Abstract
Pathogens and herbivores are key to diversity-productivity relationships. However, the extent to which plant diversity maintains higher productivity by reducing biomass loss from pathogens and herbivores remains unclear at the global scale. Based on a meta-analysis of 2315 observations from 316 studies, we show that, compared to monocultures, plant mixtures on average reduce pathogens abundance and damage to plants by 30.1% and 31.7% and those of invertebrate herbivores by 21.6% and 25.1%, while increasing plant productivity by 40.1% and 35.7%, respectively. Mixture effects on specialist pathogens and invertebrate herbivores become more negative with increasing plant taxonomic, functional, and phylogenetic diversity in mixtures, while those on generalist pathogens and invertebrate herbivores show no significant relationships with any diversity metrics. Mixture effects on pathogens decrease with stand age but those on invertebrate herbivores shift from negative to positive with stand age. Mixture effects on pathogens and invertebrate herbivores are negatively associated with those on productivity. Our findings highlight that conserving plant diversity could reduce biotic damage to plants and enhance global primary productivity.
Similar content being viewed by others
Data availability
The raw data were obtained from previously published studies cited in Supplementary Data 1. The processed data are available at Figshare under accession code https://doi.org/10.6084/m9.figshare.2803257288. Source data are provided with this paper.
Code availability
The code used in this study is available at Figshare https://doi.org/10.6084/m9.figshare.2803257288.
References
Hooper, D. U. et al. A global synthesis reveals biodiversity loss as a major driver of ecosystem change. Nature 486, 105–129 (2012).
Cardinale, B. J. et al. Biodiversity loss and its impact on humanity. Nature 486, 59–67 (2012).
Barry, K. E. et al. The future of complementarity: disentangling causes from consequences. Trends Ecol. Evol. 34, 167–180 (2019).
Otway, S. J., Hector, A. & Lawton, J. H. Resource dilution effects on specialist insect herbivores in a grassland biodiversity experiment. J. Anim. Ecol. 74, 234–240 (2005).
Keesing, F. & Ostfeld, R. S. Dilution effects in disease ecology. Ecol. Lett. 24, 2490–2505 (2021).
Field, E. et al. Associational resistance to both insect and pathogen damage in mixed forests is modulated by tree neighbour identity and drought. J. Ecol. 108, 1511–1522 (2020).
Barbosa, P. et al. Associational resistance and associational susceptibility: having right or wrong neighbors. Annu. Rev. Ecol. Evol. Syst. 40, 1–20 (2009).
Karban, R. & Agrawal, A. A. Herbivore offense. Annu. Rev. Ecol. Evol. Syst. 33, 641–664 (2002).
Wan, N. et al. Plant genetic diversity affects multiple trophic levels and trophic interactions. Nat. Commun. 13, 7312 (2022).
Barbosa, P. & Caldas, A. Do larvae of species in macrolepidopteran assemblages share traits that influence susceptibility to parasitism? Environ. Entomol. 36, 329–336 (2007).
Swenson, N. G. The assembly of tropical tree communities the advances and shortcomings of phylogenetic and functional trait analyses. Ecography 36, 264–276 (2013).
Srivastava, D. S., Cadotte, M. W., MacDonald, A. A. M., Marushia, R. G. & Mirotchnick, N. Phylogenetic diversity and the functioning of ecosystems. Ecol. Lett. 15, 637–648 (2012).
Macia-Vicente, J. G. et al. The structure of root-associated fungal communities is related to the long-term effects of plant diversity on productivity. Mol. Ecol. 32, 3763–3777 (2023).
Siemann, E., Haarstad, J. & Tilman, D. Dynamics of plant and arthropod diversity during old field succession. Ecography 22, 406–414 (1999).
van Ruijven, J., Ampt, E., Francioli, D. & Mommer, L. Do soil-borne fungal pathogens mediate plant diversity-productivity relationships? Evidence and future opportunities. J. Ecol. 108, 1810–1821 (2020).
