Tree species mixing affects soil microbial functioning indirectly via root and litter traits and soil parameters in European forests

Plant community composition influences soil microbial communities through plant trait variations that lead to changes in nutrient and organic carbon inputs into the soil by root exudates and plant litter. Although plant litter and living roots are known to influence microbial functioning independently, their relative effects are rarely measured simultaneously in naturally occurring plant communities. Here, we sought to evaluate how forest floor litter and absorptive roots affect broad functions of soil microbial communities, and how this may be influenced by tree species mixing. To do so, forest floor litter, absorptive roots and soil were sampled from mono-specific and three-species mixed stands in four mature, natural forest ecosystems across Europe. The direct effects of tree species mixing, its indirect effects via litter and root traits, and the effects of soil parameters on microbial biomass, catabolic activity and diversity, and denitrification were analysed. Results did not show direct tree mixture effects on the soil microbial parameters we measured but did suggest indirect influences via tree mixture effects on traits of above-ground litter and absorptive roots and soil parameters. Mixed forests composed of any three tree species modified soil microbial functioning by influencing nutrient availability in forest floor litter and root resource acquisition. Tree mixing also modified soil microbial functioning and catabolic diversity by influencing soil fertility and physicochemical properties. Our findings suggest an indirect but present influence of tree species mixing on the activity of heterotrophic soil microbial communities across four different forest ecosystems ranging from Mediterranean to boreal forests. Our study contributes to a better mechanistic understanding of mixed tree species effects on soil microbial functioning via the modification of forest floor litter properties and traits of absorptive roots represented by the tree community beyond simple species number consideration, and potentially via soil properties. A free Plain Language Summary can be found within the Supporting Information of this article.

Automated summary

Plant community composition influences soil microbial communities through plant trait variations, and mixed forests modify soil microbial functioning by influencing nutrient availability in forest floor litter and root resource acquisition. Tree mixing also modifies soil microbial functioning and catabolic diversity by influencing soil fertility and physicochemical properties.