Abstract
Understanding the mechanisms that shape ecosystem resistance to increasing livestock grazing pressure, a major driver of land degradation, is essential for predicting its impacts and informing sustainable land management strategies. This issue is particularly relevant in drylands, which host half of the world’s livestock production and are highly vulnerable to desertification caused by overgrazing. Here we conduct a standardized field survey across 73 dryland sites in 25 countries to simultaneously evaluate how climatic, edaphic, vegetation and grazing-related factors influence ecosystem resistance—defined here as the capacity to maintain vegetation cover under increasing grazing pressure. We found that increasing grazing pressure reduced vegetation cover in 80% of sites, with an average decline of 35%. Plant species richness emerged as the strongest predictor of ecosystem resistance, with higher richness associated with lower vegetation cover loss. Functional trait data indicated that this positive effect was mainly explained by complementarity in trait values among plants, rather than by functional redundancy. Our results indicate that conserving plant diversity is key to strengthening ecosystem resistance and sustaining dryland functioning under intensifying grazing pressure.
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Unforeseen plant phenotypic diversity in a dry and grazed world
Climate-driven ecological thresholds in China’s drylands modulated by grazing
Hotspots of biogeochemical activity linked to aridity and plant traits across global drylands
Data availability
The dataset needed to reproduce our results are available via Figshare at https://doi.org/10.6084/m9.figshare.29132654 (ref. 90).
Code availability
The R script used is available via Figshare at https://doi.org/10.6084/m9.figshare.29132654 (ref. 90).
References
Middleton, N., Stringer, L., Goudie, A. & Thomas, D. The Forgotten Billion: MDG Achievement in the Drylands (United Nations Development Programme, 2011).
United Nations. Global Drylands: A UN System-Wide Response. (United Nations Environment Programme, 2011).
Maestre, F. T. et al. Grazing and ecosystem service delivery in global drylands. Science (1979) 378, 915–920 (2022).
Google Scholar
Yahdjian, L., Sala, O. E. & Havstad, K. M. Rangeland ecosystem services: shifting focus from supply to reconciling supply and demand. Front Ecol. Environ. 13, 44–51 (2015).
Google Scholar
Asner, G. P., Elmore, A. J., Olander, L. P., Martin, R. E. & Harris, A. T. Grazing systems, ecosystem responses and global change. Annu Rev. Environ. Resour. 29, 261–299 (2004).
Google Scholar
D’Odorico, P., Bhattachan, A., Davis, K. F., Ravi, S. & Runyan, C. W. Global desertification: drivers and feedbacks. Adv. Water Resour. 51, 326–344 (2013).
Google Scholar
Schlesinger, W. H. et al. Biological feedbacks in global desertification. Science (1979) 247, 1043–1048 (1990).
Google Scholar
Biancari, L. et al. Drivers of woody dominance across global drylands. Sci. Adv. 10, eadn6007 (2024).
Google Scholar
Van Meerbeek, K., Jucker, T. & Svenning, J.-C. Unifying the concepts of stability and resilience in ecology. J. Ecol. 109, 3114–3132 (2021).
Google Scholar
Wolf, J., Chen, M. & Asrar, G. R. Global rangeland primary production and its consumption by livestock in 2000–2010. Remote Sens. 13, 3430 (2021).
Google Scholar
Gaitán, J. J. et al. Aridity and overgrazing have convergent effects on ecosystem structure and functioning in patagonian rangelands. Land Degrad. Dev. 29, 210–218 (2018).
Google Scholar
Li, C. et al. Climate-driven ecological thresholds in China’s drylands modulated by grazing. Nat. Sustain 6, 1363–1372 (2023).
Google Scholar
Oñatibia, G. R., Amengual, G., Boyero, L. & Aguiar, M. R. Aridity exacerbates grazing-induced rangeland degradation: a population approach for dominant grasses. J. Appl. Ecol. 57, 1999–2009 (2020).
Google Scholar
Milchunas, D. G., Sala, O. E. & Lauenroth, W. K. A generalized model of the effects of grazing by large herbivores on grassland community structure. Am. Nat. 132, 87–106 (1988).
Google Scholar
Adler, P. B., Milchunas, D. G., Lauenroth, W. K., Sala, O. E. & Burke, I. C. Functional traits of graminoids in semi-arid steppes: a test of grazing histories. J. Appl. Ecol. 41, 653–663 (2004).
