–504. PubMed

Richardson DM, Hui C, Nuñez MA, Pauchard A. Tree invasions: patterns, processes, challenges and opportunities. Biol. Invasions. 2014;16:473–481.

Richardson DM, Rejmánek M. Trees and shrubs as invasive alien species–a global review. Divers. Distrib. 2011;17:788–809.

Plants of the World Online. Royal Botanic Gardens Kewhttp://www.plantsoftheworldonline.org/ (2022).

van Kleunen M, et al. The Global Naturalized Alien Flora (GloNAF) database. Ecology. 2019;100:e02542. PubMed

Liang J, et al. Positive biodiversity-productivity relationship predominant in global forests. Science. 2016;354:aaf8957. PubMed

Vitousek PM, D'Antonio CM, Loope LL, Rejmanek M, Westbrooks R. Introduced species: a significant component of human-caused global change. N. Z. J. Ecol. 1997;21:1–16.

Butchart SH, et al. Global biodiversity: indicators of recent declines. Science. 2010;328:1164–1168. PubMed

Early R, et al. Global threats from invasive alien species in the twenty-first century and national response capacities. Nat. Commun. 2016;7:12485. PubMed PMC

Lockwood, J. L., Hoopes, M. F. & Marchetti, M. P. Invasion Ecology 2nd edn (John Wiley & Sons, 2013).

DiTomaso JM. Invasive weeds in rangelands: species, impacts, and management. Weed Sci. 2000;48:255–265.

Simberloff D. We can eliminate invasions or live with them. Successful management projects. Biol. Invasions. 2008;11:149–157.

Mack RN, et al. Biotic invasions: causes, epidemiology, global consequences, and control. Ecol. Appl. 2000;10:689–710.

Boyd IL, Freer-Smith PH, Gilligan CA, Godfray HC. The consequence of tree pests and diseases for ecosystem services. Science. 2013;342:1235773. PubMed

Vitousek PM, D’Antonio CM, Loope LL, Westbrooks R. Biological invasions as global environmental change. Am. Sci. 1996;84:468–478.

Diagne C, et al. High and rising economic costs of biological invasions worldwide. Nature. 2021;592:571–576. PubMed

Nunez MA, et al. Exotic mammals disperse exotic fungi that promote invasion by exotic trees. PLoS ONE. 2013;8:e66832. PubMed PMC

Richardson DM, Pyšek P. Naturalization of introduced plants: ecological drivers of biogeographical patterns. New Phytol. 2012;196:383–396. PubMed

Richardson DM, Pyšek P. Plant invasions: merging the concepts of species invasiveness and community invasibility. Prog. Phys. Geog. 2006;30:409–431.

Seabloom EW, et al. Human impacts, plant invasion, and imperiled plant species in California. Ecol. Appl. 2006;16:1338–1350. PubMed

Dayton, P. K. in Proc. Colloquium on Conservation Problems in Antarctica (ed. Parker, B. C.) 81–96 (Allen Press, 1972).

Beaury EM, Finn JT, Corbin JD, Barr V, Bradley BA. Biotic resistance to invasion is ubiquitous across ecosystems of the United States. Ecol. Lett. 2020;23:476–482. PubMed

Cadotte MW, Campbell SE, Li S-P, Sodhi DS, Mandrak NE. Preadaptation and naturalization of nonnative species: Darwin’s two fundamental insights into species invasion. Annu. Rev. Plant Biol. 2018;69:661–684. PubMed

Fridley J, et al. The invasion paradox: reconciling pattern and process in species invasions. Ecology. 2007;88:3–17. PubMed

Funk JL. The physiology of invasive plants in low-resource environments. Conserv. Physiol. 2013;1:cot026. PubMed PMC

Nuñez MA, Dickie IA. Invasive belowground mutualists of woody plants. Biol. Invasions. 2014;16:645–661.

van Kleunen M, Weber E, Fischer M. A meta-analysis of trait differences between invasive and non-invasive plant species. Ecol. Lett. 2010;13:235–245. PubMed

Pyšek, P. & Richardson, D. M. in Biological Invasions (ed. Nentwig, W.) 97–125 (Springer, 2007).

Dickie IA, et al. The emerging science of linked plant–fungal invasions. New Phytol. 2017;215:1314–1332. PubMed

Diez JM, Sullivan JJ, Hulme PE, Edwards G, Duncan RP. Darwin’s naturalization conundrum: dissecting taxonomic patterns of species invasions. Ecol. Lett. 2008;11:674–681. PubMed

Ma C, et al. Different effects of invader–native phylogenetic relatedness on invasion success and impact: a meta-analysis of Darwin’s naturalization hypothesis. Proc. R. Soc. B. 2016;283:20160663. PubMed PMC

Catford JA, Jansson R, Nilsson C. Reducing redundancy in invasion ecology by integrating hypotheses into a single theoretical framework. Divers. Distrib. 2009;15:22–40.

