Ardichvili, A. N.; Bittlingmaier, M.; Freschet, G. T.; Loreau, M.; Arnoldi, J.-F.

Species diversity potentially has a dual effect on communities: a generally positive effect on overall community biomass, reflecting the expression of species response and interaction traits, and a poorly characterised effect on mass-specific species contribution to ecosystem functions, reflecting the expression of their effect traits. Disentangling the effects of biodiversity on total biomass from those on effect trait expression would help settle a long-standing debate by clarifying how biodiversity relates to both facets of species effects on ecosystem functioning. Following the classical BEF approach, we calculate expected ecosystem function based on observed functioning in monoculture. We then derive a net biodiversity effect (NBE) and decompose it into four components: the classical complementarity and selection effects on total community biomass, and complementarity and selection effects on effect trait expression. The latter two reflect, respectively, a complementarity or facilitation in how effect traits influence the function, and how species with the highest potential for increasing the function become dominant in the community. We illustrate this NBE decomposition with three ecosystem functions (nitrogen retention capacity, soil hydraulic conductivity improvement, and forage digestibility) measured in assembled communities under controlled experimental conditions of perennial grassland plants. Regarding nitrogen retention, we find a positive complementary effect via total biomass, but a negative biodiversity effect via effect trait expression. For hydraulic conductivity improvement, biodiversity effects are mostly mediated by total biomass. As for forage digestibility, we found a positive complementarity effect on trait expression, outweighed however by a negative selection effect. This analysis reveals how biodiversity may have contrasting effects on ecosystem functions via its impact on biomass and effect trait expression. Separating between the effect of biodiversity on plant community biomass and on effect trait expression at the community level is one important step towards understanding the pathways by which diverse plant communities drive ecosystem functioning.