Abstract
Forest biodiversity plays a critical role in sustaining ecosystem functioning and buffering the effects of increased extreme weather events on forests. A global assessment of the relationship between biodiversity and photosynthesis in natural forest ecosystems, however, remains elusive. We used a large dataset of the richness of tree species from a large number of globally distributed forest plots combined with satellite retrievals of sun-induced chlorophyll fluorescence, a novel proxy for photosynthesis, to evaluate the relationship between forest biodiversity and photosynthesis and its biological mechanisms at the global scale. We found that species richness and photosynthesis were often positively correlated at the global scale, with stronger relationships in tropical forests but weaker associations in high-latitude regions. This positive relationship was mainly driven by a larger role of species richness in increasing maximal photosynthesis than in prolonging the growing season. We also found that higher light capture by increasing the complexity of community structure was the basis of this increase in forest photosynthesis. Forests with high species richness also showed higher foliar nitrogen concentrations and the maximum rate of ribulose 1,5-bisphosphate carboxylase/oxygenase carboxylation, which are two crucial traits determining photosynthetic capacity. Our observation-based findings of ecosystem carbon uptake responses to changes in biodiversity suggest that the loss of biodiversity may jeopardize ecosystem carbon uptake and the terrestrial carbon sink, and will provide important constraints to Earth-system models.
This is a preview of subscription content, access via your institution
Access options
Access Nature and 54 other Nature Portfolio journals
Get Nature+, our best-value online-access subscription
$32.99 / 30 days
cancel any time
Subscribe to this journal
Receive 12 digital issues and online access to articles
$119.00 per year
only $9.92 per issue
Buy this article
- Purchase on SpringerLink
- Instant access to the full article PDF.
USD 39.95
Prices may be subject to local taxes which are calculated during checkout






Similar content being viewed by others
Data availability
The species-richness dataset is available via GitHub at https://github.com/petrkeil/global_tree_S. All MODIS datasets used in this study are available via the Land Processes Distributed Active Archive Center at https://lpdaac.usgs.gov/product_search/. The ERA5-Land monthly climatic datasets are available via the Copernicus Climate Change Service at https://cds.climate.copernicus.eu/cdsapp. The Hansen tree-cover datasets are available via Google Earth Engine at https://glad.earthengine.app/view/global-forest-change. The MERIT DEM dataset is available at http://hydro.iis.u-tokyo.ac.jp/~yamadai/MERIT_DEM/index.html. The biomass dataset is available via Oak Ridge National Laboratory Distributed Active Archive Center at https://daac.ornl.gov/VEGETATION/guides/Global_Maps_C_Density_2010.html. All the above satellite-based data can also be obtained on Google Earth Engine. The TROPOMI SIF datasets are available via CaltechDATA at https://data.caltech.edu/records/8hm1f-w5492. The soil datasets are available via ISRIC Data Hub at https://data.isric.org/geonetwork/srv/chi/catalog.search#/home. The map of the SSCI is available via Göttingen Research Online at https://doi.org/10.25625/9NPEQA. The GEDI datasets are available via Earth Engine Data Catalog at https://developers.google.com/earth-engine/datasets/catalog/LARSE_GEDI_GEDI02_A_002_MONTHLY. The global maps of foliar N concentration are available via GitHub at https://github.com/abhirupdatta/global_maps_of_plant_traits. The global map of the maximum rate of RuBisCO carboxylation is available via Zenodo at https://doi.org/10.5281/zenodo.5090497. The model-based global tree species diversity map is available via Figshare at https://doi.org/10.6084/m9.figshare.17232491.v2. The GFBI dataset is accessible via Global Forest Biodiversity Initiative at https://www.gfbinitiative.org/. The forest age data are available via Max Planck Institute for Biogeochemistry at https://www.bgc-jena.mpg.de/geodb/projects/FileDetails.php.
Code availability
The codes that support the main findings in this study are available via Figshare at https://doi.org/10.6084/m9.figshare.22191919.v3.
