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Grassland degradation alters plant and soil biodiversity–multifunctionality relationships

Nature Plants volume 11, pages 2487–2497 (2025)Cite this article

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Abstract

Biodiversity is known to promote ecosystem multifunctionality (EMF), but how grassland degradation influences the relationship between biodiversity and EMF remains unclear. Here, using paired observations at 44 sites (a total of 792 sampling quadrats) along a 2,600 km transect, we test how moderate grassland degradation influences 20 surrogates of ecosystem functions, EMF, plant richness, soil bacterial, fungal and protist richness, and biodiversity–EMF relationships in Tibetan alpine grasslands. Our results reveal significant declines in individual ecosystem functions and EMF with moderate grassland degradation. By contrast, both plant richness and integrated soil biodiversity exhibit significant increases. The structural equation models analyses show that following degradation, the effect of soil biodiversity on EMF strengthens, whereas that of plant richness weakens. These findings offer large-scale empirical evidence that moderate grassland degradation can amplify both soil biodiversity and its functional importance, emphasizing the key role of below-ground biodiversity in supporting ecosystem functioning in degraded grasslands.

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Fig. 1: Sampling sites and vegetation community of non-degraded and degraded grasslands.
Fig. 2: Responses of ecosystem functions to grassland degradation.
Fig. 3: Comparisons of plant richness and soil biodiversity in non-degraded and degraded grasslands.
Fig. 4: SEMs illustrating the direct and indirect effects of abiotic and biotic factors on EMF.

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Data availability

Data used in this study are available from the figshare data repository https://doi.org/10.6084/m9.figshare.30119233 (ref. 93). The sequence data generated in this study have been deposited in the Genome Sequence Archive in National Genomics Data Center, China National Center for Bioinformation https://ngdc.cncb.ac.cn/bioproject/browse/PRJCA046361 (ref. 94). Source data are provided with this paper.

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Acknowledgements

The authors thank Z. Tang (Institute of Ecology, College of Urban and Environmental Science, Peking University), B. Schmid (Remote Sensing Laboratories, Department of Geography, University of Zurich) and N. A. Pichon (Swiss Federal Institute for Forest, Snow and Landscape Research WSL, Birmensdorf, Switzerland) for their comments and suggestions on the manuscript. This study was financially supported by the Strategic Priority Research Program of Chinese Academy of Sciences (XDA26020201, Y.Y.), the National Natural Science Foundation of China (32588202, Y.Y.; 32425004, Y.Y.; 32301501, X.G.) and the New Cornerstone Science Foundation through the XPLORER PRIZE (Y.Y.).

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Author notes

  1. These authors contributed equally: Xiaoxia Gao, Dianye Zhang.

Authors and Affiliations

  1. State Key Laboratory of Forage Breeding-by-Design and Utilization; Key Laboratory of Vegetation and Environmental Change, Institute of Botany, Chinese Academy of Sciences, Beijing, China

    Xiaoxia Gao, Dianye Zhang, Yunfeng Peng, Yaping Niu, Shiting Yao, Zan Wu, Qinlu Li, Lina Zhou, Yang Liu, Xuning Liu, Bin Wei, Shuqi Qin, Yutong Song, Luyao Kang & Yuanhe Yang

  2. China National Botanical Garden, Beijing, China

    Xiaoxia Gao, Dianye Zhang, Yunfeng Peng, Yaping Niu, Shiting Yao, Zan Wu, Qinlu Li, Lina Zhou, Yang Liu, Xuning Liu, Bin Wei, Shuqi Qin, Yutong Song, Luyao Kang & Yuanhe Yang

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

    Josep Peñuelas

  4. CREAF, Cerdanyola del Vallès, Catalonia, Spain

    Josep Peñuelas

  5. Ecology and Biodiversity Group, Department of Biology, Utrecht University, Utrecht, The Netherlands

    Yann Hautier

  6. Theoretical and Experimental Ecology Station, CNRS, Moulis, France

    Michel Loreau

  7. Institute of Ecology, College of Urban and Environmental Science, and State Key Laboratory for Vegetation Structure, Function and Construction (VegLab), Peking University, Beijing, China

    Michel Loreau & Shaopeng Wang

  8. University of Chinese Academy of Sciences, Beijing, China

    Yaping Niu, Bin Wei, Yutong Song & Yuanhe Yang

  9. College of Agriculture and Animal Husbandry, Qinghai University, Xining, China

    Shiting Yao

  10. School of Biological Sciences, Georgia Institute of Technology, Atlanta, GA, USA

    Lin Jiang

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Contributions

Y.Y. conceived the idea. Y.Y., D.Z. and X.G. designed the research. Y.P. and Y.Y. designed the field sampling. Y.P., X.G., Q.L., Z.W., Y.N., S.Y. and X.L. performed field samplings. X.G., Z.W., L.Z., Y.N., Y.L. and S.Y. performed the experiments. X.G. and D.Z. analysed the data. X.G., D.Z. and Y.Y. wrote the paper with inputs from J.P., Y.H., M.L., S.W., L.J., B.W., S.Q., Y.S. and L.K.

