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Aims <p>Shrub encroachment is widespread in temperate grasslands and strongly modifies local resource availability and plant-plant interactions. However, its effects on the foliar elemental composition and coupling of coexisting herbs, and how these effects shift with aridity, remain poorly understood.</p> Methods <p>We conducted a large-scale transect survey across the Inner Mongolian steppe, comparing herbaceous plants growing beneath shrub canopies and in adjacent open patches along an aridity gradient. Foliar macro- and micro-element concentrations were measured for two herbaceous functional groups (grasses <i>vs</i>. forbs). We then developed a multidimensional framework that integrates biogeochemical niche analysis with multi-element network methods to assess how shrubs mediate the foliar elemental composition and elemental coupling of understory herbaceous species.</p> Results <p>The presence of <i>Caragana</i>&#xa0;shrubs&#xa0;generally reduced elemental accumulation in understory herbs, but this inhibitory effect weakened with increasing aridity. Grasses and forbs exhibited distinct responses to shrub presence and aridity, leading to greater biogeochemical niche partitioning between functional groups under more arid conditions. Shrub-induced microhabitat modifications altered the&#xa0;network of inter-element relationships in herbs, increasing node and edge counts but decreasing clustering coefficient and edge density, a shift toward elemental decoupling. Through this network restructuring, shrubs indirectly suppressed herbaceous biomass accumulation.</p> Conclusions <p>Our findings highlight that herbaceous multi-element strategies and coupling patterns are significantly shaped by shrub-mediated effects along aridity gradients. In addition, the multidimensional elemental framework can provide a novel perspective for understanding shrub-herb interactions and their climate responses in temperate grasslands.</p>

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Shrub-mediated shifts in herbaceous foliar elemental composition and coupling along an aridity gradient across the Inner Mongolian steppe

  • 期刊论文

  • 发表日期：

    2026年6月25日

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  • Peng He,
  • Jiang Chen,
  • Josep Peñuelas,
  • Jordi Sardans,
  • Lina Xie,
  • Heyong Liu,
  • Liang Man,
  • Chengcang Ma,
  • Xingguo Han,
  • Yong Jiang,
  • Mai-He Li

摘要

Aims

Shrub encroachment is widespread in temperate grasslands and strongly modifies local resource availability and plant-plant interactions. However, its effects on the foliar elemental composition and coupling of coexisting herbs, and how these effects shift with aridity, remain poorly understood.

Methods

We conducted a large-scale transect survey across the Inner Mongolian steppe, comparing herbaceous plants growing beneath shrub canopies and in adjacent open patches along an aridity gradient. Foliar macro- and micro-element concentrations were measured for two herbaceous functional groups (grasses vs. forbs). We then developed a multidimensional framework that integrates biogeochemical niche analysis with multi-element network methods to assess how shrubs mediate the foliar elemental composition and elemental coupling of understory herbaceous species.

Results

The presence of Caragana shrubs generally reduced elemental accumulation in understory herbs, but this inhibitory effect weakened with increasing aridity. Grasses and forbs exhibited distinct responses to shrub presence and aridity, leading to greater biogeochemical niche partitioning between functional groups under more arid conditions. Shrub-induced microhabitat modifications altered the network of inter-element relationships in herbs, increasing node and edge counts but decreasing clustering coefficient and edge density, a shift toward elemental decoupling. Through this network restructuring, shrubs indirectly suppressed herbaceous biomass accumulation.

Conclusions

Our findings highlight that herbaceous multi-element strategies and coupling patterns are significantly shaped by shrub-mediated effects along aridity gradients. In addition, the multidimensional elemental framework can provide a novel perspective for understanding shrub-herb interactions and their climate responses in temperate grasslands.