Abstract
Grassland canopy height is one of the most important traits for determining plant diversity and community structure, directly affecting the resource use efficiency of livestock in grassland ecosystems. However, broad-scale changes in grassland canopy height are seldom reported due to the complex effects of species aggregation on both interspecific and intraspecific structures. Here, we decouple grassland aboveground biomass into vertical and horizontal allocations, thereby offering a pathway to mirror changes in grassland canopy height. Grassland aboveground biomass is estimated using a machine learning algorithm by fusing climatic factors, satellite-driving metrics, and 8-year consecutive ground-truth surveys; the horizontal allocation of grassland aboveground biomass is derived from optimized linear spectral mixture analysis. We find that changes in horizontal biomass allocation primarily accounted for increases in Chinese grassland aboveground biomass from 2001 to 2022, resulting in a significant decline in grassland canopy height. The decline in grassland canopy height is shaped by reduced radiation and, more importantly, by the combined effects of warming and grazing, while also being related to variations in plant diversity. The dwarfing grassland community with declining canopy height may increase the impact of livestock disturbances, thus diminishing the resistance of grassland ecosystems to climate fluctuations.
Similar content being viewed by others
Data availability
The MOD09A1 data can be accessed at https://doi.org/10.5067/MODIS/MOD09A1.061, and the MOD15A2H data from https://doi.org/10.5067/MODIS/MOD15A2H.061. GEDI Level 2 A and Level 2B products are available at https://doi.org/10.5067/GEDI/GEDI02_A.002 and https://doi.org/10.5067/GEDI/GEDI02_B.002, respectively. The ERA5-Land dataset can be downloaded at https://cds.climate.copernicus.eu/. The CO2 data can be accessed at https://doi.org/10.24381/a90c7e33. The field measurements of aboveground biomass, fractional vegetation cover, canopy height, and species richness are collected from the National Inventory of Grassland Resources by the National Forestry and Grassland Administration of China. The grazing intensity data are available at https://doi.org/10.6084/m9.figshare.26195684. The data for continuous interannual measurements of aboveground biomass and fractional vegetation cover are primarily obtained from the China Ecosystem Research Network (www.nesdc.org.cn). The gridded dataset of grassland canopy height across China (2001–2022) generated in this study is available in the Zenodo repository [https://doi.org/10.5281/zenodo.18454934]49.
Code availability
The codes in this study are available in the Zenodo repository [https://doi.org/10.5281/zenodo.18453938]50.
References
Lang, N., Jetz, W., Schindler, K. & Wegner, J. D. A high-resolution canopy height model of the Earth. Nat. Ecol. Evol. 7, 1778–1789 (2023).
Falster, D. S. & Westoby, M. Plant height and evolutionary games. Trends Ecol. Evol. 18, 337–343 (2003).
Moles, A. T. et al. Global patterns in plant height. J. Ecol. 97, 923–932 (2009).
Quan, Q. et al. Plant height as an indicator for alpine carbon sequestration and ecosystem response to warming. Nat. Plants 1, 11 (2024).
Li, D. Grassland plants under warming and precipitation change. Plant Growth and Regulation: Alterations to Sustain Unfavorable Conditions 1 (2018).
Arft, A. et al. Responses of tundra plants to experimental warming: meta-analysis of the international tundra experiment. Ecol. Monogr. 69, 491–511 (1999).
Yan, R. et al. Impacts of differing grazing rates on canopy structure and species composition in Hulunber meadow steppe. Rangel. Ecol. Manag. 68, 54–64 (2015).
Li, H. et al. A machine learning scheme for estimating fine-resolution grassland aboveground biomass over China with Sentinel-1/2 satellite images. Remote Sens. Environ. 311, 114317 (2024).
Li, F., Chen, W., Zeng, Y., Zhao, Q. & Wu, B. Improving estimates of grassland fractional vegetation cover based on a pixel dichotomy model: A case study in Inner Mongolia. China Remote Sens. 6, 4705–4722 (2014).
Camps-Valls, G. et al. A unified vegetation index for quantifying the terrestrial biosphere. Sci. Adv. 7, eabc7447 (2021).
