Diversity-biomass relationships are shaped by tree mycorrhizal associations and stand structural diversity at different spatial scales

Diversity-biomass relationships (DBRs) in terrestrial ecosystems tend to vary across spatial scales, but, particularly in hyperdiverse forests, the mechanisms driving these trends remain uncertain. Until now, few have simultaneously investigated the connections between tree species diversity, stand structural diversity, mycorrhizal associations, and ecosystem functioning. In addition, DBRs have only been studied at limited spatial scales, with limited focus on the direct and indirect effects of environmental factors. We addressed these research gaps using a 30-ha forest dynamics plot located in Pu'er City, Southwest China. Through piecewise structural equation models, we quantified the direct effects of tree species diversity (α, β, γ), stand structural diversity, mycorrhizal associations (AM, EcM), and the environmental factors (soil fertility and topography), as well as the indirect effects of the environmental factors on aboveground tree biomass across spatial scales ranging from 400 to 230,400 ​m2. We hypothesized that complex interactions among these factors underpin the variation in DBRs in natural ecosystems across spatial scales. Our results showed that environmental conditions indirectly affected the tree biomass via changes in tree species diversity, and these effects became stronger as the spatial scale increased. At small to moderate spatial scales, environmental factors were more predictive of tree biomass than tree species diversity (or its components); the effects of stand structural diversity on biomass also gradually increased with spatial scale. Conversely, from the intermediate to the largest spatial scales, mycorrhizal associations gradually became the best predictors of DBR dynamics. Our research offers novel empirical evidence demonstrating the importance of environmental conditions, structural diversity, and mycorrhizal associations in shaping cross-scale DBRs. Future comprehensive studies should consider these factors to assess the mechanisms shaping scale-dependent DBRs in complex natural ecosystems.