Tree species mixing begets admixture of soil microbial communities: Variations along bulk soil, rhizosphere soil and root tissue

Tree species can substantially impact the structure and functioning of soil microbial communities. However, how tree species mixing affects soil microbial communities along different soil compartment niches remains unknown, despite the theoretical (e.g., plant-soil feedback) and practical (e.g., rhizosphere engineering) importance of such knowledge in understanding and managing forest ecosystems. In this study, we analyzed soil samples taken from three microhabitats (bulk soil, rhizosphere soil, and root tissue) of pure conifer species (Pinus mas-soniana) plantations (PP), pure broadleaf species (Castanopsis hystrix) plantations (PC), and mixed plantation forest of the two species (MF) to understand the effects of tree species mixing on core soil microbial communities. We found that the diversity of soil bacterial and fungal communities was not affected by tree species mixing in most cases, but community composition (relative abundance and functional composition) was largely shaped by tree species mixing. Non-metric multidimensional scaling (NMDS) analysis revealed that the core bacterial and fungal communities of MF were between those of PP and PC. This suggests that tree species mixing can lead to a combination of soil microbial communities associated with each of the two tree species, potentially resulting in higher nutrient contents in the bulk soil of MF and lower phosphorus and carbon limitations in both the bulk and rhizosphere soils of MF compared to those of PP and PC. The Mantel test and redundancy analysis (RDA) showed that pH and NO3--N content were the main environmental variables influencing soil microbial communities. Overall, the effects of tree species mixing varied markedly with microhabitat and season, with rhizosphere soil having the highest microbial diversity and network complexity and root tissue having the lowest. These results suggest that the mixing effects of tree species are more pronounced in the rhizosphere and dry season, on which silvicultural practices should focus.