References
Hooper, D. U. et al. A global synthesis reveals biodiversity loss as a major driver of ecosystem change. Nature 486, 105–108 (2012).
Maestre, F. T. et al. Plant species richness and ecosystem multifunctionality in global drylands. Science 335, 214–218 (2012).
Duffy, J. E., Godwin, C. M. & Cardinale, B. J. Biodiversity effects in the wild are common and as strong as key drivers of productivity. Nature 549, 261–264 (2017).
Grace, J. B. et al. Integrative modelling reveals mechanisms linking productivity and plant species richness. Nature 529, 390–393 (2016).
van der Plas, F. Biodiversity and ecosystem functioning in naturally assembled communities. Biol. Rev. 94, 1220–1245 (2019).
Liang, J. et al. Positive biodiversity-productivity relationship predominant in global forests. Science 354, aaf8957 (2016).
Mori, A. S. et al. Biodiversity–productivity relationships are key to nature-based climate solutions. Nat. Clim. Chang. 11, 543–550 (2021).
Wardle, D. A., Bardgett, R. D., Callaway, R. M. & Van Der Putten, W. H. Terrestrial ecosystem responses to species gains and losses. Science 332, 1273–1277 (2011).
Fernández-Martínez, M. et al. Diagnosing destabilization risk in global land carbon sinks. Nature https://doi.org/10.1038/s41586-023-05725-1 (2023).
IPCC. Climate Change 2013: The Physical Science Basis (eds Stocker, T. F. et al.) (Cambridge Univ. Press, 2013).
Fernández-Martínez, M. et al. The role of climate, foliar stoichiometry and plant diversity on ecosystem carbon balance. Glob. Chang. Biol. 26, 7067–7078 (2020).
Tilman, D., Isbell, F. & Cowles, J. M. Biodiversity and ecosystem functioning. Annu. Rev. Ecol. Evol. Syst. 45, 471–493 (2014).
Gonzalez, A. et al. Scaling-up biodiversity-ecosystem functioning research. Ecol. Lett. 23, 757–776 (2020).
Loreau, M. & Hector, A. Partitioning selection and complementarity in biodiversity experiments. Nature 412, 72–76 (2001).
Huang, Y. et al. Impacts of species richness on productivity in a large-scale subtropical forest experiment. Science 362, 80–83 (2018).
Sapijanskas, J., Paquette, A., Potvin, C., Kunert, N. & Loreau, M. Tropical tree diversity enhances light capture through crown plasticity and spatial and temporal niche differences. Ecology 95, 2479–2492 (2014).
Hooper, D. U. & Vitousek, P. M. The effects of plant composition and diversity on ecosystem processes. Science 277, 1302–1305 (1997).
Tobner, C. M. et al. Functional identity is the main driver of diversity effects in young tree communities. Ecol. Lett. 19, 638–647 (2016).
Zuppinger-Dingley, D. et al. Selection for niche differentiation in plant communities increases biodiversity effects. Nature 515, 108–111 (2014).
Dronova, I. & Taddeo, S. Remote sensing of phenology: towards the comprehensive indicators of plant community dynamics from species to regional scales. J. Ecol. https://doi.org/10.1111/1365-2745.13897 (2022).
Duarte, M. M. et al. High tree diversity enhances light interception in tropical forests. J. Ecol. 109, 2597–2611 (2021).
Williams, L. J., Paquette, A., Cavender-Bares, J., Messier, C. & Reich, P. B. Spatial complementarity in tree crowns explains overyielding in species mixtures. Nat. Ecol. Evol. 1, 63 (2017).
Chen, X. et al. Tree diversity increases decadal forest soil carbon and nitrogen accrual. Nature https://doi.org/10.1038/s41586-023-05941-9 (2023).
Chen, X., Chen, H. Y. H. & Chang, S. X. Meta-analysis shows that plant mixtures increase soil phosphorus availability and plant productivity in diverse ecosystems. Nat. Ecol. Evol. 6, 1112–1121 (2022).