Levine JM, D'Antonio CM. Elton revisited: a review of evidence linking diversity and invasibility. Oikos. 1999;87:15–26.

Tilman D. Community invasibility, recruitment limitation, and grassland biodiversity. Ecology. 1997;78:81–92.

Fine PV. The invasibility of tropical forests by exotic plants. J. Trop. Ecol. 2002;18:687–705.

Elton, C. S. The Ecology of Invasion by Animals and Plants 1st edn (Methuen, 1958).

Kennedy TA, et al. Biodiversity as a barrier to ecological invasion. Nature. 2002;417:636–638. PubMed

Mungi NA, Qureshi Q, Jhala YV. Role of species richness and human impacts in resisting invasive species in tropical forests. J. Ecol. 2021;109:3308–3321.

Stohlgren TJ, Jarnevich C, Chong GW, Evangelista PH. Scale and plant invasions: a theory of biotic acceptance. Preslia. 2006;78:405–426.

Zeiter M, Stampfli A. Positive diversity–invasibility relationship in species-rich semi-natural grassland at the neighbourhood scale. Ann. Bot. 2012;110:1385–1393. PubMed PMC

Zanne AE, et al. Functional biogeography of angiosperms: life at the extremes. New Phytol. 2018;218:1697–1709. PubMed

Kraft NJ, et al. Community assembly, coexistence and the environmental filtering metaphor. Funct. Ecol. 2015;29:592–599.

Keddy PA. Assembly and response rules: two goals for predictive community ecology. J. Veg. Sci. 1992;3:157–164.

Gallien L, Carboni M, Münkemüller T. Identifying the signal of environmental filtering and competition in invasion patterns–a contest of approaches from community ecology. Methods Ecol. Evol. 2014;5:1002–1011.

Divíšek J, et al. Similarity of introduced plant species to native ones facilitates naturalization, but differences enhance invasion success. Nat. Commun. 2018;9:4631. PubMed PMC

MacArthur R, Levins R. The limiting similarity, convergence, and divergence of coexisting species. Am. Nat. 1967;101:377–385.

Park DS, Potter D. A test of Darwin’s naturalization hypothesis in the thistle tribe shows that close relatives make bad neighbors. Proc. Natl Acad. Sci. USA. 2013;110:17915–17920. PubMed PMC

Schaefer H, Hardy OJ, Silva L, Barraclough TG, Savolainen V. Testing Darwin’s naturalization hypothesis in the Azores. Ecol. Lett. 2011;14:389–396. PubMed

Darwin, C. On the Origin of Species by Means of Natural Selection, or Preservation of Favoured Races in the Struggle for Life (John Murray, 1859). PubMed PMC

Hubbell, S. P. The Unified Neutral Theory of Biodiversity and Biogeography (Princeton Univ. Press, 2001). PubMed

Pinto‐Ledezma JN, et al. Testing Darwin’s naturalization conundrum based on taxonomic, phylogenetic, and functional dimensions of vascular plants. Ecol. Monogr. 2020;90:e01420.

Theoharides KA, Dukes JS. Plant invasion across space and time: factors affecting nonindigenous species success during four stages of invasion. New Phytol. 2007;176:256–273. PubMed

Hulme PE. Trade, transport and trouble: managing invasive species pathways in an era of globalization. J. Appl. Ecol. 2009;46:10–18.

Simberloff D, Von Holle B. Postitive interactions of nonindigenous species: invasion meltdown? Biol. Invasions. 1999;1:21–23.

Pyšek P, Křivánek M, Jarošík V. Planting intensity, residence time, and species traits determine invasion success of alien woody species. Ecology. 2009;90:2734–2744. PubMed

Rejmánek M. Invasive trees and shrubs: where do they come from and what we should expect in the future? Biol. Invasions. 2014;16:483–498.

Chong KY, et al. Are terrestrial biological invasions different in the tropics? Annu. Rev. Ecol. Evol. Syst. 2021;52:291–314.

Smith JL, Perino JV. Osage orange (Maclura pomifera): history and economic uses. Econ. Bot. 1981;35:24–41.

Pyšek P, et al. The global invasion success of Central European plants is related to distribution characteristics in their native range and species traits. Divers. Distrib. 2009;15:891–903.

