Differential effects of functional traits on aboveground biomass in semi-natural grasslands

Despite increasing evidence on the importance of species functional characteristics for ecosystem processes, two major hypotheses suggest different mechanisms: the 'mass ratio hypothesis' assumes that functional traits of the dominant species determine ecosystem processes, while the 'complementarity hypothesis' predicts that resource niches may be used more completely when a community is functionally more diverse. Here, we present a method which uses two different groups of biotic predictor variables being (1) abundance-weighted mean (=aggregated) trait values and (2) functional trait diversity based on Rao's quadratic diversity (FDQ) to test the competing hypotheses on biodiversity-ecosystem functioning relationships after accounting for co-varying abiotic factors. We applied this method to data recorded on biodiversity-biomass relationships and environmental variables in 35 semi-natural temperate grasslands and used a literature-based matrix of fourteen plant functional traits to assess the explanatory power of models including different sets of predictor variables. Aboveground community biomass did not correlate with species richness. Abiotic factors, in particular soil nitrogen concentration, explained about 50% of variability in aboveground biomass. The best model incorporating functional trait diversity explained only about 30%, while the best model based on aggregated trait values explained about 54% of variability in aboveground biomass. The inclusion of all predictor variable groups in a combined model increased the predictive power to about 75%. This model comprised soil nitrogen concentration as abiotic factor, aggregated traits being indicative for species competitive dominance (rooting depth, leaf distribution, specific leaf area, perennial life cycle) and functional trait diversity in vegetative plant height, leaf area and life cycle. Our study strongly suggests that abiotic factors, trait values of the dominant species and functional trait diversity in combination may best explain differences in aboveground community biomass in natural ecosystems and that their isolated consideration may be misleading.