Stand spatial structure and microbial diversity are key drivers of soil multifunctionality during secondary succession in degraded karst forests

Studying the relationship between biodiversity and ecosystem multifunctionality (the ability of ecosystems to provide multiple ecosystem functions) (BEMF) is a current hotspot in ecology research. Previous studies on BEMF emphasized the role of plant and microbial diversity but rarely mention stand spatial structure. To investigate the effect of stand spatial structure on BEMF, this study established 30 forest dynamic plots in three natural restoration stages (shrubbery, secondary growth forest, and old -growth forest) in Maolan National Nature Reserve, Guizhou province, China. A positive response in soil multifunctionality (SMF), plant species diversity, stand spatial structure, and fungal beta diversity ( p < 0.05) followed natural restoration. However, bacterial beta diversity showed a negative response ( p < 0.05), while microbial alpha diversity remained unchanged ( p > 0.05). These results based on a structural equation model showed that plant species diversity had no direct or indirect effect on SMF, soil microbial diversity was the only direct driver of SMF, and stand spatial structure indirectly affected SMF through soil microbial diversity. The random forest model showed that soil microbial beta diversity and the Shannon -Wiener index of the diameter at breast height for woody plant species were the optimal variables to characterize SMF and soil microbial diversity, respectively. These results suggested that natural restoration promoted SMF, and microbial diversity had a direct positive effect on SMF. In the meantime, stand spatial structure had a significant indirect effect on SMF, while plant species diversity did not. Future work on degraded karst forest restoration should direct more attention to the role of the stand spatial structure and emphasize the importance of biodiversity.