Multiple abiotic and biotic drivers of aboveground biomass shift with forest stratum

Insights into the underlying ecological mechanisms for diversity-biomass relationships across forest strata (i.e., overstorey and understorey) are crucial to understand the importance of vertical stratification on ecosystem function in natural forests. Yet, it remains unclear how multiple abiotic (i.e., soil nutrients) and biotic (i.e., biodiversity, functional identity and stand structural complexity) factors simultaneously determine aboveground biomass in each individual forest stratum and whole-community. To address this knowledge gap, we disentangled the relative effects of soil nutrients (soil fertility hypothesis), taxonomic, functional trait and evolutionary diversity (niche complementarity hypothesis), stand structural complexity (niche differentiation hypothesis based on tree sizes), and community-weighted mean trait values (mass ratio hypothesis) on aboveground biomass across forest strata and whole-community. We used forest inventory, functional traits and environmental factors datasets from 125 subtropical forest plots in Eastern China. Multiple linear regression models were performed for the selection of best predictors within each biotic group, and structural equation modelling was used to evaluating how multiple abiotic and biotic drivers determine aboveground biomass. In the overstorey, aboveground biomass was positively related to the community-weighted mean of tree height (i.e., functional dominance) and stand density but was negatively related to functional evenness on nutrient-rich soils. In the understorey, aboveground biomass was positively related to phylogenetic species richness and stand-level tree mean diameter (i.e., a proxy for forest growth) but was negatively related to Shannon's species diversity on nutrient-poor soils. Understorey aboveground biomass was also determined by overstorey stand structure and functional dominance through direct and indirect effects via understorey biotic drivers. These results suggest that functional dominance and stand structural complexity are the main biotic drivers of overstorey aboveground biomass. Whereas, functional dominance and stand structural complexity of overstorey, soil nutrients, and niche complementarity among understorey species with the conservative strategy determine understorey aboveground biomass. Whole-community aboveground biomass might be resulting from the superior role of overstorey functional dominance. This study highlights that the niche complementarity, mass ratio and soil nutrients effects are important for driving aboveground biomass, but in different ways across overstorey and understorey strata in natural subtropical forests. We argue that the relative effect of biodiversity loss on aboveground biomass depends critically on the functional and evolutionary identity of the lost species in the specific forest stratum. Better insights can be gained into forest management and biodiversity conservation by considering overstorey and understorey strata separately in complex natural forests.