Functional groups differ in trait means, but not in trait plasticity to species richness in local grassland communities.

Despite growing interest in incorporating intraspecific variation of functional traits in community-level studies, it remains unclear whether species classified into functional groups based on interspecific trait differences are similar regarding their variation in trait expression in response to varying plant diversity and composition in local communities. In a large biodiversity experiment (Jena Experiment) designed on a trait-based a priori definition of functional groups (grasses, legumes, small herbs, tall herbs), we studied means, extent of variation (coefficient of variation across communities) and plasticity to increased plant diversity (slopes over a logarithmic species richness ranging from 1, 2, 4, 8 and 16 to 60 species) for nine functional traits. Species means and extent of variation in traits related to nitrogen (N) acquisition and N use differed among functional groups and were more similar in phylogenetically closely related species than expected by chance. Species in the same functional group showed a weak phylogenetic signal and varied widely in means and extent of variation in traits related to shoot architecture and to a smaller extent in leaf traits related to carbon acquisition. This indicated that functional groups were less distinguishable in light than in nitrogen acquisition strategies. The direction and degree of trait plasticity to increasing species richness did not show a phylogenetic signal and were not different among functional groups, but varied largely among species within functional groups. Correlation structures in trait means, extent of trait variation and trait plasticity revealed functional tradeoffs in the acquisition of nitrogen and light across species. While correlations between trait means and extent of trait variation varied from trait to trait (positive, negative or unrelated), trait means and trait plasticity were mostly unrelated. Our results suggest that the concept of functional groups is viable, but context-specific trait measurements are required to improve our understanding about the functional significance of intraspecific trait variation and interspecific trait differences in local plant communities.

Origin context of trait data matters for predictions of community performance in a grassland biodiversity experiment.

Plant functional traits may explain the positive relationship between species richness and ecosystem functioning, but species-level trait variation in response to growth conditions is often ignored in trait-based predictions of community performance. In a large grassland biodiversity experiment (Jena Experiment), we measured traits on plants grown as solitary individuals, in monocultures or in mixtures. We calculated two measures of community-level trait composition, i.e., community-weighted mean traits (CWM) and trait diversity (Rao's quadratic entropy; FD) based on different contexts in which traits were measured (trait origins). CWM and FD values of the different measurement origins were then compared regarding their power to predict community biomass production and biodiversity effects quantified with the additive partitioning method. Irrespective of trait origin, models combining CWM and FD values as predictors best explained community biomass and biodiversity effects. CWM values based on monoculture, mixture-mean or community-specific trait data were similarly powerful predictors, but predictions became worse when trait values originated from solitary-grown individuals. FD values based on monoculture traits were the best predictors of community biomass and net biodiversity effects, while FD values based on community-specific traits were the best predictors for complementarity and selection effects. Traits chosen as best CWM predictors were not strongly affected by trait origin but traits chosen as best FD predictors varied strongly dependent on trait origin and altered the predictability of community performance. We conclude that by adjusting their functional traits to species richness and even specific community compositions, plants can change community-level trait compositions, thereby also changing community biomass production and biodiversity effects. Incorporation of these plastic trait adjustments of plants in trait-based ecology can improve its predictive power in explaining biodiversity-ecosystem functioning relationships.

Functionally and phylogenetically diverse plant communities key to soil biota.

Recent studies assessing the role of biological diversity for ecosystem functioning indicate that the diversity of functional traits and the evolutionary history of species in a community, not the number of taxonomic units, ultimately drives the biodiversity--ecosystem-function relationship. Here, we simultaneously assessed the importance of plant functional trait and phylogenetic diversity as predictors of major trophic groups of soil biota (abundance and diversity), six years from the onset of a grassland biodiversity experiment. Plant functional and phylogenetic diversity were generally better predictors of soil biota than the traditionally used species or functional group richness. Functional diversity was a reliable predictor for most biota, with the exception of soil microorganisms, which were better predicted by phylogenetic diversity. These results provide empirical support for the idea that the diversity of plant functional traits and the diversity of evolutionary lineages in a community are important for maintaining higher abundances and diversity of soil communities.

Using plant functional traits to explain diversity-productivity relationships.

BACKGROUND: The 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 FINDINGS: We 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 (FD(Q)) 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 FD(Q) (36 and 38% of explained variation respectively). FD(Q) 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 FD(Q) led to significant model improvement compared to a separate consideration of different components of functional trait composition. Effects of FD(Q) 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/SIGNIFICANCE: Our 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.

Density-independent mortality and increasing plant diversity are associated with differentiation of Taraxacum officinale into r- and K-strategists.