Burdon, J. J., Thrall, P. H. & Ericson, L. The current and future dynamics of disease in plant communities. Annu. Rev. Phytopathol. 44, 19–39 (2006).
Peralta, A. L., Sun, Y. M., McDaniel, M. D. & Lennon, J. T. Crop rotational diversity increases disease suppressive capacity of soil microbiomes. Ecosphere 9, e02235 (2018).
Schuldt, A. et al. Tree diversity promotes functional dissimilarity and maintains functional richness despite species loss in predator assemblages. Oecologia 174, 533–543 (2014).
Haas, S. E., Hooten, M. B., Rizzo, D. M. & Meentemeyer, R. K. Forest species diversity reduces disease risk in a generalist plant pathogen invasion. Ecol. Lett. 14, 1108–1116 (2011).
Marquis, R. J., Lill, J. T. & Piccinni, A. Effect of plant architecture on colonization and damage by leaftying caterpillars of Quercus alba. Oikos 99, 531–537 (2002).
Singh, B. K. et al. Climate change impacts on plant pathogens, food security and paths forward. Nat. Rev. Microbiol. 21, 640–656 (2023).
Skendzic, S., Zovko, M., Zivkovic, I. P., Lesic, V. & Lemic, D. The impact of climate change on agricultural insect pests. Insects 12, 440 (2021).
Power, A. G. & Mitchell, C. E. Pathogen spillover in disease epidemics. Am. Naturalist 164, S79–S89 (2004).
Kim, T. N. & Underwood, N. Plant neighborhood effects on herbivory: damage is both density and frequency dependent. Ecology 96, 1431–1437 (2015).
Altizer, S., Ostfeld, R. S., Johnson, P. T. J., Kutz, S. & Harvell, C. D. Climate change and infectious diseases: from evidence to a predictive framework. Science 341, 514–519 (2013).
Roy, B. A., Güsewell, S. & Harte, J. Response of plant pathogens and herbivores to a warming experiment. Ecology 85, 2570–2581 (2004).
Liu, X. et al. Dilution effect of plant diversity on infectious diseases: latitudinal trend and biological context dependence. Oikos 129, 457–465 (2020).
Field, E., Hector, A., Barsoum, N. & Koricheva, J. Tree diversity reduces pathogen damage in temperate forests: a systematic review and meta-analysis. Forest Ecol. Manag. 578, 122398 (2025).
Jactel, H., Moreira, X. & Castagneyrol, B. Tree diversity and forest resistance to insect pests: patterns, mechanisms, and prospects. Annu. Rev. Entomol. 66, 277–296 (2021).
Wan, N. et al. Global synthesis of effects of plant species diversity on trophic groups and interactions. Nat. Plants 6, 503–510 (2020).
Yousefi, M. et al. The effectiveness of intercropping and agri-environmental schemes on ecosystem service of biological pest control: a meta-analysis. Agronomy Sustain. Dev. 44, 15 (2024).
Hahn, P. G. & Cammarano, J. H. Environmental context and herbivore traits mediate the strength of associational effects in a meta-analysis of crop diversity. J. Appl. Ecol. 60, 875–885 (2023).
Koricheva, J. & Hayes, D. The relative importance of plant intraspecific diversity in structuring arthropod communities: a meta-analysis. Funct. Ecol. 32, 1704–1717 (2018).
Zhang, Y., Chen, H. Y. H. & Reich, P. B. Forest productivity increases with evenness, species richness and trait variation: a global meta-analysis. J. Ecol. 100, 742–749 (2012).
Jactel, H. et al. Positive biodiversity-productivity relationships in forests: climate matters. Biol. Lett. 14, 20170747 (2018).
Chen, C., Xiao, W. Y. & Chen, H. Y. H. Meta-analysis reveals global variations in plant diversity effects on productivity. Nature 638, 435–440 (2025).