Huete, A. R., Jackson, R. D. & Post, D. F. Spectral response of a plant canopy with different soil backgrounds. Remote Sens. Environ. 17, 37–53 (1985).
De Conto, T., Armston, J. & Dubayah, R. Characterizing the structural complexity of the Earth’s forests with spaceborne lidar. Nat. Commun. 15, 8116 (2024).
Bu, J. & Xiao, J. Upscaling eddy covariance measurements of carbon and water fluxes to the continental scale by incorporating GEDI-derived canopy structural complexity metrics. Remote Sens. Environ. 329, 114930 (2025).
Malambo, L. & Popescu, S. C. Assessing the agreement of ICESat-2 terrain and canopy height with airborne lidar over US ecozones. Remote Sens. Environ. 266, 112711 (2021).
Xu, C. et al. Correction of UAV LiDAR-derived grassland canopy height based on scan angle. Front. Plant Sci. 14, 1108109 (2023).
Lang, N., Schindler, K. & Wegner, J. D. Country-wide high-resolution vegetation height mapping with Sentinel-2. Remote Sens. Environ. 233, 111347 (2019).
Meilhac, J., Deschamps, L., Maire, V., Flajoulot, S. & Litrico, I. Both selection and plasticity drive niche differentiation in experimental grasslands. Nat. Plants 6, 28–33 (2019).
Pérez-Ramos, I. M., Matías, L., Gómez-Aparicio, L. & Godoy, Ó. Functional traits and phenotypic plasticity modulate species coexistence across contrasting climatic conditions. Nat. Commun. 10, 2555 (2019).
Poorter, H. & Sack, L. Pitfalls and possibilities in the analysis of biomass allocation patterns in plants. Front. Plant Sci. 3, 259 (2012).
Gruntman, M., Groß, D., Májeková, M. & Tielbörger, K. Decision-making in plants under competition. Nat. Commun. 8, 2235 (2017).
McConnaughay, K. & Coleman, J. Biomass allocation in plants: ontogeny or optimality? A test along three resource gradients. Ecology 80, 2581–2593 (1999).
Liang, M., Liang, C., Hautier, Y., Wilcox, K. R. & Wang, S. Grazing-induced biodiversity loss impairs grassland ecosystem stability at multiple scales. Ecol. Lett. 24, 2054–2064 (2021).
Chen, C. et al. China and India lead in the greening of the world through land-use management. Nat. Sustain 2, 122–129 (2019).
MacDougall, A. S. et al. Widening global variability in grassland biomass since the 1980s. Nat. Ecol. Evol. 8, 1877–1888 (2024).
Luo, W. et al. Plant traits and soil fertility mediate productivity losses under extreme drought in C3 grasslands. Ecology 102, e03465 (2021).
Díaz, S. et al. Plant trait responses to grazing – a global synthesis. Glob. Change Biol. 13, 313–341 (2007).
Gross, N. et al. Unforeseen plant phenotypic diversity in a dry and grazed world. Nature 632, 808–814 (2024).
Díaz, S., Noy-Meir, I. & Cabido, M. Can grazing response of herbaceous plants be predicted from simple vegetative traits?. J. Appl. Ecol. 38, 497–508 (2001).
Zhang, M. et al. Experimental impacts of grazing on grassland biodiversity and function are explained by aridity. Nat. Commun. 14, 5040 (2023).
Ainsworth, E. A. & Long, S. P. What have we learned from 15 years of free-air CO2 enrichment (FACE)? A meta-analytic review of the responses of photosynthesis, canopy properties and plant production to rising CO2. N. Phytol. 165, 351–372 (2005).
Song, J. et al. Elevated CO2 does not stimulate carbon sink in a semi-arid grassland. Ecol. Lett. 22, 458–468 (2019).
Feldman, A. F. et al. Large global-scale vegetation sensitivity to daily rainfall variability. Nature 636, 380–384 (2024).
Sloat, L. L. et al. Increasing importance of precipitation variability on global livestock grazing lands. Nat. Clim. Change 8, 214–218 (2018).
