J Plant Ecol ›› Advance articles     DOI:10.1093/jpe/rtag195

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Divergent effects of litter manipulation and nitrogen enrichment on aboveground- belowground stability are mediated by soil biodiversity in contrasting grassland states

Shuaifei Wang1, 2, Wanjie Chen1, 2, Yongfei Bai3, 4, and Dima Chen1, 2*   

  1. 1 Inner Mongolia Key Laboratory of Grassland Ecology, School of Ecology and Environment, Inner Mongolia University, Hohhot, China
    2 Ministry of Education Key Laboratory of Ecology and Resource Use of the Mongolian Plateau, School of Ecology and Environment, Inner Mongolia University, Hohhot, China
    3 State Key Laboratory of Vegetation and Environmental Change, Institute of Botany, Chinese Academy of Sciences, Beijing, China
    4 University of Chinese Academy of Sciences, Beijing, China
    *Corresponding author:
    Prof. Dima Chen
    School of Ecology and Environment, Inner Mongolia University, Hohhot, China
    Email: chendima@imu.edu.cn
  • Received:2026-01-25 Revised:2026-05-19 Accepted:2026-08-11 Online:2026-08-18 Published:2026-08-18
  • Supported by:
    This study was supported by the National Natural Science Foundation of China (42177272, 32401430, 32201404 32301441 and 32501474), the Natural Science Foundation of Inner Mongolia (2025ZD006 and 2024QN03034), and the Postdoctoral Fellowship Program of China Postdoctoral Science Foundation (GZC20251663).

Abstract: Grassland degradation poses a severe threat to semi-arid ecosystems. However, the interactive effects of litter management and nitrogen (N) enrichment on ecosystem stability remain unclear, especially when jointly considering above- and belowground processes. We coupled N enrichment with a seven-year asymmetric litter manipulation experiment in Inner Mongolian grasslands. Over three consecutive years, we assessed how these drivers influenced plant and soil biodiversity and the stability of above- and belowground multifunctionality. In the undegraded grassland, both litter removal and N enrichment reduced bacterial diversity and belowground functionality, yet whole- system stability remained largely unchanged. In the degraded grassland, however, N enrichment reduced belowground functional stability by simplifying soil communities and amplifying functional variability, whereas litter addition buffered these destabilizing effects, consistent with improved moisture retention. Notably, soil biodiversity, rather than plant diversity, emerged as the primary biotic regulator of stability, but the dominant stabilizing taxa shifted with grassland state: nematode and bacterial diversity underpinned stability in the undegraded and degraded grassland, respectively. Above- and belowground stability were decoupled, with belowground processes exhibiting greater sensitivity to both litter and N manipulation. Consequently, management for stability in semi-arid grasslands should explicitly prioritize soil biodiversity over plant diversity alone. In undegraded systems, minimizing litter removal better safeguards belowground functioning. In degraded systems, strategic litter addition enhances resistance to N enrichment by alleviating moisture limitation but continued N inputs risk long-term destabilization via soil biotic homogenization. Monitoring soil bioindicators (e.g., nematode trophic diversity and functional bacterial groups) can provide early warnings and guide adaptive management strategies.

Key words: Soil biodiversity-stability relationship, Litter manipulation, Ecosystem functionality, Aboveground-belowground decoupling, Functional group compensation, Multi-trophic interactions