Luo, X. & Keenan, T. F. Global evidence for the acclimation of ecosystem photosynthesis to light. Nat. Ecol. Evol. 4, 1351–1357 (2020).
Jung, M. et al. Compensatory water effects link yearly global land CO2 sink changes to temperature. Nature 541, 516–520 (2017).
Miao, W. et al. Effects of biodiversity, stand factors and functional identity on biomass and productivity during the restoration of subtropical forests in Central China. J. Plant Ecol. 15, 385–398 (2022).
Lepš, J. What do the biodiversity experiments tell us about consequences of plant species loss in the real world? Basic Appl. Ecol. 5, 529–534 (2004).
Wardle, D. A. & Jonsson, M. Biodiversity effects in real ecosystems – a response to Duffy. Front. Ecol. Environ. 8, 10–11 (2010).
Wardle, D. A. Do experiments exploring plant diversity–ecosystem functioning relationships inform how biodiversity loss impacts natural ecosystems? J. Veg. Sci. 27, 646–653 (2016).
Jochum, M. et al. The results of biodiversity–ecosystem functioning experiments are realistic. Nat. Ecol. Evol. 4, 1485–1494 (2020).
Keil, P. & Chase, J. M. Global patterns and drivers of tree diversity integrated across a continuum of spatial grains. Nat. Ecol. Evol. 3, 390–399 (2019).
Guanter, L. et al. Global and time-resolved monitoring of crop photosynthesis with chlorophyll fluorescence. Proc. Natl Acad. Sci. USA 111, E1327–E1333 (2014).
Zeng, Y. et al. Optical vegetation indices for monitoring terrestrial ecosystems globally. Nat. Rev. Earth Environ. 3, 477–493 (2022).
Badgley, G., Field, C. B. & Berry, J. A. Canopy near-infrared reflectance and terrestrial photosynthesis. Sci. Adv. 3, 1–6 (2017).
Liang, J. et al. Co-limitation towards lower latitudes shapes global forest diversity gradients. Nat. Ecol. Evol. https://doi.org/10.1038/s41559-022-01831-x (2022).
Olson, D. M. et al. Terrestrial ecoregions of the world: a new map of life on Earth. Bioscience 51, 933–938 (2001).
Liu, D., Wang, T., Peñuelas, J. & Piao, S. Drought resistance enhanced by tree species diversity in global forests. Nat. Geosci. https://doi.org/10.1038/s41561-022-01026-w (2022).
Doughty, R. et al. TROPOMI reveals dry-season increase of solar-induced chlorophyll fluorescence in the Amazon forest. Proc. Natl Acad. Sci. USA 116, 22393–22398 (2019).
Ratcliffe, S. et al. Biodiversity and ecosystem functioning relations in European forests depend on environmental context. Ecol. Lett. 20, 1414–1426 (2017).
Searle, E. B. & Chen, H. Y. H. Complementarity effects are strengthened by competition intensity and global environmental change in the central boreal forests of Canada. Ecol. Lett. 23, 79–87 (2020).
Xia, J. et al. Joint control of terrestrial gross primary productivity by plant phenology and physiology. Proc. Natl Acad. Sci. USA 112, 2788–2793 (2015).
Oehri, J., Schmid, B., Schaepman-Strub, G. & Niklaus, P. A. Biodiversity promotes primary productivity and growing season lengthening at the landscape scale. Proc. Natl Acad. Sci. USA 114, 10160–10165 (2017).
Wolf, A. A., Zavaleta, E. S. & Selmants, P. C. Flowering phenology shifts in response to biodiversity loss. Proc. Natl Acad. Sci. USA 114, 3463–3468 (2017).
Gough, C. M., Atkins, J. W., Fahey, R. T. & Hardiman, B. S. High rates of primary production in structurally complex forests. Ecology 100, 1–6 (2019).
