BackgroundThe different hypotheses proposed to explain positive species richness–productivity relationships, i.e. selection effect and complementarity effect, imply that plant functional characteristics are at the core of a mechanistic understanding of biodiversity effects.Methodology/Principal FindingsWe used two community-wide measures of plant functional composition, (1) community-weighted means of trait values (CWM) and (2) functional trait diversity based on Rao’s quadratic diversity (FDQ) to predict biomass production and measures of biodiversity effects in experimental grasslands (Jena Experiment) with different species richness (2, 4, 8, 16 and 60) and different functional group number and composition (1 to 4; legumes, grasses, small herbs, tall herbs) four years after establishment. Functional trait composition had a larger predictive power for community biomass and measures of biodiversitity effects (40–82% of explained variation) than species richness per se (<1–13% of explained variation). CWM explained a larger amount of variation in community biomass (80%) and net biodiversity effects (70%) than FDQ (36 and 38% of explained variation respectively). FDQ explained similar proportions of variation in complementarity effects (24%, positive relationship) and selection effects (28%, negative relationship) as CWM (27% of explained variation for both complementarity and selection effects), but for all response variables the combination of CWM and FDQ led to significant model improvement compared to a separate consideration of different components of functional trait composition. Effects of FDQ were mainly attributable to diversity in nutrient acquisition and life-history strategies. The large spectrum of traits contributing to positive effects of CWM on biomass production and net biodiversity effects indicated that effects of dominant species were associated with different trait combinations.Conclusions/SignificanceOur results suggest that the identification of relevant traits and the relative impacts of functional identity of dominant species and functional diversity are essential for a mechanistic understanding of the role of plant diversity for ecosystem processes such as aboveground biomass production.

Summary 1.Positive effects of biodiversity on plant productivity may result from diversity-induced changes in the size or density of individual plants, yet these two possibilities have never been tested at the same time in a biodiversity experiment with a large species pool. Here, we distinguish between size effects and density effects on plant productivity, using data from 198 experimental grassland communities that contained 1-16 species. Plant modules such as tillers or rosettes were defined as relevant units, being equivalent to plant individuals in the majority of species. 2. In agreement with previous studies, we found positive effects of species richness on above-ground productivity. We show that this positive biodiversity effect resulted from diversity-induced increases in module density rather than from increases in module size. In contrast, variation in productivity within diversity levels was related to module size rather than module density. 3. The size-density relationships varied among plant functional groups and among species but their average response to increasing species richness paralleled the pattern observed at the level of the entire plant communities: species richness had a positive effect on above-ground species biomass and species module density but not on species module size. Twenty-four out of 26 overyielding species had denser populations and 25 out of 28 underyielding species had smaller modules in mixtures than in monocultures. 4. Synthesis . In grasslands, an increase in community productivity must involve an increase in plant size or density. We found that diversity-induced increases in productivity were related to diversityinduced increases in density, whereas diversity-independent increases in productivity were related to increases in plant size. Our results suggest that increased density of overyielding species in mixtures was the main driver of the positive biodiversity-productivity relationship in our experiment. We conclude that the mechanisms leading to enhanced productivity of species-rich as compared with species-poor communities cannot be derived from mechanisms explaining high productivity within communities that contain a particular number of species.

Invasive species often evolve rapidly in response to the novel biotic and abiotic conditions in their introduced range. Such adaptive evolutionary changes might play an important role in the success of some invasive species. Here, we investigated whether introduced European populations of the South African ragwort Senecio inaequidens (Asteraceae) have genetically diverged from native populations. We carried out a greenhouse experiment where 12 South African and 11 European populations were for several months grown at two levels of nutrient availability, as well as in the presence or absence of a generalist insect herbivore. We found that, in contrast to a current hypothesis, plants from introduced populations had a significantly lower reproductive output, but higher allocation to root biomass, and they were more tolerant to insect herbivory. Moreover, introduced populations were less genetically variable, but displayed greater plasticity in response to fertilization. Finally, introduced populations were phenotypically most similar to a subset of native populations from mountainous regions in southern Africa. Taking into account the species' likely history of introduction, our data support the idea that the invasion success of Senecio inaequidens in Central Europe is based on selective introduction of specific preadapted and plastic genotypes rather than on adaptive evolution in the introduced range. [References: 41