Warner, E. et al. Young mixed planted forests store more carbon than monocultures-a meta-analysis. Front. For. Glob. Change 6, 1226514 (2023).
Beillouin, D., Ben-Ari, T., Malézieux, E., Seufert, V. & Makowski, D. Positive but variable effects of crop diversification on biodiversity and ecosystem services. Glob. Change Biol. 27, 4697–4710 (2021).
Chao, A., Chiu, C.-H. & Jost, L. Unifying species diversity, phylogenetic diversity, functional diversity, and related similarity and differentiation measures through Hill numbers. Annu. Rev. Ecol. Evol. Syst. 45, 297–324 (2014).
Chao, A. et al. Measuring temporal change in alpha diversity: a framework integrating taxonomic, phylogenetic and functional diversity and the standardization. Methods Ecol. Evol. 12, 1926–1940 (2021).
Hantsch, L., Braun, U., Scherer-Lorenzen, M. & Bruelheide, H. Species richness and species identity effects on occurrence of foliar fungal pathogens in a tree diversity experiment. Ecosphere 4, 81 (2013).
Rottstock, T., Joshi, J., Kummer, V. & Fischer, M. Higher plant diversity promotes higher diversity of fungal pathogens, while it decreases pathogen infection per plant. Ecology 95, 1907–1917 (2014).
Benitez, M.-S., Hersh, M. H., Vilgalys, R. & Clark, J. S. Pathogen regulation of plant diversity via effective specialization. Trends Ecol. Evol. 28, 705–711 (2013).
Gilbert, G. S. & Webb, C. O. Phylogenetic signal in plant pathogen-host range. Proc. Natl. Acad. Sci. USA 104, 4979–4983 (2007).
Rutten, G. et al. More diverse tree communities promote foliar fungal pathogen diversity, but decrease infestation rates per tree species, in a subtropical biodiversity experiment. J. Ecol. 109, 2068–2080 (2021).
Bernays, E. A., Bright, K. L., Gonzalez, N. & Angel, J. Dietary mixing in a generalist herbivore: tests of two hypotheses. Ecology 75, 1997–2006 (1994).
Unsicker, S. B., Oswald, A., Koehler, G. & Weisser, W. W. Complementarity effects through dietary mixing enhance the performance of a generalist insect herbivore. Oecologia 156, 313–324 (2008).
Loranger, J. et al. Predicting invertebrate herbivory from plant traits: evidence from 51 grassland species in experimental monocultures. Ecology 93, 2674–2682 (2012).
Parker, J. D., Burkepile, D. E., Lajeunesse, M. J. & Lind, E. M. Phylogenetic isolation increases plant success despite increasing susceptibility to generalist herbivores. Diversity Distrib. 18, 1–9 (2012).
Egorov, E., Gossner, M. M., Meyer, S. T., Weisser, W. W. & Brändle, M. Does plant phylogenetic diversity increase invertebrate herbivory in managed grasslands? Basic Appl. Ecol. 20, 40–50 (2017).
Wang, G. et al. Dilution of specialist pathogens drives productivity benefits from diversity in plant mixtures. Nat. Commun. 14, 8417 (2023).
Boyes, K. N., Hietala-Henschell, K. G., Barton, A. P., Storer, A. J. & Marshall, J. M. Linking tree growth rate, damage repair, and susceptibility to a genus-specific pest infestation. J. For. Res. 30, 1935–1941 (2019).
Morgan, R. E., de Groot, P. & Smith, S. M. Susceptibility of pine plantations to attack by the pine shoot beetle (Tomicus piniperda) in southern Ontario. Can. J. For. Res. 34, 2528–2540 (2004).
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).
Scherber, C. et al. Effects of plant diversity on invertebrate herbivory in experimental grassland. Oecologia 147, 489–500 (2006).
