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Global functional shifts in trees driven by alien naturalization and native extinction

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Abstract

Human activities are driving simultaneous native extinctions and alien naturalizations, reshaping global tree diversity with major implications for ecosystem structure and function. Here we analysed functional traits and environmental niches of 31,001 tree species worldwide, comparing naturalized, threatened and non-threatened species to assess current patterns and project future shifts under intensified extinction and naturalization. Future tree-rich ecosystems are projected to become increasingly dominated by fast-growing, high-resource-use species with acquisitive traits, while slow-growing, conservative species face greater extinction risk. Although group means along the main functional axes do not differ significantly, naturalized species occupy broader functional and environmental spaces and thrive in colder and more variable climates, whereas threatened species are more specialized to warm, stable and nutrient-rich environments, with non-threatened species intermediate. Projected naturalizations expand local functional diversity, but their acquisitive strategies could reduce long-term ecosystem stability, while extinctions cause pronounced contractions of functional and environmental trait space, especially in climatically variable regions. Overall, our findings reveal an accelerating global shift towards faster-growing tree communities, with likely consequences for carbon storage and biodiversity, underscoring the need to safeguard slow-growing species and limit the dominance of acquisitive trees.

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Fig. 1: Functional and environmental spaces of tree species.
Fig. 2: Shifts of the occupied functional and environmental PC spaces between current and future projection scenarios, represented as the occupied space in the 0.99 quantile distribution (representing the existing trait and environmental boundaries at the global scale) for each of the three tree groups.
Fig. 3: Shifts in functional and environmental spectra.

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Data availability

The data that support the findings of this study are available via GitHub at https://github.com/kun-ecology/global_tree.fun-env.space and are mirrored on Zenodo at https://doi.org/10.5281/zenodo.17662284 (ref. 104). The trait data were obtained from ref. 42, and the environmental data were extracted from ref. 43.

Code availability

R scripts for reproducing the analyses and figures are available via GitHub at https://github.com/kun-ecology/global_tree.fun-env.space and are mirrored on Zenodo at https://doi.org/10.5281/zenodo.17662284 (ref. 104).

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Acknowledgements

This project was supported by the Natural Science Foundation of China (32171588 and 32471676), the Innovation Program of the Shanghai Municipal Education Commission (2023ZKZD36), the Fundamental and Interdisciplinary Disciplines Breakthrough Plan of the Ministry of Education of China (JYB2025XDXM904). We also consider this work a contribution to the Center for Ecological Dynamics in a Novel Biosphere (ECONOVO), funded by the Danish National Research Foundation (grant no. DNRF173 to J.-C.S.). J.M.S.-D. is supported by the programme RYC2022-035668-I, funded by MCIU/AEI/10.13039/501100011033 and FSE+. J. Pisek is supported by the Estonian Research Council (PRG1405) and by the Estonian Ministry of Education and Research, Centre of Excellence for Sustainable Land Use (TK232). C.B. was supported by the National Research Foundation of Korea grant funded by the Korea government (MSIT) (2022R1A2C1003504). A.G.G. was supported by the ANID PIA/BASAL (FB210006) and FONDECYT (1240874). Ü.N. is supported by Estonian Ministry of Education and Research, Centre of Excellence AgroCropFuture (TK200).

Author information

Authors and Affiliations

  1. Zhejiang Tiantong Forest Ecosystem National Observation and Research Station, Institute of Eco-Chongming & Research Center for Global Change and Ecological Forecasting, School of Ecological and Environmental Sciences, East China Normal University, Shanghai, P. R. China

    Wen-Yong Guo

  2. Zhejiang Zhoushan Island Ecosystem Observation and Research Station, Zhoushan, P. R. China

    Wen-Yong Guo

  3. State Key Laboratory of Estuarine and Coastal Research, East China Normal University, Shanghai, P. R. China

    Wen-Yong Guo

  4. Botanical Institute of Barcelona (CSIC-CMCNB), Barcelona, Spain

    Josep M. Serra-Diaz

  5. Northeast Institute of Geography and Agroecology, Chinese Academy Sciences, Changchun, P. R. China

    Kun Guo

  6. Plant Ecology and Nature Conservation Group, Wageningen University, Wageningen, the Netherlands

    Coline C. F. Boonman & Fons van der Plas

  7. School of Geography, University of Nottingham, Nottingham, UK

    Franziska Schrodt

  8. Department of Integrative Biology, University of South Florida, St. Petersburg, FL, USA

    Brian S. Maitner

  9. Department of Ecology and Evolution and Eversource Energy Center, University of Connecticut, Storrs, CT, USA

    Cory Merow

  10. CEFE, Univ Montpellier, CNRS, EPHE, IRD, Montpellier, France

    Cyrille Violle

  11. School of Environmental Sciences, University of Guelph, Guelph, Ontario, Canada

    Madhur Anand

  12. Department of Biology, University of Copenhagen, Copenhagen, Denmark

    Hans Henrik K. Bruun

  13. Department of Biological Sciences, Gyeongkuk National University, Andong, Republic of Korea

    Chaeho Byun

  14. Department of Geography, King’s College London, London, UK

    Jane A. Catford

  15. Fenner School of Environment & Society, Australian National University, Canberra, Australian Capital Territory, Australia

    Jane A. Catford

  16. Department of Biotechnology and Life Sciences, University of Insubria, Varese, Italy

    Bruno E. L. Cerabolini

  17. Herbario Luis Fournier Origgi, Centro de Investigación en Biodiversidad y Ecología Tropical, Universidad de Costa Rica, San José, Costa Rica

