Tree species diversity increases fine root productivity through increased soil volume filling

Although fine roots (< 2 mm in diameter) account for a major share of the production of terrestrial ecosystems, diversity effects on fine root productivity and their mechanisms remain unclear. We hypothesized that: (i) fine root productivity increases with tree species diversity, (ii) higher fine root productivity is a result of greater soil volume filling due to species-specific patterns of root placement and proliferation, and (iii) differences in fine root productivity and soil volume filling associated with tree species diversity are more pronounced in summer when plants are physiologically active and demand for water and nutrients is at its greatest. We investigated the effects of tree species diversity on fine root productivity and soil volume filling of boreal forest stands that have grown naturally for 85 years on similar sites. Annual fine root production was 19-83% higher in evenly mixed- than single-species-dominated stands, and increased with tree species evenness, but not tree species richness. Fine root biomass was higher in evenly mixed- than single-species-dominated stands in summer months, but not in spring or fall. Higher fine root productivity in evenly mixed- than single-species-dominated stands was realized by filling more soil volume horizontally and vertically in the forest floor in the mixtures of deep- and shallow-rooted species vs. the deeper mineral soil in the mixtures of deep-rooted species. Synthesis. Our results provide some of the first direct evidence for below-ground species complementarity in heterogeneous natural forests, by demonstrating that tree species evenness increases fine root productivity by filling/exploiting the soil environment more completely in space and time, driven by differences in the inherent rooting traits of the component species and variations of root growth within species.