Wang, Y. et al. Grassland changes and adaptive management on the Qinghai–Tibetan Plateau. Nat. Rev. Earth Environ. 3, 668–683 (2022).
Yang, H. et al. Community structure and composition in response to climate change in a temperate steppe: steppe community responses to climate change. Glob. Change Biol. 17, 452–465 (2011).
Maestre, F. T. et al. Grazing and ecosystem service delivery in global drylands. Science 378, 915–920 (2022).
Yu, Q. et al. Contrasting drought sensitivity of Eurasian and North American grasslands. Nature 639, 114–118 (2025).
Isbell, F. et al. Biodiversity increases the resistance of ecosystem productivity to climate extremes. Nature 526, 574–577 (2015).
Tilman, D., Reich, P. B. & Knops, J. M. H. Biodiversity and ecosystem stability in a decade-long grassland experiment. Nature 441, 629–632 (2006).
Li, L., Chen, J., Han, X., Zhang, W. & Shao, C. Grassland Ecosystems of China: A Synthesis and Resume. 2 (Springer Singapore, Singapore, 2020).
Hufkens, K. et al. Productivity of North American grasslands is increased under future climate scenarios despite rising aridity. Nat. Clim. Change 6, 710–714 (2016).
Bardgett, R. D. et al. Combatting global grassland degradation. Nat. Rev. Earth Environ. 2, 720–735 (2021).
Agustí-Panareda, A. et al. Technical note: the CAMS greenhouse gas reanalysis from 2003 to 2020. Atmos. Chem. Phys. 23, 3829–3859 (2023).
Wang, D. et al. A long-term high-resolution dataset of grasslands grazing intensity in China. Sci. Data 11, 1194 (2024).
Wang, G. et al. Simulating the spatiotemporal variations in aboveground biomass in Inner Mongolian grasslands under environmental changes. Atmos. Chem. Phys. 21, 3059–3071 (2021).
Mohammed, R., Rawashdeh, J. & Abdullah, M. Machine learning with oversampling and undersampling techniques: overview study and experimental results. in 2020 11th international conference on information and communication systems (ICICS). 243–248 (IEEE, 2020).
Raftery, A. E., Madigan, D. & Hoeting, J. A. Bayesian model averaging for linear regression models. J. Am. Stat. Assoc. 92, 179–191 (1997).
Lefcheck, J. S. piecewiseSEM: Piecewise structural equation modelling in r for ecology, evolution, and systematics. Methods Ecol. Evol. 7, 573–579 (2016).
Li, H. et al. Gridded dataset of grassland canopy height across China (2001–2022). Zenodo https://doi.org/10.5281/zenodo.18454934 (2026).
Li, H. et al. Codes for Declining grassland canopy height in China under asymmetric biomass allocation. Zenodo https://doi.org/10.5281/zenodo.18453938 (2026).
Acknowledgments
This project was supported by the National Natural Science Foundation of China (42471426). F.L. was supported by the Chinese Universities Scientific Fund (2025TC044) and the Science and Technology Program of Inner Mongolia Autonomous Region (2025YFDZ0055).
Author information
Authors and Affiliations
Contributions
H.L. conducted model simulations, analysed the results, and drafted the text. X.H. conducted driver analysis and drafted the text. F.L. designed the study, analysed the results, and edited the text. Y.Z. contributed to the improvement of the study design and the drafting of the text. K.L. and J.W. (Jie Wang) revised the manuscript. J.W. (Jiating Wang) contributed to the collection of ground-truth data.
Corresponding authors
Ethics declarations
Competing interests
The authors declare no competing interests.
Peer review
Peer review information
Nature Communications thanks Nasem Badreldin and Tatiana Kuplich 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.
Rights and permissions
Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, 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 changes were made. 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/4.0/.
About this article
Cite this article
Li, H., Hu, X., Li, F. et al. Declining grassland canopy height in China under asymmetric biomass allocation. Nat Commun (2026). https://doi.org/10.1038/s41467-026-70275-9
Received:
Accepted:
Published:
DOI: https://doi.org/10.1038/s41467-026-70275-9


