Community-aggregated plant traits interact with soil nutrient heterogeneity to determine ecosystem functioning

Background and aims: Spatial distribution of soil nutrients (soil heterogeneity) and availability have strong effects on above- and belowground plant functional traits. Although there is ample evidence on the tight links between functional traits and ecosystem functioning, the role played by soil heterogeneity and availability as modulators of such relationship is poorly known. Methods: We conducted a factorial experiment in microcosms containing grasses, legumes and non-legume forbs communities differing in composition to evaluate how soil heterogeneity and availability (50 and 100 mg N) affect the links between traits and ecosystem functioning. Community-aggregated specific leaf area (SLA_agg) and specific root length (SRL_agg) were measured as both relevant response traits to soil heterogeneity and availability, and significant effect traits affecting ecosystem functioning (i. e., belowground biomass, β-glucosidase and acid phosphatase activities, and in situ N availability rate). Results: SRL_agg was negatively and significantly associated to β-glucosidase, phosphatase and N availability rate in the high nutrient availability and heterogeneous distribution scenario. We found a significant negative relationship between SLA_agg and availability rate of mineral-N under low nutrient availability conditions. Conclusions: Soil heterogeneity modulated the effects of both traits and nutrient availability on ecosystem functioning. Specific root length was the key trait associated with soil nutrient cycling and belowground biomass in contrasted heterogeneous soil conditions. The inclusion of soil heterogeneity into the trait-based response-effect framework may help to scale from plant communities to the ecosystem level.