Randolph, S. E. & Dobson, A. D. M. Pangloss revisited: a critique of the dilution effect and the biodiversity-buffers-disease paradigm. Parasitology 139, 847–863 (2012).
Moher, D., Liberati, A., Tetzlaff, J., Altman, D. G. & Grp, P. Preferred reporting items for systematic reviews and meta-analyses: the PRISMA Statement. J. Clin. Epidemiol. 62, 1006–1012 (2009).
Chen, C., Chen, H. Y. H., Chen, X. & Huang, Z. Meta-analysis shows positive effects of plant diversity on microbial biomass and respiration. Nat. Commun. 10, 1332 (2019).
Fick, S. E. & Hijmans, R. J. WorldClim 2: new 1-km spatial resolution climate surfaces for global land areas. Int. J. Climatol. 37, 4302–4315 (2017).
Ali, J. G. & Agrawal, A. A. Specialist versus generalist insect herbivores and plant defense. Trends Plant Sci. 17, 293–302 (2012).
Mordecai, E. A. Pathogen impacts on plant communities: unifying theory, concepts, and empirical work. Ecol. Monogr. 81, 429–441 (2011).
Swanepoel, L. H. et al. A systematic review of rodent pest research in Afro-Malagasy small-holder farming systems: are we asking the right questions? Plos One 12, e0174554 (2017).
Kattge, J. et al. TRY - a global database of plant traits. Glob. Change Biol. 17, 2905–2935 (2011).
Maitner, B. S. et al. The BIEN R package: a tool to access the Botanical Information and Ecology Network (BIEN) database. Methods Ecol. Evol. 9, 373–379 (2018).
Denelle, P., Weigelt, P. & Kreft, H. GIFT—an R package to access the Global Inventory of Floras and Traits. Methods Ecol. Evol. 14, 2738–2748 (2023).
Komatsu, K. J. et al. CoRRE Trait Data: a dataset of 17 categorical and continuous traits for 4079 grassland species worldwide. Sci. Data 11, 795 (2024).
Iversen C. M. et al. Fine-root ecology database (FRED): a global collection of root trait data with coincident site, vegetation, edaphic, and climatic data, version 3. https://doi.org/10.25581/ornlsfa.014/1459186 (2021).
Guerrero-Ramirez, N. R. et al. Global root traits (GRooT) database. Glob. Ecol. Biogeogr. 30, 25–37 (2021).
Buhtz, A., Hohe, A., Schwarz, D. & Grosch, R. Effects of Verticillium dahliae on tomato root morphology considering plant growth response and defence. Plant Pathol. 66, 667–676 (2017).
González-Teuber, M., Palma-Onetto, V., Aguilera-Sammaritano, J. & Mithöfer, A. Roles of leaf functional traits in fungal endophyte colonization: potential implications for host-pathogen interactions. J. Ecol. 109, 3972–3987 (2021).
Schuldt, A. et al. Plant traits affecting herbivory on tree recruits in highly diverse subtropical forests. Ecol. Lett. 15, 732–739 (2012).
Hedges, L. V., Gurevitch, J. & Curtis, P. S. The meta-analysis of response ratios in experimental ecology. Ecology 80, 1150–1156 (1999).
Loreau, M. & Hector, A. Partitioning selection and complementarity in biodiversity experiments. Nature 412, 72–76 (2001).
Pittelkow, C. M. et al. Productivity limits and potentials of the principles of conservation agriculture. Nature 517, 365–482 (2015).
Ma, Z. & Chen, H. Y. H. Effects of species diversity on fine root productivity in diverse ecosystems: a global meta-analysis. Glob. Ecol. Biogeogr. 25, 1387–1396 (2016).
Nakagawa, S. et al. A robust and readily implementable method for the meta-analysis of response ratios with and without missing standard deviations. Ecol. Lett. 26, 232–244 (2023).
Viechtbauer, W. Conducting meta-analyses in R with the metafor Package. J. Stat. Softw. 36, 1–48 (2010).