    Eduardo Chacón-Madrigal

  18. Herbario Nacional, Departamento de Historia Natural, Museo Nacional de Costa Rica, San José, Costa Rica

    Eduardo Chacón-Madrigal

  19. Department of Biology, University of Pisa, Pisa, Italy

    Daniela Ciccarelli

  20. Department of Ecology—Evolutionary Biology, Faculty of Biology—Biotechnology, VNUHCM—University of Science, Ho Chi Minh City, Vietnam

    Anh Tuan Dang-Le

  21. Vietnam National University, Ho Chi Minh City, Vietnam

    Anh Tuan Dang-Le

  22. Center for Biodiversity Conservation and Endangered Species, Ho Chi Minh City, Vietnam

    Anh Tuan Dang-Le

  23. FSC—International Center—Forest Stewardship Council, Bonn, Germany

    Arildo S. Dias

  24. Embrapa Maranhão, Brazilian Agricultural Research Corporation (Embrapa), São Luís, Brazil

    Aelton B. Giroldo

  25. Departamento de Ciencias Ambientales y Recursos Naturales Renovables, Facultad de Ciencias Agronómicas, Universidad de Chile, Santiago, Chile

    Alvaro G. Gutiérrez

  26. Instituto de Ecología y Biodiversidad, Santiago, Chile

    Alvaro G. Gutiérrez

  27. Institute of Botany, Ulm University, Ulm, Germany

    Steven Jansen

  28. German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig, Leipzig, Germany

    Jens Kattge

  29. Max Planck Institute for Biogeochemistry, Jena, Germany

    Jens Kattge

  30. Trees and Forest Genetic Resources and Biodiversity, World Agroforestry, CIFOR-ICRAF, Nairobi, Kenya

    Roeland Kindt

  31. Department of Plant & Environmental Sciences, Weizmann Institute of Science, Rehovot, Israel

    Tamir Klein

  32. Forest Ecology and Management Group, Wageningen University, Wageningen, the Netherlands

    Koen Kramer

  33. Land Life Company, Amsterdam, the Netherlands

    Koen Kramer

  34. Environmental Research Institute, University of Waikato, Hamilton, New Zealand

    Christopher H. Lusk

  35. Department of Physical and Environmental Sciences, University of Toronto Scarborough, Toronto, Ontario, Canada

    Adam R. Martin

  36. Department of Botany and Biodiversity Research Centre, University of British Columbia, Vancouver, British Columbia, Canada

    Sean T. Michaletz

  37. Department of Biology, Vrije Universiteit Brussel, Brussels, Belgium

    Vanessa Minden

  38. Research Center for Advanced Science and Technology, University of Tokyo, Tokyo, Japan

    Akira S. Mori

  39. Estonian University of Life Sciences, Tartu, Estonia

    Ülo Niinemets

  40. Division of Forest and Biomaterials Science, Graduate School of Agriculture, Kyoto University, Kyoto, Japan

    Yusuke Onoda

  41. CREAF, Barcelona, Spain

    Josep Peñuelas

  42. CSIC, Global Ecology Unit CREAF-CSIC-UAB, Barcelona, Spain

    Josep Peñuelas

  43. Tartu Observatory, University of Tartu, Tõravere, Estonia

    Jan Pisek

  44. Department of Ecology, Radboud Institute for Biological and Environmental Sciences, Faculty of Science, Radboud University, Nijmegen, the Netherlands

    Bjorn J. M. Robroek

  45. Department of Biology, Algoma University, Sault Ste. Marie, Ontario, Canada

    Brandon Schamp

  46. Centre for Environmental Sciences (CMK), Hasselt University, Hasselt, Belgium

    Nadejda A. Soudzilovskaia

  47. Canadian Forest Service, Natural Resources Canada, Quebec City, Quebec, Canada

    Nelson Thiffault

  48. Department of Ecology and Evolutionary Biology, University of Arizona, Tucson, AZ, USA

    Brian J. Enquist

  49. Santa Fe Institute, Santa Fe, NM, USA

    Brian J. Enquist

  50. Center for Ecological Dynamics in a Novel Biosphere (ECONOVO), Department of Biology, Aarhus University, Aarhus, Denmark

    Jens-Christian Svenning

Authors
  1. Wen-Yong Guo
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  2. Josep M. Serra-Diaz
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  40. Jens-Christian Svenning
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Contributions

W.-Y.G. conceived the project. J.M.S.-D., W.-Y.G., C.C.F.B., R.K. and all others collected the data. K.G. and W.-Y.G. analysed the data. W.-Y.G., K.G. and J.-C.S. interpreted the data and wrote the manuscript. All authors contributed data, discussed the results, revised the manuscript drafts, contributed to writing and approved the final manuscript.

Corresponding author

Correspondence to Wen-Yong Guo.

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The authors declare no competing interests.

Peer review

Peer review information

Nature Plants thanks Wei Huang, Ana Novoa and the other, anonymous, reviewer(s) for their contribution to the peer review of this work.

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Supplementary information

Supplementary Information

Supplementary Tables 1–5 and Figs. 1–8.

Reporting Summary

Supplementary Table 6

Potential naturalized tree species obtained from DAPC.

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Guo, WY., Serra-Diaz, J.M., Guo, K. et al. Global functional shifts in trees driven by alien naturalization and native extinction. Nat. Plants (2026). https://doi.org/10.1038/s41477-025-02207-2

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