Google Scholar
Adler, P. B., Milchunas, D. G., Sala, O. E., Burke, I. C. & Lauenroth, W. K. Plant traits and ecosystem grazing effects: comparison of U.S. sagebrush steppe and patagonian steppe. Ecol. Appl. 15, 774–792 (2005).
Google Scholar
Oñatibia, G. R. & Aguiar, M. R. On the early warning signal of degradation in drylands: patches or plants?. J. Ecol. 111, 428–435 (2023).
Google Scholar
Flombaum, P. & Sala, O. E. Higher effect of plant species diversity on productivity in natural than artificial ecosystems. Proc. Natl Acad. Sci. USA 105, 6087–6090 (2008).
Google Scholar
Maestre, F. T. et al. Plant species richness and ecosystem multifunctionality in global drylands. Science 335, 214–218 (2012).
Google Scholar
Tilman, D., Isbell, F. & Cowles, J. M. Biodiversity and Ecosystem Functioning. Annu Rev. Ecol. Evol. Syst. 45, 471–493 (2014).
Google Scholar
Binder, S., Isbell, F., Polasky, S., Catford, J. A. & Tilman, D. Grassland biodiversity can pay. Proc. Natl Acad. Sci. USA 115, 3876–3881 (2018).
Google Scholar
Schaub, S. et al. Plant diversity effects on forage quality, yield and revenues of semi-natural grasslands. Nat. Commun. 11, 768 (2020).
Google Scholar
Schaub, S., Buchmann, N., Lüscher, A. & Finger, R. Economic benefits from plant species diversity in intensively managed grasslands. Ecol. Econ. 168, 106488 (2020).
Google Scholar
McNaughton, S. J. Ecology of a grazing ecosystem: the serengeti. Ecol. Monogr. 55, 259–294 (1985).
Google Scholar
Pillar, V. D. et al. Functional redundancy and stability in plant communities. J. Veg. Sci. 24, 963–974 (2013).
Google Scholar
Hempson, G. P., Archibald, S. & Bond, W. J. The consequences of replacing wildlife with livestock in Africa. Sci. Rep. 7, 17196 (2017).
Google Scholar
Manier, D. J. & Hobbs, N. T. Large herbivores in sagebrush steppe ecosystems: livestock and wild ungulates influence structure and function. Oecologia 152, 739–750 (2007).
Google Scholar
Maestre, F. T. et al. The BIODESERT survey: assessing the impacts of grazing on the structure and functioning of global drylands. Web. Ecol. 22, 75–96 (2022).
Google Scholar
McNaughton, S. J., Oesterheld, M. & Sala, O. E. Large herbivores in rangelands. Nature 364, 293 (1993).
Google Scholar
Oesterheld, M., Loreti, J., Semmartin, M. & Paruelo, J. M. Grazing, fire, and climate effects on primary productivity of grasslands and savannas. In Ecosystems of the World 16: Ecosystems of Disturbed Grounds (ed. Walker, L. R.) 287–306 (Elsevier, 1999).
Fensham, R. J. & Fairfax, R. J. Water-remoteness for grazing relief in Australian arid-lands. Biol. Conserv. 141, 1447–1460 (2008).
Google Scholar
Schrama, M. et al. Cessation of grazing causes biodiversity loss and homogenization of soil food webs. Proc. R. Soc. B 290, 20231345 (2023).
Google Scholar
Gross, N. et al. Unforeseen plant phenotypic diversity in a dry and grazed world. Nature 632, 808–814 (2024).
Google Scholar
Aguiar, M. R. & Sala, O. E. Patch structure, dynamics and implications for the functioning of arid ecosystems. Trends Ecol. Evol. 14, 273–277 (1999).
Google Scholar
Ludwig, J. A. & Tongway, D. J. Spatial organisation of landscapes and its function in semi-arid woodlands. Aust. Landsc. Ecol. 10, 51–63 (1995).
Google Scholar
Bartoń, K. MuMIn: multi-model inference. MuMIn https://doi.org/10.32614/CRAN.package (2023).
Isbell, F. et al. Biodiversity increases the resistance of ecosystem productivity to climate extremes. Nature 526, 574–577 (2015).
Google Scholar
Wagg, C. et al. Biodiversity–stability relationships strengthen over time in a long-term grassland experiment. Nat. Commun. 13, 7752 (2022).
Google Scholar
Bailey, D. W. et al. Synthesis paper: targeted livestock grazing: prescription for healthy rangelands. Rangel. Ecol. Manag 72, 865–877 (2019).
