Survival rates of large trees determine forest biomass dynamics. Survival rates of small trees have been linked to mechanisms that maintain biodiversity across tropical forests. How species survival rates change with size offers insight into the links between biodiversity and ecosystem function across tropical forests. We tested patterns of size-dependent tree survival across the tropics using data from 1,781 species and over 2 million individuals to assess whether tropical forests can be characterized by size-dependent life-history survival strategies. We found that species were classifiable into four ‘survival modes’ that explain life-history variation that shapes carbon cycling and the relative abundance within forests. Frequently collected functional traits, such as wood density, leaf mass per area and seed mass, were not generally predictive of the survival modes of species. Mean annual temperature and cumulative water deficit predicted the proportion of biomass of survival modes, indicating important links between evolutionary strategies, climate and carbon cycling. The application of survival modes in demographic simulations predicted biomass change across forest sites. Our results reveal globally identifiable size-dependent survival strategies that differ across diverse systems in a consistent way. The abundance of survival modes and interaction with climate ultimately determine forest structure, carbon storage in biomass and future forest trajectories.

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The authors thank the many people involved in establishing and maintaining all the plots utilized in these analyses. A detailed list of funding sources for each plot is available in the Supplementary Information. The development of this project benefited from ForestGEO workshops in 2015, 2016 and 2017 (NSF DEB-1046113 to S.J.D.). Contributions by C.X., J.Q.C., S.J.D. and N.M. were supported by the Next-Generation Ecosystem Experiments (NGEE-Tropics) project, funded by the US Department of Energy, Office of Biological and Environmental Research. S.M.M. was partially funded by NSF - EF1137366. D.J.J. was supported by Los Alamos National Laboratory (Director’s Post-doctoral Fellowship).

Author information


  1. Earth and Environmental Science, Los Alamos National Laboratory, Los Alamos, NM, USA

    • Daniel J. Johnson
    •  & Chonggang Xu
  2. School of Forest Resources and Conservation, Gainesville, FL, USA

    • Daniel J. Johnson
  3. Forest Global Earth Observatory, Smithsonian Environmental Research Center, Edgewater, MD, USA

    • Jessica Needham
    •  & Sean M. McMahon
  4. Department of Civil and Environmental Engineering, University of California, Irvine, CA, USA

    • Elias C. Massoud
  5. Forest Global Earth Observatory, Smithsonian Tropical Research Institute, Washington, DC, USA

    • Stuart J. Davies
    •  & David Kenfack
  6. Forest Global Earth Observatory, Smithsonian Conservation Biology Institute, Front Royal, VA, USA

    • Kristina J. Anderson-Teixeira
  7. Research Office, Department of National Parks, Wildlife and Plant Conservation, Bangkok, Thailand

    • Sarayudh Bunyavejchewin
  8. Climate and Ecosystems Science Division, Lawrence Berkeley National Laboratory, Berkeley, CA, USA

    • Jeffery Q. Chambers
  9. Department of Natural Resources and Environmental Studies, National Dong Hwa University, Hualien, Taiwan

    • Chia-Hao Chang-Yang
  10. Department of Life Science, Tunghai University, Taichung, Taiwan

    • Jyh-Min Chiang
  11. Department of Botany and Plant Physiology, University of Buea, Buea, Cameroon

    • George B. Chuyong
  12. Field Museum, Chicago, IL, USA

    • Richard Condit
  13. Institute of Pacific Islands Forestry, USDA Forest Service, Hilo, HI, USA

    • Susan Cordell
    •  & Christian P. Giardina
  14. Forest Research Institute Malaysia, Kepong, Selangor Darul Ehsan, Malaysia

    • Christine Fletcher
    • , Abdul Rahman Kassim
    •  & Musalmah Nasardin
  15. Department of Geography, University of Hawai‘i at Mānoa, Honolulu, HI, USA

    • Thomas W. Giambelluca
  16. Department of Botany, University of Peradeniya, Peradeniya, Sri Lanka

    • Nimal Gunatilleke
    •  & Savitri Gunatilleke
  17. Institute of Ecology and Evolutionary Biology, National Taiwan University, Taipei, Taiwan

    • Chang-Fu Hsieh
  18. Center for Tropical Forest Science-Forest Global Earth Observatory, Smithsonian Tropical Research Institute, Panama, Republic of Panama

    • Stephen Hubbell
  19. Department of Natural Resources and Environmental Management, University of Hawaii at Manoa, Honolulu, HI, USA

    • Faith Inman-Narahari
    •  & Creighton M. Litton
  20. Department of Agricultural and Biological Engineering, University of Florida, Gainesville, FL, USA

    • Masatoshi Katabuchi
  21. Nanyang Technological University, Singapore, Singapore

    • Shawn Lum
  22. Sarawak Forestry Department, Kuching, Sarawak, Malaysia

    • Mohizah Mohamad
    •  & Sylvester Tan
  23. Institute of Biology, University of the Philippines Diliman, Quezon City, Philippines

    • Perry S. Ong
  24. Department of Biology, University of Hawaii, Hilo, HI, USA

    • Rebecca Ostertag
  25. Department of Ecology and Evolutionary Biology, University of California, Los Angeles, Los Angeles, CA, USA

    • Lawren Sack
  26. Department of Biology, University of Maryland, Baltimore, MD, USA

    • Nathan G. Swenson
    •  & Maria Natalia Umaña
  27. National Dong Hwa University, Hualian, Taiwan

    • I Fang Sun
  28. School of Biological Sciences, Washington State University, Vancouver, WA, USA

    • Duncan W. Thomas
  29. Centre for Ecology & Hydrology, Bush Estate, Penicuik, Midlothian, UK

    • Jill Thompson
  30. Department of Ecology, Evolution & Environmental Biology, Columbia University, New York, NY, USA

    • Maria Uriarte
  31. Escuela de Ciencias Biológicas, Pontificia Universidad Católica del Ecuador, Quito, Ecuador

    • Renato Valencia
  32. Far Eastern University, Manila, Philippines

    • Sandra Yap
  33. Institute for Tropical Ecosystem Studies, College of Natural Sciences, University of Puerto Rico, Río Piedras, Puerto Rico

    • Jess Zimmerman
  34. Pacific Northwest National Laboratory, Richland, WA, USA

    • Nate G. McDowell


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D.J.J., J.N., C.X., N.M. and S.M.M. conceived and designed the analyses, D.J.J. and J.N. performed the analyses, D.J.J., C.X., S.J.D., N.M., J.N., S.M.M. wrote the initial draft. E.C.M. and C.X. provided FATES simulations. K.J.A.-T., S.B., C.H.C.-Y., J.Q.C., J.-M.C., G.B.C., R.C., S.C., C.F., F.I.-N., C.P.G., S.G., N.G., T.W.G., C.-F.H., S.H., A.R.K., M.K., D.K., C.M.L., S.L., E.C.M., M.M., N.G.M., P.S.O., R.O., L.S., N.G.S., I.F.S., S.T., D.W.T., J.T., M.N.U., M.U., R.V., S.Y. and J.K.Z. contributed data, provided site-specific information and helped revise the manuscript. All authors contributed to the final version of the manuscript.

Competing interests

The authors declare no competing interests.

Corresponding author

Correspondence to Daniel J. Johnson.

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