Corresponding author

Correspondence to Yuanhe Yang.

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Nature Plants thanks Eric Allan, Ling Wang and the other, anonymous, reviewer(s) for their contribution to the peer review of this work.

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Extended data

Extended Data Fig. 1 Schematic diagram of the sampling design.

At a specific site, three blocks were selected, and within each block a 30 × 30 m plot was established in both non-degraded and degraded grasslands. Within each plot, three 1 × 1 m quadrats were randomly chosen to conduct vegetation surveys and collect plant and soil samples (9 soil cores at 0-10 cm depth). Altitude map derived from the Shuttle Radar Topography Mission (SRTM)-Digital Elevation Model (DEML)95. Drone images of contrasting non-degraded and degraded grasslands show that plant productivity and the proportion of grasses and sedges (for example, S. purpurea and K. humilis) decreases while the proportion of forbs (for example, L. nanum and S. chamaejasme) increases upon degradation. Photos were captured by Qinlu Li in Babao Town (37.86°N, 100.44°E, altitude 3663 m), July 2021. ND, non-degraded grasslands; DG, degraded grasslands.

Extended Data Fig. 2 Comparisons of ecosystem multifunctionality (EMF) between non-degraded (ND) and degraded grasslands (DG).

a, Averaged approach; b, multiple-threshold approach; c, d, hill-number based approaches when q = 1 (c) and q = 2 (d). The open circles shown are the site-level values. The ends of a box denote the 25th and 75th quartiles, and whiskers extend to 1.5 times the standard deviation (n = 44 independent sites). The horizontal line within each box represents the median value. Linear mixed models with a correction for environmental covariates were employed to conduct statistical comparisons between non-degraded and degraded grasslands across the 44 sites. Climate and soil factors were treated as covariates, site and grassland type (alpine steppe, alpine meadow or swamp meadow) were treated as random factors. All the reported two-sided P values were not corrected for multiple comparisons, and detailed F-statistics were presented in Supplementary table 1.

Source data

Extended Data Fig. 3 Comparisons of plant community in non-degraded (ND) and degraded grasslands (DG).

a, Cover of grasses and sedges; b, cover of forbs. The dashed lines indicate median values; solid lines represent the values for each site (n = 44 independent sites). Linear mixed models with a correction for environmental covariates were employed to conduct statistical comparisons between non-degraded and degraded grasslands across the 44 sites (cover of grasses and sedges: F1,46.72 = 276.01, P = 3.10 × 10−21; cover of forbs: F1, 44.72 = 52.66, P = 4.40 × 10−9; the two-sided P values were not corrected for multiple comparisons). Climate and soil factors were treated as covariates, site and grassland type (alpine steppe, alpine meadow or swamp meadow) were treated as random factors.

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Extended Data Fig. 4 Differences in the relative abundance of copiotrophic and oligotrophic bacteria between non-degraded (ND) and degraded grasslands (DG).

a, Copiotrophic bacteria; b, oligotrophic bacteria. The ends of a box denote the 25th and 75th quartiles, and whiskers extend to 1.5 times the standard deviation (n = 44 independent sites). The horizontal line within each box represents the median value. The open circles represent the values for each site. Linear mixed models with a correction for environmental covariates were employed to conduct statistical comparisons between non-degraded and degraded grasslands across the 44 sites (copiotrophic bacteria: F1,51.14 = 5.69, P = 0.0209; oligotrophic bacteria: F1, 51.01 = 6.46, P = 0.0141; the two-sided P values were not corrected for multiple comparisons). Climate and soil factors were treated as covariates, site and grassland type (alpine steppe, alpine meadow or swamp meadow) were treated as random factors. The demarcations of copiotrophic and oligotrophic bacteria were as follows31, oligotrophs: Acidobacteriota, Actinobacteriota, Planctomycetota, Chloroflexi, Verrucomicrobiota, Cyanobacteria; copiotrophs: Firmicutes, Gemmatimonadota, Bacteroidota, Proteobacteria. Copiotrophic bacteria are generally fast-slowing and resource acquisitive, while oligotrophic bacteria tend to be slow-growing and resource conservative31.

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Extended Data Fig. 5 Differences in cover of clonal and non-clonal plants in non-degraded (ND) and degraded grasslands (DG).

a, Clonal plants; b, non-clonal plants. The horizontal line within each box represents the median value; the open circles represent the values for each site (n = 44 independent sites). Linear mixed models with a correction for environmental covariates were employed to conduct statistical comparisons between non-degraded and degraded grasslands across the 44 sites (cover of clonal plants: F1,47.77 = 265.38, P = 3.89 × 10−21; cover of non-clonal plants: F1, 44.67 = 39.42, P = 1.24 × 10−7; the two-sided P values were not corrected for multiple comparisons). Climate and soil factors were treated as covariates, site and grassland type (alpine steppe, alpine meadow or swamp meadow) were treated as random factors.