Google Scholar
Li, Y. et al. Adaptive grazing enhances plant species richness and density in the soil seed bank in a semi-arid grassland. Ecol. Indic. 163, 112113 (2024).
Google Scholar
Teague, R. & Kreuter, U. Managing grazing to restore soil health, ecosystem function, and ecosystem services. Front. Sustain. Food Syst. 4, 34187 (2020).
Google Scholar
Su, J., Xu, F. & Zhang, Y. Grassland biodiversity and ecosystem functions benefit more from cattle than sheep in mixed grazing: A meta-analysis. J. Environ. Manag. 337, 117769 (2023).
Google Scholar
Deraison, H., Badenhausser, I., Loeuille, N., Scherber, C. & Gross, N. Functional trait diversity across trophic levels determines herbivore impact on plant community biomass. Ecol. Lett. 18, 1346–1355 (2015).
Google Scholar
Gross, N., Suding, K. N., Lavorel, S. & Roumet, C. Complementarity as a mechanism of coexistence between functional groups of grasses. J. Ecol. 95, 1296–1305 (2007).
Google Scholar
Strauss, S. Y. & Agrawal, A. A. The ecology and evolution of plant tolerance to herbivory. Trends Ecol. Evol. 14, 179–185 (1999).
Google Scholar
Valencia, E. et al. Functional diversity enhances the resistance of ecosystem multifunctionality to aridity in Mediterranean drylands. N. Phytol. 206, 660–671 (2015).
Google Scholar
Laliberté, E. & Legendre, P. A distance-based framework for measuring functional diversity from multiple traits. Ecology 91, 299–305 (2010).
Google Scholar
Ricotta, C. et al. Measuring the functional redundancy of biological communities: a quantitative guide. Methods Ecol. Evol. 7, 1386–1395 (2016).
Google Scholar
Biancari, L., Oñatibia, G. R., Gaitán, J. J. & Aguiar, M. R. Coexistence of grasses and shrubs in Patagonian steppes. Norm or exception?. J. Veg. Sci. 34, e13177 (2023).
Google Scholar
Eldridge, D. J. et al. Impacts of shrub encroachment on ecosystem structure and functioning: towards a global synthesis. Ecol. Lett. 14, 709–722 (2011).
Google Scholar
Eldridge, D. J., Soliveres, S., Bowker, M. A. & Val, J. Grazing dampens the positive effects of shrub encroachment on ecosystem functions in a semi-arid woodland. J. Appl. Ecol. 50, 1028–1038 (2013).
Google Scholar
Kulmatiski, A. & Beard, K. H. Woody plant encroachment facilitated by increased precipitation intensity. Nat. Clim. Chang 3, 833–837 (2013).
Google Scholar
Maestre, F. T. et al. Biogeography of global drylands. N. Phytol. 231, 540–558 (2021).
Google Scholar
Maestre, F. T., Callaway, R. M., Valladares, F. & Lortie, C. J. Refining the stress-gradient hypothesis for competition and facilitation in plant communities. J. Ecol. 97, 199–205 (2009).
Google Scholar
Graff, P. & Aguiar, M. R. Testing the role of biotic stress in the stress gradient hypothesis. Process. Patterns Arid Rangel. Oikos 120, 1023–1030 (2011).
Brooker, R. W. et al. Facilitation in plant communities: the past, the present, and the future. J. Ecol. 96, 18–34 (2008).
Google Scholar
Callaway, R. M., Kikodze, D., Chiboshvili, M. & Khetsuriani, L. Unpalatable plants protect neighbors from grazing and increase plant community diversity. Ecology 86, 1856–1862 (2005).
Google Scholar
Milchunas, D. G. & Noy-Meir, I. Grazing refuges, external avoidance of herbivory and plant diversity. Oikos 99, 113–130 (2002).
Google Scholar
Biancari, L., Aguiar, M. R. & Cipriotti, P. A. Influence of plant neighborhood on patch dynamics. Oikos e11307 (2025); https://doi.org/10.1002/oik.11307
Kéfi, S. et al. Spatial vegetation patterns and imminent desertification in Mediterranean arid ecosystems. Nature 449, 213–217 (2007).
Google Scholar
Isbell, F. I. & Wilsey, B. J. Increasing native, but not exotic, biodiversity increases aboveground productivity in ungrazed and intensely grazed grasslands. Oecologia 165, 771–781 (2011).