Nakagawa, S., Yang, Y., Macartney, E. L., Spake, R. & Lagisz, M. Quantitative evidence synthesis: a practical guide on meta-analysis, meta-regression, and publication bias tests for environmental sciences. Environ. Evid. 12, 8 (2023).
Bates, D., Maechler, M., Bolker, B. M. & Walker, S. C. Fitting linear mixed-effects models using lme4. J. Stat. Softw. 67, 1–48 (2015).
Johnson, J. B. & Omland, K. S. Model selection in ecology and evolution. Trends Ecol. Evol. 19, 101–108 (2004).
Zuur, A. F., Ieno, E. N. & Elphick, C. S. A protocol for data exploration to avoid common statistical problems. Methods Ecol. Evol. 1, 3–14 (2010).
Chen, X., Chen, H. Y. H., Searle, E. B., Chen, C. & Reich, P. B. Negative to positive shifts in diversity effects on soil nitrogen over time. Nat. Sustain. 4, 225–234 (2021).
Vetter, D. Handbook of meta-analysis in ecology and evolution. Q. Rev. Biol. 89, 53–54 (2014).
Adams, D. C., Gurevitch, J. & Rosenberg, M. S. Resampling tests for meta-analysis of ecological data. Ecology 78, 1277–1283 (1997).
Dodhia, R. M. Applied multiple regression/correlation analysis for the behavioral sciences. J. Educ. Behav. Stat. 30, 227–229 (2005).
Lefcheck, J. S. PIECEWISESEM: piecewise structural equation modelling in R for ecology, evolution, and systematics. Methods Ecol. Evol. 7, 573–579 (2016).
Grace, J. Structural Equation Modeling and Natural Systems (Cambridge University Press, 2006).
Huang, C., Chen, H., Cheng, W., Chen, X. & Ma, Z. Meta-analysis shows that plant mixtures reduce pathogens and invertebrate herbivores and increase plant productivity. Figshare, https://doi.org/10.6084/m9.figshare.28032572 (2026).
Acknowledgements
This research was supported by National Natural Science Foundation of China (32572024 to Z.M., 32301457 to W.C.), Shenzhen Science and Technology Program to Z.M. (JCYJ20230807111116034), Guangdong Basic and Applied Basic Research Foundation to Z.M. (2023A1515010643) and Fundamental Research Funds for the Central Universities, Sun Yat-sen University to Z.M. (23lgbj009). We thank all the researchers whose data are used in this meta-analysis.
Author information
Authors and Affiliations
Contributions
C.H. and Z.M. conceptualized the study. C.H. contributed to data collection. C.H. and Z.M. conducted the data analysis and drafted the initial manuscript. H.C., W.C., and X.C. contributed to multiple rounds of revisions of the manuscript. All authors reviewed the manuscript and approved the final version for publication.
Corresponding author
Ethics declarations
Competing interests
The authors declare no competing interests.
Peer review
Peer review information
Nature Communications thanks Pilar Fernandez, and the other, anonymous, reviewer(s) for their contribution to the peer review of this work. A peer review file is available.
Additional information
Publisher’s note Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
Source data
Rights and permissions
Open Access This article is licensed under a Creative Commons Attribution-NonCommercial-NoDerivatives 4.0 International License, which permits any non-commercial use, sharing, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if you modified the licensed material. You do not have permission under this licence to share adapted material derived from this article or parts of it. The images or other third party material in this article are included in the article’s Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by-nc-nd/4.0/.
About this article
Cite this article
Huang, C., Chen, H.Y.H., Wenda, C. et al. Meta-analysis shows that plant mixtures reduce pathogens and invertebrate herbivores and increase plant productivity. Nat Commun (2026). https://doi.org/10.1038/s41467-026-70609-7
Received:
Accepted:
Published:
DOI: https://doi.org/10.1038/s41467-026-70609-7