Svenning JC, et al. The influence of interspecific interactions on species range expansion rates. Ecography. 2014;37:1198–1209. PubMed PMC

Amarasekare P, Simon MW. Latitudinal directionality in ectotherm invasion success. Proc. R. Soc. B. 2020;287:20191411. PubMed PMC

Park DS, Feng X, Maitner BS, Ernst KC, Enquist BJ. Darwin’s naturalization conundrum can be explained by spatial scale. Proc. Natl Acad. Sci. USA. 2020;117:10904–10910. PubMed PMC

Tournadre J. Anthropogenic pressure on the open ocean: The growth of ship traffic revealed by altimeter data analysis. Geophys. Res. Lett. 2014;41:7924–7932.

de Lima RAF, et al. Making forest data fair and open. Nat. Ecol. Evol. 2022;6:656–658. PubMed

Moles AT, et al. Invasions: the trail behind, the path ahead, and a test of a disturbing idea. J. Ecol. 2012;100:116–127.

Peng S, Kinlock NL, Gurevitch J, Peng S. Correlation of native and exotic species richness: a global meta‐analysis finds no invasion paradox across scales. Ecology. 2019;100:e02552. PubMed PMC

Dickie IA, et al. Conflicting values: ecosystem services and invasive tree management. Biol. Invasions. 2014;16:705–719.

Finnoff D, Shogren JF, Leung B, Lodge D. Take a risk: preferring prevention over control of biological invaders. Ecol. Econ. 2007;62:216–222.

Bivand, R. & Piras, G. Comparing implementations of estimation methods for spatial econometrics. J. Stat. Softw. 63, 1–36 (2015).

Kalwij JM. Review of ‘The Plant List, a working list of all plant species’. J. Veg. Sci. 2012;23:998–1002.

Gorelick N, et al. Google Earth Engine: planetary-scale geospatial analysis for everyone. Remote Sens. Environ. 2017;202:18–27.

Beech E, Rivers M, Oldfield S, Smith PP. GlobalTreeSearch: the first complete global database of tree species and country distributions. J. Sustain. For. 2017;36:454–489.

Catford JA, Vesk PA, Richardson DM, Pyšek P. Quantifying levels of biological invasion: towards the objective classification of invaded and invasible ecosystems. Glob. Change Biol. 2012;18:44–62.

Guo Q, et al. A unified approach for quantifying invasibility and degree of invasion. Ecology. 2015;96:2613–2621. PubMed

Van Den Hoogen J, et al. Soil nematode abundance and functional group composition at a global scale. Nature. 2019;572:194–198. PubMed

Bastin J-F, et al. The global tree restoration potential. Science. 2019;365:76–79. PubMed

Karger DN, et al. Climatologies at high resolution for the earth’s land surface areas. Sci. Data. 2017;4:170122. PubMed PMC

Stewart S, et al. Climate extreme variables generated using monthly time‐series data improve predicted distributions of plant species. Ecography. 2021;44:626–639.

Hengl T, et al. SoilGrids250m: Global gridded soil information based on machine learning. PLoS ONE. 2017;12:e0169748. PubMed PMC

Global Ports. All Layers and Tables (GLOBAL/GlobalPorts)https://gis.wfp.org/arcgis/rest/services/GLOBAL/GlobalPorts/MapServer/layers (2021).

Global Airports. The World Bank Data Cataloghttps://datacatalog.worldbank.org/search/dataset/0038117 (2021).

Maynard DS, et al. Global relationships in tree functional traits. Nat. Commun. 2022;13:3185. PubMed PMC

Joswig JS, et al. Climatic and soil factors explain the two-dimensional spectrum of global plant trait variation. Nat. Ecol. Evol. 2022;6:36–50. PubMed PMC

Florczyk, A. J. et al. GHSL Data Package 2019: Public Release GHS P2019 (European Commission, Joint Research Centre, 2019).

Owen NR, Gumbs R, Gray CL, Faith DP. Global conservation of phylogenetic diversity captures more than just functional diversity. Nat. Commun. 2019;10:859. PubMed PMC

Mazel F, et al. Prioritizing phylogenetic diversity captures functional diversity unreliably. Nat. Commun. 2018;9:2888. PubMed PMC

Tucker CM, et al. A guide to phylogenetic metrics for conservation, community ecology and macroecology. Biol. Rev. 2017;92:698–715. PubMed PMC

Jin Y, Qian HV. PhyloMaker: an R package that can generate very large phylogenies for vascular plants. Ecography. 2019;42:1353–1359.

Strauss SY, Webb CO, Salamin N. Exotic taxa less related to native species are more invasive. Proc. Natl Acad. Sci. USA. 2006;103:5841–5845. PubMed PMC

Cazzolla Gatti R, et al. The number of tree species on Earth. Proc. Natl Acad. Sci. USA. 2022;119:e2115329119. PubMed PMC

Pyšek, P. & Richardson, D. M. in Biological Invasions (ed. Nentwig, W.) 97–125 (Springer, 2008).

