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 δ13C 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 δ15N values of grasses decreased when growing with legumes suggesting the use of depleted legume-derived N, while decreasing δ15N 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 coexistenc

Trait means, trait plasticity and trait differences to other species jointly explain species performances in grasslands of varying diversity

Functional traits may help to explain the great variety of species performances in plant communities, but it is not clear whether the magnitude of trait values of a focal species or trait differences to co‐occurring species are key for trait‐based predictions. In addition, trait expression within species is often plastic, but this variation has been widely neglected in trait‐based analyses. We studied functional traits and plant biomass of 59 species in 66 experimental grassland mixtures of varying species richness (Jena Experiment). We related mean species performances (species biomass and relative yield RY) and their plasticities along the diversity gradient to trait‐based pedictors involving mean species traits (Tmean), trait plasticities along the diversity gradient (Tslope), extents of trait variation across communities (TCV; coefficient of variation) and hierarchical differences (Tdiff) and trait distances (absolute values of trait differences Tdist) between focal and co‐occurring species. Tmean (30–55%) and Tdiff (30–33%) explained most variation in mean species performances and their plasticities, but Tslope (20–25%) was also important in explaining mean species performances. The mean species traits and the trait differences between focal species and neighbors with the greatest explanatory power were related to plant size and stature (shoot length, mass:height ratios) and leaf photosynthetic capacity (specific leaf area, stable carbon isotopes and leaf nitrogen concentration). The contribution of trait plasticities in explaining species performances varied in direction (positive or negative) and involved traits related to photosynthetic capacity, nitrogen acquisition (nitrogen concentrations and stable isotopes) as well as structural stability (shoot carbon concentrations). Our results suggest that incorporating plasticity in trait expression as well as trait differences to co‐occurring species is critical for extending trait‐based analyses to understand the assembly of plant communities and the contribution of individual species in structuring 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

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

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 (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/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

Mixed-effects model analysis of characteristics measured on plants grown from cuttings and seeds of resident and colonizer populations of <i>T. officinale</i> in the Jena Experiment, 5 years after sowing.

<p>Note: Models were fitted by stepwise inclusion of model terms. Listed are the results of likelihood-ratio tests that were applied to assess model improvement (L ratio) and the statistical significance of these tests (p values).</p

Effects of r- and K-selection regime on plant characteristics of <i>T. officinale</i>.

<p>Shoot biomass (a), root biomass (b), shoot:root ratio (c), aboveground vegetative biomass ( = leaves) (d), reproductive biomass (e), leaf number (f), maximum leaf length (g), flower head number (h), flower head diameter (i), seed mass per flower head (k), average seed mass (l), seed mass per plant individual (m), seed number per plant individual (n), first day of flowering (o), first day of seed maturity (p), and flower stalk length at seed maturity (q) were measured in plants grown from cuttings and seeds of resident populations (grey bars) and colonizer populations (white bars) of <i>T. officinale</i> in the Jena Experiment, 5 years after sowing. Values are means across populations ±1 SE for plants grown from cuttings and from seeds.</p

Species richness effects on germination rates of seeds of <i>T. officinale</i>.

<p>Material was collected from colonizer populations (open circles) and resident populations (closed circles) in the Jena Experiment, 5 years after sowing. Values are means per species-richness level ±1 SE.</p

Data from: Trait means, trait plasticity and trait differences to other species jointly explain species performances in grasslands of varying diversity

Functional traits may help to explain the great variety of species performances in plant communities, but it is not clear whether the magnitude of trait values of a focal species or trait differences to co-occurring species are key for trait-based predictions. In addition, trait expression within species is often plastic, but this variation has been widely neglected in trait-based analyses. We studied functional traits and plant biomass of 59 species in 66 experimental grassland mixtures of varying species richness (Jena Experiment). We related mean species performances (species biomass and relative yield RY) and their plasticities along the diversity gradient to trait-based pedictors involving mean species traits (Tmean), trait plasticities along the diversity gradient (Tslope), extents of trait variation across communities (TCV; coefficient of variation) and hierarchical differences (Tdiff) and trait distances (absolute values of trait differences Tdist) between focal and co-occurring species. Tmean (30–55%) and Tdiff (30–33%) explained most variation in mean species performances and their plasticities, but Tslope (20–25%) was also important in explaining mean species performances. The mean species traits and the trait differences between focal species and neighbors with the greatest explanatory power were related to plant size and stature (shoot length, mass:height ratios) and leaf photosynthetic capacity (specific leaf area, stable carbon isotopes and leaf nitrogen concentration). The contribution of trait plasticities in explaining species performances varied in direction (positive or negative) and involved traits related to photosynthetic capacity, nitrogen acquisition (nitrogen concentrations and stable isotopes) as well as structural stability (shoot carbon concentrations). Our results suggest that incorporating plasticity in trait expression as well as trait differences to co-occurring species is critical for extending trait-based analyses to understand the assembly of plant communities and the contribution of individual species in structuring plant communities

Species performance measures

This file contains measures of species performance (population-level biomass, relative yields, performance plasticities) for 60 study species based on aboveground biomass harvested in the mixtures of the Jena Experiment at estimated peak canopy development (late May) in the study years 2005-2009. All abbreviations of variables are explained in the data file; abbreviations of species names are explained in the READ me file