Zhang, L., Liu, X., Zhou, S. & Shipley, B. Explaining variation in productivity requires intraspecific variability in plant height among communities. J. Plant Ecol. 15, 310–319 (2022).
Godlee, J. L. et al. Structural diversity and tree density drives variation in the biodiversity–ecosystem function relationship of woodlands and savannas. New Phytol. 232, 579–594 (2021).
Fahey, R. T. et al. Defining a spectrum of integrative trait-based vegetation canopy structural types. Ecol. Lett. 22, 2049–2059 (2019).
Ishii, H. & Asano, S. The role of crown architecture, leaf phenology and photosynthetic activity in promoting complementary use of light among coexisting species in temperate forests. Ecol. Res. 25, 715–722 (2010).
Williams, L. J. et al. Enhanced light interception and light use efficiency explain overyielding in young tree communities. Ecol. Lett. 24, 996–1006 (2021).
Williams, L. J., Cavender-Bares, J., Paquette, A., Messier, C. & Reich, P. B. Light mediates the relationship between community diversity and trait plasticity in functionally and phylogenetically diverse tree mixtures. J. Ecol. 108, 1617–1634 (2020).
He, M. et al. Global spectrum of vegetation light‐use efficiency. Geophys. Res. Lett. 49, 1–9 (2022).
Musavi, T. et al. Stand age and species richness dampen interannual variation of ecosystem-level photosynthetic capacity. Nat. Ecol. Evol. 1, 0048 (2017).
Farquhar, G. D., von Caemmerer, S. & Berry, J. A. A biochemical model of photosynthetic CO2 assimilation in leaves of C3 species. Planta 149, 78–90 (1980).
Chen, J. M. et al. Global datasets of leaf photosynthetic capacity for ecological and earth system research. Earth Syst. Sci. Data Discuss. 2022, 1–26 (2022).
Luo, X. et al. Global variation in the fraction of leaf nitrogen allocated to photosynthesis. Nat. Commun. 12, 1–10 (2021).
Williams, L. J. et al. Remote spectral detection of biodiversity effects on forest biomass. Nat. Ecol. Evol. 5, 46–54 (2021).
Guiz, J. et al. Long-term effects of plant diversity and composition on plant stoichiometry. Oikos 125, 613–621 (2016).
Sardans, J. et al. Empirical support for the biogeochemical niche hypothesis in forest trees. Nat. Ecol. Evol. 5, 184–194 (2021).
Peñuelas, J. et al. The bioelements, the elementome, and the biogeochemical niche. Ecology 100, 1–15 (2019).
Fernández-Martínez, M. et al. Bryophyte C:N:P stoichiometry, biogeochemical niches and elementome plasticity driven by environment and coexistence. Ecol. Lett. 24, 1375–1386 (2021).
Peaucelle, M. et al. Spatial variance of spring phenology in temperate deciduous forests is constrained by background climatic conditions. Nat. Commun. 10, 1–10 (2019).
Zhang, W. P. et al. Interspecific interactions affect N and P uptake rather than N:P ratios of plant species: evidence from intercropping. J. Plant Ecol. 15, 223–236 (2022).
Cavender-Bares, J. et al. Integrating remote sensing with ecology and evolution to advance biodiversity conservation. Nat. Ecol. Evol. https://doi.org/10.1038/s41559-022-01702-5 (2022).
Brun, P. et al. The productivity-biodiversity relationship varies across diversity dimensions. Nat. Commun. 10, 5691 (2019).
Pau, S., Gillespie, T. W. & Wolkovich, E. M. Dissecting NDVI-species richness relationships in Hawaiian dry forests. J. Biogeogr. 39, 1678–1686 (2012).
Madonsela, S., Cho, M. A., Ramoelo, A., Mutanga, O. & Naidoo, L. Estimating tree species diversity in the savannah using NDVI and woody canopy cover. Int. J. Appl. Earth Obs. Geoinf. 66, 106–115 (2018).
Barry, K. E. et al. A graphical null model for scaling biodiversity–ecosystem functioning relationships. J. Ecol. 109, 1549–1560 (2021).