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Extended Data Fig. 6 Relationships between soil biodiversity and plant richness in non-degraded (ND) and degraded grasslands (DG).

Soil biodiversity was calculated by averaging biodiversity across the three taxa (soil bacteria, fungi and protists) after standardization33,73. Linear mixed models (n = 44 independent sites) were used to determine the linkages between soil biodiversity and plant richness, with grassland type (alpine steppe, alpine meadow or swamp meadow) being treated as the random factor (non-degraded grasslands: F1,42 = 27.68, P = 4.55 × 10−6; degraded grasslands: F1,41.71 = 24.97, P = 1.09 × 10−5; the two-sided P values were not corrected for multiple comparisons). Shaded areas are the 95% confidence intervals; the open circles are the values for each site. Soil biodiversity still exhibited significant linkages with plant richness when considering the environmental covariates (climate and soil factors).

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Extended Data Fig. 7 Differences in cover of poisonous weeds in non-degraded (ND) and degraded grasslands (DG).

The ends of a box denote the 25th and 75th quartiles, and whiskers extend to 1.5 times the standard deviation. The open circles represent the values for each site (n = 44 independent sites). Linear mixed models with a correction for environmental covariates were employed to conduct statistical comparisons between non-degraded and degraded grasslands across the 44 sites (F1,48.06 = 34.34, P = 4.08 × 10−7; the two-sided P value was not corrected for multiple comparisons). Climate and soil factors were treated as covariates, site and grassland type (alpine steppe, alpine meadow or swamp meadow) were treated as random factors.

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Extended Data Fig. 8 Comparisons of plant and microbial community evenness between non-degraded (ND) and degraded grasslands (DG).

a, Plant evenness; b, bacteria evenness; c, fungi evenness; d, protist evenness. The ends of a box denote the 25th and 75th quartiles, and whiskers extend to 1.5 times the standard deviation (n = 44 independent sites). The horizontal line within each box represents the median value. The open circles represent the values for each site. Linear mixed models with a correction for environmental covariates were employed to conduct statistical comparisons between non-degraded and degraded grasslands across the 44 sites (plant: F1,48.52 = 143.60, P = 4.15 × 10−16; bacteria: F1, 42.24 = 11.05, P = 0.0018; fungi: F1, 48.86 = 4.45, P = 0.0400; protist: F1, 50.95 = 0.25, P = 0.6211; the two-sided P values were not corrected for multiple comparisons). Climate and soil factors were treated as covariates, site and grassland type (alpine steppe, alpine meadow or swamp meadow) were treated as random factors.

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Extended Data Fig. 9 Comparisons of microbial interspecific cohesion between non-degraded (ND) and degraded grasslands (DG).

a, Microbial positive cohesion; b, microbial negative cohesion. The ends of a box denote the 25th and 75th quartiles, and whiskers extend to 1.5 times the standard deviation (n = 44 independent sites). The open circles represent the values for each site, the horizontal line within each box represents the median value. Linear mixed models with a correction for environmental covariates were employed to conduct statistical comparisons between non-degraded and degraded grasslands across the 44 sites (positive cohesion: F1,49.67 = 24.93, P = 7.71 × 10−6; negative cohesion: F1, 41.20 = 0.11, P = 0.7464; the two-sided P values were not corrected for multiple comparisons). Climate and soil factors were treated as covariates, site and grassland type (alpine steppe, alpine meadow or swamp meadow) were treated as random factors.

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Extended Data Fig. 10 Comparisons of microbial community composition between non-degraded (ND) and degraded grasslands (DG).

a, Bacterial community composition; b, fungal community composition; c, protist community composition. Data shown in this figure are the relative abundance of microbes at the level of phylum. Community composition of bacteria, fungi and protist all exhibited significant changes following degradation according to permutational multivariable analysis of variance (all P < 0.05). Linear mixed models with a correction for environmental covariates were employed to conduct statistical comparisons of the relative abundance of each microbial group between non-degraded and degraded grasslands across the 44 sites. Climate and soil factors were treated as covariates, site and grassland type (alpine steppe, alpine meadow or swamp meadow) were treated as random factors. The plus (+) and minus (-) signs denote that microbial taxa significantly increase or decrease, respectively, following grassland degradation. All the reported two-sided P values were not corrected for multiple comparisons.

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Gao, X., Zhang, D., Peng, Y. et al. Grassland degradation alters plant and soil biodiversity–multifunctionality relationships. Nat. Plants 11, 2487–2497 (2025). https://doi.org/10.1038/s41477-025-02147-x

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