Google Scholar
Jawuoro, S. O., Koech, O. K., Karuku, G. N. & Mbau, J. S. Plant species composition and diversity depending on piospheres and seasonality in the southern rangelands of Kenya. Ecol. Process 6, 16 (2017).
Google Scholar
Eldridge, D. J. et al. Dung predicts the global distribution of herbivore grazing pressure in drylands. Nat. Food 6, 253–259 (2025).
Google Scholar
Lee, M. A. A global comparison of the nutritive values of forage plants grown in contrasting environments. J. Plant Res. 131, 641–654 (2018).
Google Scholar
McIntyre, S. The role of plant leaf attributes in linking land use to ecosystem function in temperate grassy vegetation. Agric. Ecosyst. Environ. 128, 251–258 (2008).
Google Scholar
Pontes, L. D. S., Sousanna, J.-F., Louault, F., Andueza, D. & Carrère, P. Leaf traits affect the above-ground productivity and quality of pasture grasses. Funct. Ecol. 21, 844–853 (2007).
Google Scholar
Yeoh, H.-H. & Wee, Y.-C. Leaf protein contents and nitrogen-to-protein conversion factors for 90 plant species. Food Chem. 49, 245–250 (1994).
Google Scholar
Díaz, S. et al. Plant trait responses to grazing—a global synthesis. Glob. Chang Biol. 13, 313–341 (2007).
Google Scholar
Díaz, S. et al. The global spectrum of plant form and function. Nature 529, 167–171 (2016).
Google Scholar
Schneider, C. A., Rasband, W. S. & Eliceiri, K. W. NIH Image to ImageJ: 25 years of image analysis. Nat. Methods 9, 671–675 (2012).
Google Scholar
Waterman, P. G. & Mole, S. Analysis of Phenolic Plant Metabolites (Wiley-Blackwell, 1994).
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).
Google Scholar
Wang, L. et al. Dryland productivity under a changing climate. Nat. Clim. Chang 12, 981–994 (2022).
Google Scholar
Kettler, T. A., Doran, J. W. & Gilbert, T. L. Simplified method for soil particle-size determination to accompany soil-quality analyses. Soil Sci. Soc. Am. J. 65, 849–852 (2001).
Google Scholar
Abanda, P. A., Compton, J. S. & Hannigan, R. E. Soil nutrient content, above-ground biomass and litter in a semi-arid shrubland, South Africa. Geoderma 164, 128–137 (2011).
Google Scholar
Begon, M., Townsend, C. R. & Harper, J. L. Ecology: From Individuals to Ecosystems (Blackwell Publishing, 2006).
Naeem, S. Species redundancy and ecosystem reliability. Conserv. Biol. 12, 39–45 (1998).
Google Scholar
Pavoine, S. adiv: an R package to analyse biodiversity in ecology. Methods Ecol. Evol. 11, 1106–1112 (2020).
Google Scholar
Burnham, K. P. & Anderson, D. R. Model Selection and Multimodel Inference: A Practical Information-Theoretic Approach (Springer, 2002); https://doi.org/10.1007/b97636
Graham, M. H. Confronting multicollinearity in ecological multiple regression. Ecology 84, 2809–2815 (2003).
Google Scholar
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).
Google Scholar
GraphPad Prism Software (GraphPad, 2025); https://www.graphpad.com
Schielzeth, H. Simple means to improve the interpretability of regression coefficients. Methods Ecol. Evol. 1, 103–113 (2010).
Google Scholar
Harrison, X. A. et al. A brief introduction to mixed effects modelling and multi-model inference in ecology. PeerJ 6, e4794 (2018).
Google Scholar
Dray, S. & Dufour, A.-B. The ade4 package: implementing the duality diagram for ecologists. J. Stat. Softw. 22, 1–20 (2007).
Google Scholar
Bivand, R. S., Pebesma, E. & Gómez-Rubio, V. Applied Spatial Data Analysis with R (Springer, 2013); https://doi.org/10.1007/978-1-4614-7618-4
Bivand, R. R. Packages for analyzing spatial data: a comparative case study with areal data. Geogr. Anal. 54, 488–518 (2022).
Google Scholar
R Core Team. R: A Language and Environment for Statistical Computing. (R Foundation for Statistical Computing, 2024); https://www.r-project.org
Rosseel, Y. lavaan: an R package for structural equation modeling. J. Stat. Softw. 48, 1–36 (2012).