Petchey OL, Gaston KJ. Functional diversity (FD), species richness and community composition. Ecol. Lett. 2002;5:402–411.

Paradis E, Schliep K. ape 5.0: an environment for modern phylogenetics and evolutionary analyses in R. Bioinformatics. 2019;35:526–528. PubMed

Wickham H, et al. Welcome to the Tidyverse. J. Open Source Softw. 2019;4:1686.

Bittinger, K. Abdiv: Alpha and beta diversity measures. R package version 0.2.0 https://cran.r-project.org/web/packages/abdiv/index.html (2020).

Calaway, R., Analytics, R., Weston, S., Tenenbaum, D. & Calaway, M. doParallel: Foreach parallel adaptor for the ‘parallel’ package. R package version 1.0.17 https://cran.r-project.org/web/packages/doParallel/index.html (2015).

Microsoft & Weston, S. foreach: Provides foreach looping construct. R package version 1.5.1. https://CRAN.R-project.org/package=foreach (2020).

Pearse WD, et al. Pez: phylogenetics for the environmental sciences. Bioinformatics. 2015;31:2888–2890. PubMed

Breiman L. Bagging predictors. Mach. Learn. 1996;24:123–140.

Crase B, Liedloff AC, Wintle BA. A new method for dealing with residual spatial autocorrelation in species distribution models. Ecography. 2012;35:879–888.

Portier J, Gauthier S, Robitaille A, Bergeron Y. Accounting for spatial autocorrelation improves the estimation of climate, physical environment and vegetation’s effects on boreal forest’s burn rates. Landsc. Ecol. 2018;33:19–34. PubMed PMC

Bjornstad, O. N. ncf: Spatial covariance functions. R package version 1.3-2 https://cran.r-project.org/web/packages/ncf/index.html (2022).

Bivand, R. S., Pebesma, E. J., Gómez-Rubio, V. & Pebesma, E. J. Applied spatial data analysis with R (Springer, 2008).

Lundberg SM, et al. From local explanations to global understanding with explainable AI for trees. Nat. Mach. Intell. 2020;2:56–67. PubMed PMC

Lundberg, S. M. & Lee, S.-I. in Advances in Neural Information Processing Systems 30 (eds Guyon, I. et al.) (NIPS, 2017).

Wright, M. N., Wager, S. & Probst, P. Ranger: A fast implementation of random forests. R package version 0.12 (2020).

Greenwell, B. fastshap: Fast approximate Shapley values. R package version 0.0.7 https://github.com/bgreenwell/fastshap (2020).

Roberts DR, et al. Cross‐validation strategies for data with temporal, spatial, hierarchical, or phylogenetic structure. Ecography. 2017;40:913–929.

Li J. Assessing the accuracy of predictive models for numerical data: not r nor r2, why not? Then what? PLoS ONE. 2017;12:e0183250. PubMed PMC

Platt, J. in Advances in Large Margin Classifiers, Vol. 10 (eds Smola, A. J. et al.) 61–74 (MIT Press, 1999).

Niculescu-Mizil, A. & Caruana, R. in Proc. 22nd International Conference on Machine Learning 625–632 (Association for Computing Machinery, 2005).

Meyer H, Pebesma E. Predicting into unknown space? Estimating the area of applicability of spatial prediction models. Methods Ecol. Evol. 2021;12:1620–1633.

FAO. Global Forest Resources Assessment 2020: Main Report (Food and Agriculture Organization of the United Nations, 2020).

R Core Team. R: A Language and Environment for Statistical Computing. http://www.R-project.org/ (R Foundation for Statistical Computing, 2019).

Bates, D., Maechler, M., Bolker, B. & Walker, S. Fitting linear mixed-effects models using lme4. J. Stat. Softw.10.18637/jss.v067.i01 (2015).

Kunzetsova, A., Brockhoff, P. & Christensen, R. lmerTest package: tests in linear mixed effect models. J. Stat. Softw.10.18637/jss.v082.i13 (2017).

Zeileis, A. et al. betareg: Beta regression. R package version 3.1-4 https://cran.r-project.org/web/packages/betareg/index.html (2021).

Wickham, H., Chang, W. & Wickham, M. H. ggplot2: Create elegant data visualisations using the grammar of graphics. R package version 2 https://search.r-project.org/CRAN/refmans/ggplot2/html/ggplot2-package.html (2016).

Stage dependence of Elton's biotic resistance hypothesis of biological invasions