Zhang, Z., Zhang, Y. & Zhang, Y. Generating high-resolution total canopy SIF emission from TROPOMI data: algorithm and application. Remote Sens. Environ. 295, 113699 (2023).
Anderegg, L. D. L. et al. Representing plant diversity in land models: an evolutionary approach to make ‘Functional Types’ more functional. Glob. Chang. Biol. 28, 2541–2554 (2022).
Fisher, R. A. et al. Vegetation demographics in Earth System Models: a review of progress and priorities. Glob. Chang. Biol. 24, 35–54 (2018).
Sakschewski, B. et al. Resilience of Amazon forests emerges from plant trait diversity. Nat. Clim. Chang. 6, 1032–1036 (2016).
Jung, M. et al. Recent decline in the global land evapotranspiration trend due to limited moisture supply. Nature 467, 951–954 (2010).
Wang, S. et al. Recent global decline of CO2 fertilization effects on vegetation photosynthesis. Science 370, 1295–1300 (2020).
Soto-Navarro, C. et al. Mapping co-benefits for carbon storage and biodiversity to inform conservation policy and action. Philos. Trans. R. Soc. B 375, 20190128 (2020).
Luby, I. H., Miller, S. J. & Polasky, S. When and where to protect forests. Nature 609, 89–93 (2022).
Yang, X. et al. Global patterns of potential future plant diversity hidden in soil seed banks. Nat. Commun. 12, 1–8 (2021).
Hansen, M. C. et al. High-resolution global maps of 21st-century forest cover change. Science 342, 850–853 (2013).
García-Palacios, P., Gross, N., Gaitán, J. & Maestre, F. T. Climate mediates the biodiversity–ecosystem stability relationship globally. Proc. Natl Acad. Sci. USA 115, 8400–8405 (2018).
Porcar-Castell, A. et al. Chlorophyll a fluorescence illuminates a path connecting plant molecular biology to Earth-system science. Nat. Plants 7, 998–1009 (2021).
Köhler, P. et al. Global retrievals of solar-induced chlorophyll fluorescence with TROPOMI: first results and intersensor comparison to OCO-2. Geophys. Res. Lett. 45, 456–10,463 (2018).
Magney, T. S., Barnes, M. L. & Yang, X. On the covariation of chlorophyll fluorescence and photosynthesis across scales. Geophys. Res. Lett. 47, 1–7 (2020).
Ryu, Y., Jiang, C., Kobayashi, H. & Detto, M. MODIS-derived global land products of shortwave radiation and diffuse and total photosynthetically active radiation at 5 km resolution from 2000. Remote Sens. Environ. 204, 812–825 (2018).
Spawn, S. A., Sullivan, C. C., Lark, T. J. & Gibbs, H. K. Harmonized global maps of above and belowground biomass carbon density in the year 2010. Sci. Data 7, 1–22 (2020).
Yamazaki, D. et al. A high-accuracy map of global terrain elevations. Geophys. Res. Lett. 44, 5844–5853 (2017).
Poggio, L. et al. SoilGrids 2.0: producing soil information for the globe with quantified spatial uncertainty. SOIL 7, 217–240 (2021).
Roerink, G. J., Menenti, M. & Verhoef, W. Reconstructing cloudfree NDVI composites using Fourier analysis of time series. Int. J. Remote Sens. 21, 1911–1917 (2000).
Dechant, B. et al. Canopy structure explains the relationship between photosynthesis and sun-induced chlorophyll fluorescence in crops. Remote Sens. Environ. 241, 111733 (2020).
Zhang, Z. et al. Sensitivity of estimated total canopy SIF emission to remotely sensed LAI and BRDF products. J. Remote Sens. 2021, 1–18 (2021).
Zhang, Z. et al. The potential of satellite FPAR product for GPP estimation: an indirect evaluation using solar-induced chlorophyll fluorescence. Remote Sens. Environ. 240, 111686 (2020).