Google Scholar
Biancari, L. et al. Plant diversity enhances ecosystem resistance to increasing grazing pressure in global drylands. Figshare https://doi.org/10.6084/m9.figshare.29132654 (2025).
Acknowledgements
We thank all participants of the BIODESERT global field survey. This survey was funded by the European Research Council (ERC grant agreement 647038), awarded to F.T.M. F.T.M., L.B. and E.G. acknowledge support by the King Abdullah University of Science and Technology (KAUST). M.R.A., G.R.O. and L.Y. acknowledge support by the University of Buenos Aires and CONICET. E.V. was supported by the Spanish Ministry of Science, Innovation and Universities (grant nos. PID2022-140398NA-I00 and CNS2024-154579).
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Conceptualization: L.B., F.T.M., Y.L.B.P., N.G., G.R.O., L.Y. and M.R.A. Methodology: F.T.M., N.G., Y.L.B.P., D.J.E. and H.S. Investigation: F.T.M., Y.L.B.P., N.G., H.S., D.J.E., E.V., X.M., V.O., B.G., S.A., C.P., E.G., M.G.G., J.J.G., J.M.V., B.J.M., G.R.O. and L.Y. Formal analysis: L.B., G.R.O., H.S., Y.L.B.P. and N.G. Writing—original draft: L.B., F.T.M., G.R.O., L.Y. and M.R.A. Writing—review and editing: L.B., F.T.M., G.R.O., M.R.A., L.Y., X.M., E.V., H.S., D.J.E., C.P., N.G., Y.L.B.P., J.M.V., V.O., B.G., S.A., E.G., M.G.G., J.J.G. and B.J.M. Supervision: F.T.M.
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Extended data
Extended Data Fig. 1 Location of the 73 experimental sites surveyed.
Background colors represent aridity index for drylands (areas with an aridity index [mean annual precipitation/potential evapotranspiration] lower than 0.65). Pictures illustrate examples of low grazing plots (left) and high grazing plots (right) across four sites. Photo credits: Matthew Bowker (USA), Juan J. Gaitan (Argentina), Alice Nunes (Portugal), David J. Eldridge (Australia).
Extended Data Fig. 2 Effects of grazing pressure on species richness (A), Shannon diversity index (B), and Pielou evenness index (C).
Mean and 95% confidence intervals are shown (n = 73 sites, each representing a pair of low- and high-grazing plots). Differences between grazing pressures were evaluated using paired t tests (two-sided). For panel A (richness): t = 1.19, P = 0.238, mean difference = −0.99 [95 % CI = −2.64 to 0.67]. For panel B (diversity): t = 1.34, P = 0.183, mean difference = −0.096 [95 % CI = −0.237 to 0.046]. For panel C (evenness): t = 0.90, P = 0.373, mean difference = −0.022 [95 % CI = −0.070 to 0.027]. None of the tests were statistically significant (P > 0.05).
Extended Data Fig. 3 Importance of predictor variables to explain the response of vegetation cover to increasing grazing pressure (resistance) including (A) Shannon’s diversity index, and (B) Pielou’s evenness index.
Importance is based on the sum of Akaike weights of all models where each predictor is present using a multimodel inference approach. MAP = mean annual precipitation, MAT = mean annual temperature, SF = soil fertility, SAC = soil sand content, DIV = Shannon’s diversity index, EVE = Pielou’s evenness index, RWC = relative woody cover, FQ = forage quality, HR = herbivore richness, and LS = dominant livestock species.
Extended Data Fig. 4 Effects of key predictors on the response to increasing grazing pressure (resistance).
Structural equation model showing the relationships among aridity (estimated as 1-Aridity Index), herbivore richness, soil sand content, relative woody cover, latitude, plant species richness, and resistance (estimated as a log response ratio: ln[vegetation cover high grazing pressure/vegetation cover low grazing pressure]). Numbers on arrows are fully standardized path coefficients. Blue and red arrows indicate positive and negative relationships, respectively. Asterisks indicate significance level: ** p<0.01 and *** p<0.001.
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Biancari, L., Oñatibia, G.R., Le Bagousse-Pinguet, Y. et al. Plant diversity enhances ecosystem resistance to increasing grazing pressure in global drylands.
Nat Ecol Evol (2026). https://doi.org/10.1038/s41559-025-02952-9
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DOI: https://doi.org/10.1038/s41559-025-02952-9
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