Temporal stability of aboveground biomass is governed by species asynchrony in temperate forests

Understanding the effects of plant species diversity and trait composition on aboveground biomass is a central focus of ecology and has important implications for biodiversity conservation. However, the simultaneous direct and indirect effects of soil nutrients, species asynchrony, functional trait diversity, and trait composition for explaining the community temporal stability of aboveground biomass remain underrepresented in natural forests. Here, we hypothesized that species asynchrony relative to soil nutrients, functional trait diversity, and trait composition plays a central role in stabilizing the community temporal stability of natural forests. We tested this hypothesis using a structural equation model based on 10-year continuous monitoring data (i.e., three-time repeated forest inventories) in both second-growth and old-growth temperate forests in northeast China. Our results showed that the community temporal stability of aboveground biomass was driven by a strong direct positive effect of species asynchrony in both second-growth and old-growth temperate forests, whereas functional trait diversity and composition (i.e. community-weighted mean of leaf nitrogen content) were of additional importance in an old-growth forest only. Functional trait diversity decreased community-weighted mean of leaf nitrogen content in an old-growth forest, whereas this relationship was non-significant in a second-growth forest. Soil nutrients had non-significant effects on the community temporal stability of both second-growth and old-growth forests. Species asynchrony was the direct determinant of the community temporal stability of aboveground biomass in temperate forests. The direct effect of species asynchrony increased with forest succession, implying that temporal niche differentiation and facilitation increase over time. This study suggests that managing forests with mixtures of both early and late successional species or shade intolerant and tolerant species, not only species diversity, is important for maintaining forest stability in a changing environment. We argue that the species asynchrony effect is crucial to understand the underlying ecological mechanisms for a diversity-biomass relationship in natural forests.