Moreno-Martínez, Á. et al. A methodology to derive global maps of leaf traits using remote sensing and climate data. Remote Sens. Environ. 218, 69–88 (2018).
Yang, H. et al. Climatic and biotic factors influencing regional declines and recovery of tropical forest biomass from the 2015/16 El Niño. Proc. Natl Acad. Sci. USA 119, e2101388119 (2022).
Marselis, S. M., Keil, P., Chase, J. M. & Dubayah, R. The use of GEDI canopy structure for explaining variation in tree species richness in natural forests. Environ. Res. Lett. 17, 045003 (2022).
Ehbrecht, M. et al. Global patterns and climatic controls of forest structural complexity. Nat. Commun. 12, 1–12 (2021).
Zhai, L., Coyle, D. R., Li, D. & Jonko, A. Fire, insect and disease-caused tree mortalities increased in forests of greater structural diversity during drought. J. Ecol. https://doi.org/10.1111/1365-2745.13830 (2022).
Besnard, S. et al. Mapping global forest age from forest inventories, biomass and climate data. Earth Syst. Sci. Data 13, 4881–4896 (2021).
Dormann, F. C. et al. Methods to account for spatial autocorrelation in the analysis of species distributional data: a review. Ecography 30, 609–628 (2007).
Gross, N. et al. Functional trait diversity maximizes ecosystem multifunctionality. Nat. Ecol. Evol. 1, 0132 (2017).
Norton, A. J. et al. Hydrologic connectivity drives extremes and high variability in vegetation productivity across Australian arid and semi-arid ecosystems. Remote Sens. Environ. 272, 112937 (2022).
Lefcheck, J. S. piecewiseSEM: piecewise structural equation modelling in r for ecology, evolution, and systematics. Methods Ecol. Evol. 7, 573–579 (2016).
Acknowledgements
This research was supported by the National Science Foundation of China (42125105), the Spanish Government TED2021-132627B-I00 funded by MCIN/AEI/10.13039/501100011033 and the European NextGenerationEU/PRTR, the Fundación Ramón Areces grant CIVP20A6621 and the Catalan government project SGR2021-01333. M.F.-M. was supported by the European Research Council project ERC-StG-2022-101076740 STOIKOS and a Ramón y Cajal fellowship (RYC2021-031511-I) funded by the Spanish Ministry of Science and Innovation, the NextGenerationEU program of the European Union, the Spanish plan of recovery, transformation and resilience and the Spanish Agency of Research.
Author information
Authors and Affiliations
Contributions
Y.Z. designed the research. R.C. performed the analysis. R.C. and Y.Z. drafted the paper. J.P., Z.Z., and M.F.-M. contributed to the interpretation of the results and to the writing of the paper. W.J. and G.L. contributed to the writing of the paper.
Corresponding authors
Ethics declarations
Competing interests
The authors declare no competing interests.
Peer review
Peer review information
Nature Plants thanks Arthur Gessler, Xiaojuan Liu and the other, anonymous, reviewer(s) for their contribution to the peer review of this work.
Additional information
Publisher’s note Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
Supplementary information
Supplementary Information
Supplementary Figs. 1–25 and Tables 1–8.
Rights and permissions
Springer Nature or its licensor (e.g. a society or other partner) holds exclusive rights to this article under a publishing agreement with the author(s) or other rightsholder(s); author self-archiving of the accepted manuscript version of this article is solely governed by the terms of such publishing agreement and applicable law.
About this article
Cite this article
Cao, R., Zhang, Y., Fernández-Martínez, M. et al. Global evidence for a positive relationship between tree species richness and ecosystem photosynthesis. Nat. Plants 11, 1429–1440 (2025). https://doi.org/10.1038/s41477-025-02046-1
Received:
Accepted:
Published:
Version of record:
Issue date:
DOI: https://doi.org/10.1038/s41477-025-02046-1
This article is cited by
-
… of the Year
Nature Plants (2025)


