Moderate ground surface heterogeneity sustains higher aboveground biomass by enhancing structural diversity in fragmented forests

doi:10.1093/jpe/rtag267

J Plant Ecol ›› Advance articles DOI:10.1093/jpe/rtag267

Moderate ground surface heterogeneity sustains higher aboveground biomass by enhancing structural diversity in fragmented forests

Fulin Wang¹, Xu Han¹, Xihong Yang¹, Min Jiao¹, Xiaodong Li¹, Xianqi Tang¹, Jingjing You¹, Yutong Xiao¹, Mingjuan Yu¹, Mingming Zhang¹, Pan Wu², Jun Mu⁴, Yuejun He^1,2,3,*

1. Forestry College, Research Center of Forest Ecology, Guizhou University, Guiyang 550025, China;
2. Key Laboratory of Karst Georesources and Environment, Ministry of Education, College of Resources and Environmental Engineering, Guizhou University, Guiyang 550025, China;
3. Guizhou Xishui National Nature Reserve Administration Bureau, Xishui 564600, China;
4. Guizhou Guiyang Urban Ecosystem Observation and Research Station, National Forestry and Grassland Administration, Guiyang 550025, China
^*Corresponding authors:
Yuejun He,E-mail: hyj1358@163.com

Received:2025-11-24 Revised:2026-09-03 Accepted:2026-10-14 Published:2026-11-06
Supported by:
This study was supported by the National Natural Science Foundation of China (NSFC: 32260268); the Key Program of Guizhou Provincial Basic Research Foundation (No. ZD [2025] 072); the project of Central Funding for National Nature Reserves (CFNR-2025-XS-R01； CFNR-2026-XS-R01); and the National Key Research and Development Project of China [2025YFC2609500].

Abstract

Abstract: Forests are key components of global ecosystems, regulating nutrient cycling and energy flow. In karst landscapes, pronounced ground surface heterogeneity generates highly uneven resource distributions, thereby shaping forest community structure and biomass accumulation. Although multiple dimensions of biodiversity are known to influence aboveground biomass (AGB), how ground surface heterogeneity mediates biodiversity and AGB across fine spatial scales in fragmented forests remains unclear. In this study, we quantified ground surface heterogeneity using an entropy-based index across 212 subplots nested within 53 plots in a subtropical karst forest. By integrating multivariate biodiversity metrics and community-weighted traits, we evaluated the pathways through which ground surface heterogeneity regulates biodiversity and AGB. Our analyses revealed that structural diversity was the strongest predictor of AGB, whereas functional composition also contributed positively, suggesting that both selection and complementarity effects were involved. Moreover, at the plot scale, biodiversity and AGB exhibited unimodal responses to ground surface heterogeneity, indicating that AGB peaked at intermediate levels of heterogeneity, potentially reflecting enhanced structural differentiation and resource complementarity. Taken together, these findings demonstrate a clear hierarchical pathway that ground surface heterogeneity drives variations in biodiversity, which subsequently translates into enhanced stand structural diversity, ultimately maintaining higher AGB. This study underscores the central role of moderate ground surface heterogeneity in sustaining forest structural diversity and biomass accumulation, highlighting the importance of integrating habitat variability and structural metrics into forest management and restoration strategies in fragmented ecosystems.

This study investigated the relationships among ground surface heterogeneity, biodiversity, and aboveground biomass in karst forests in southwestern China. Moderate heterogeneity was associated with higher biomass, with structural diversity emerging as an important predictor, highlighting its role in understanding biomass patterns in fragmented forests.

Key words: aboveground biomass, ground surface heterogeneity, structural diversity, functional composition, intermediate heterogeneity hypothesis, fragmented forests

Fulin Wang, Xu Han, Xihong Yang, Min Jiao, Xiaodong Li, Xianqi Tang, Jingjing You, Yutong Xiao, Mingjuan Yu, Mingming Zhang, Pan Wu, Jun Mu, Yuejun He. Moderate ground surface heterogeneity sustains higher aboveground biomass by enhancing structural diversity in fragmented forests[J]. J Plant Ecol, DOI: 10.1093/jpe/rtag267.

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Moderate ground surface heterogeneity sustains higher aboveground biomass by enhancing structural diversity in fragmented forests

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