BACKGROUND: Differential selection between clones of apomictic species may result in ecological differentiation without mutation and recombination, thus offering a simple system to study adaptation and life-history evolution in plants. METHODOLOGY/PRINCIPAL FINDINGS: We caused density-independent mortality by weeding to colonizer populations of the largely apomictic Taraxacum officinale (Asteraceae) over a 5-year period in a grassland biodiversity experiment (Jena Experiment). We compared the offspring of colonizer populations with resident populations deliberately sown into similar communities. Plants raised from cuttings and seeds of colonizer and resident populations were grown under uniform conditions. Offspring from colonizer populations had higher reproductive output, which was in general agreement with predictions of r-selection theory. Offspring from resident populations had higher root and leaf biomass, fewer flower heads and higher individual seed mass as predicted under K-selection. Plants grown from cuttings and seeds differed to some degree in the strength, but not in the direction, of their response to the r- vs. K-selection regime. More diverse communities appeared to exert stronger K-selection on resident populations in plants grown from cuttings, while we did not find significant effects of increasing species richness on plants grown from seeds. CONCLUSIONS/SIGNIFICANCE: Differentiation into r- and K-strategists suggests that clones with characteristics of r-strategists were selected in regularly weeded plots through rapid colonization, while increasing plant diversity favoured the selection of clones with characteristics of K-strategists in resident populations. Our results show that different selection pressures may result in a rapid genetic differentiation within a largely apomictic species. Even under the assumption that colonizer and resident populations, respectively, happened to be r- vs. K-selected already at the start of the experiment, our results still indicate that the association of these strategies with the corresponding selection regimes was maintained during the 5-year experimental period.

Foliar and soil δ15N values reveal increased nitrogen partitioning among species in diverse grassland communities.

Plant and soil nitrogen isotope ratios (δ¹5N) were studied in experimental grassland plots of varying species richness. We hypothesized that partitioning of different sources of soil nitrogen among four plant functional groups (legumes, grasses, small herbs, tall herbs) should increase with diversity. Four years after sowing, all soils were depleted in ¹5N in the top 5 cm whereas in non-legume plots soils were enriched in ¹5N at 5-25 cm depth. Decreasing foliar δ¹5N and Δδ¹5N (= foliar δ¹5N-soil δ¹5N) values in legumes indicated increasing symbiotic N2 fixation with increasing diversity. In grasses, foliar Δδ¹5N also decreased with increasing diversity suggesting enhanced uptake of N depleted in ¹5N. Foliar Δδ¹5N values of small and tall herbs were unaffected by diversity. Foliar Δδ¹5N values of grasses were also reduced in plots containing legumes, indicating direct use of legume-derived N depleted in ¹5N. Increased foliar N concentrations of tall and small herbs in plots containing legumes without reduced foliar δ¹5N indicated that these species obtained additional mineral soil N that was not consumed by legumes. These functional group and species specific shifts in the uptake of different N sources with increasing diversity indicate complementary resource use in diverse communities.

Differential effects of plant diversity on functional trait variation of grass species.

BACKGROUND AND AIMS: Functional trait differences and trait adjustment in response to influences of the biotic environment could reflect niche partitioning among species. In this study, we tested how variation in above-ground plant traits, chosen as indicators for light and nitrogen acquisition and use, differs among taxonomically closely related species (Poaceae) to assess their potential for niche segregation at increasing plant diversity. METHODS: Traits of 12 grass species were measured in experimental grasslands (Jena Experiment) of varying species richness (from 1 to 60) and presence of particular functional groups (grasses, legumes, tall herbs and small herbs). KEY RESULTS: Grass species increased shoot and leaf length, investment into supporting tissue (stem mass fraction) and specific leaf area as well as reduced foliar δ(13)C values with increasing species richness, indicating higher efforts for light acquisition. These species-richness effects could in part be explained by a higher probability of legume presence in more diverse communities. Leaf nitrogen concentrations increased and biomas s : N ratios in shoots decreased when grasses grew with legumes, indicating an improved nitrogen nutrition. Foliar δ(15)N values of grasses decreased when growing with legumes suggesting the use of depleted legume-derived N, while decreasing δ(15)N values with increasing species richness indicated a shift in the uptake of different N sources. However, efforts to optimize light and nitrogen acquisition by plastic adjustment of traits in response to species richness and legume presence, varied significantly among grass species. It was possible to show further that trait adjustment of grass species increased niche segregation in more diverse plant communities but that complementarity through niche separation may differ between light and nutrient acquisition. CONCLUSIONS: The results suggest that even among closely related species such as grasses different strategies are used to cope with neighbours. This lack in redundancy in turn may facilitate complementary resource use and coexistence.