1. Leaf functional traits provide important insights into plants' responses to different environments. Leaf traits have been increasingly studied within-species in the last decade, following the growing realisation that neglecting the intra-specific scale can result in misreading plants' response to environmental change. However, while likely to lead to similar pitfalls, within-individual leaf traits are under-researched despite being the scale at which elementary interactions shape ecosystem processes. 2. To address this critical lack of understanding at the local scale, we assessed leaf trait variation in a large biodiversity-ecosystem functioning experiment in subtropical China. We used optical spectroscopy to determine nine morphological and biochemical traits of >5800 leaves from 414 trees representing 14 species. We evaluated the relative importance of the intra-individual level for total leaf trait variation, and the interacting effect of two trait variation's drivers, soil nutrient availability, and a local species richness gradient. 3. Comparing the amount of trait variation at the between-species, between-individuals and intra-individual levels, we found that intra-individual variation accounted on average for >25% of total trait variation. Additionally, intra-individual variation was the most prominent component of intra-specific variation. We found partial support for positive effects of soil nutrient availability and species diversity on intra-individual trait variation, and a strong interdependence of both effects. Contrary to the amplifying effects we expected, trait variation increased with soil nutrient availability at intermediate diversity, but decreased at low and high diversity. 4. Our findings quantify the relevance of intra-individual level for leaf trait variation, and expose a complex interaction between its drivers. In particular, interactive effects of soil nutrient availability and local species diversity on trait variation suggest responses ranging from alleviating competition to enhancing complementarity. Taken together, our work highlights the importance of integrating an intra-individual perspective to understand trait-based mechanisms in biodiversity-ecosystem functioning relationships.
Forest ecosystems are critical for their carbon sequestration potential. Increasing tree diversity has been shown to enhance both forest productivity and litter decomposition. Litter diversity increases litter decomposability by increasing the diversity of substrates offered to decomposers. However, the relative importance of litter decomposability and decomposer community in mediating tree diversity effects on decomposition remains unknown. Moreover, tree diversity modulation of litterfall spatial distribution, and consequently litter decomposition, has rarely been tested. We studied tree diversity effects on leaf litter decomposition and its mediation by the amount of litterfall, litter species richness and decomposability, and soil microorganisms in a large‐scale tree diversity experiment in subtropical China. Furthermore, we examined how litter functional identity and diversity affect leaf litter decomposability. Finally, we tested how leaf functional traits, tree biomass, and forest spatial structure drive the litterfall spatial distribution. We found evidence that tree species richness increased litter decomposition by increasing litter species richness and the amount of litterfall. We showed that soil microorganisms in this subtropical forest perform 84–87% of litter decomposition. Moreover, changes in the amount of litterfall and microbial decomposition explained 19–37% of the decomposition variance. Additionally, up to 20% of the microbial decomposition variance was explained by litter decomposability, while litter decomposability itself was determined by litter functional identity, diversity, and species richness. Tree species richness increased litter species richness and the amount of litterfall (+200% from monoculture to eight‐species neighborhood). We further demonstrated that the amount of species‐specific litterfall increased with increasing tree proximity and biomass, and was modulated by leaf functional traits. These litterfall drivers increased the spatial heterogeneity of litter distribution, and thus litter decomposition. We highlighted multiple biomass‐ and diversity‐mediated effects of tree diversity on ecosystem properties driving forest nutrient cycling. We conclude that considering spatial variability in biotic properties will improve our mechanistic understanding of ecosystem functioning.
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