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ISSN:0030-1299 ISSN:1600-0706

Below-ground resource partitioning is among the most prominent hypotheses for driving the positive biodiversity-ecosystem function relationship. However, experimental tests of this hypothesis in biodiversity experiments are scarce, and the available evidence is not consistent. We tested the hypothesis that resource partitioning in space, in time or in both space and time combined drives the positive effect of diversity on both plant productivity and total community resource uptake. At the community level, we predicted that total community resource uptake and biomass production above- and below-ground will increase with increased species richness or functional group richness. We predicted that, at the species level, resource partition breadth will become narrower, and that overlap between the resource partitions of different species will become smaller with increasing species richness or functional group richness. We applied multiple resource tracers (Li and Rb as potassium analogues, the water isotopologues-H2 18O and 2H2O, and 15N) in three seasons at two depths across a species and functional group richness gradient at a grassland biodiversity experiment. We used this multidimensional resource tracer study to test if plant species partition resources with increasing plant diversity across space, time or both simultaneously. At the community level, total community resource uptake of nitrogen and potassium and above- and below-ground biomass increased significantly with increasing species richness but not with increasing functional group richness. However, we found no evidence that resource partition breadth or resource partition overlap decreased with increasing species richness for any resource in space, time or both space and time combined. Synthesis. These findings indicate that below-ground resource partitioning may not drive the enhanced resource uptake or biomass production found here. Instead, other mechanisms such as facilitation, species-specific biotic feedback or above-ground resource partitioning are likely necessary for enhanced overall ecosystem function.

Summary 1.Niche complementarity and competitive disparity are driving mechanisms behind plant community assembly and productivity. Consequently, there is great interest in predicting species complementarity and their competitive differences from their functional traits as dissimilar species may compete less and result in more complete use of resources. 2.Here we assessed the role of trait dissimilarities on species complementarity and competitive disparities within an experimental gradient of plant species richness and functional trait dissimilarity. Communities were assembled using three pools of grass and forb species based on a priori knowledge of traits related to (1) above- and belowground spatial differences in resource acquisition, (2) phenological differences, or (3) both. Complementarity and competitive disparities were assessed by partitioning the overyielding in mixed species communities into species complementarity and dominance effects. 3.Community overyielding and the underlying complementarity and competitive dominance varied strongly among the three plant species pools. Overyielding and complementarity was greatest among species that were assembled based on their variation in both spatial and phenological traits. Competitive dominance was greatest when species were assembled based on spatial resource-acquisition traits alone. 4.In communities that were assembled based on species variation in only spatial or phenological traits greater competitive dominance was predicted by greater differences SLA and flowering initiation respectively, while greater complementarity was predicted by greater dissimilarity in leaf area and flowering senescence, respectively. Greater differences in leaf area could also be linked to greater species complementarity in communities assembled based on variation in both phenological and spatial traits, but trait dissimilarity was unrelated to competitive dominance in these communities. 5.Our results indicate that complementarity and competitive disparity among species are both driven by trait dissimilarities. However, the identity of the traits that drives the complementarity and competitive disparity depends on the trait variation among species that comprise the community. Moreover, we demonstrate that communities assembled with the greater variation in both spatial and phenological traits show the greatest complementarity among species. This article is protected by copyright. All rights reserved.

Summary 1.Efficient extraction of soil water is essential for the productivity of plant communities. However, research on the complementary use of resources in mixed plant communities, and especially the impact of plant species richness on root water uptake, is limited. So far, these investigations have been hindered by a lack of methods allowing for the estimation of root water uptake profiles. 2.The overarching aim of our study was to determine whether diverse grassland plant communities in general exploit soil water more deeply and whether this shift occurs all the time or only during times of enhanced water demand. 3.Root water uptake was derived by analyzing the diurnal decrease of soil water content separately at each measurement depth, thus yielding root water uptake profiles for 12 experimental grasslands communities with two different levels of species richness (4 and 16 sown species). Additional measurements of leaf water potential, stomatal conductance, and root traits were used to identify differences in water relations between plant functional groups. 4.Although the vertical root distribution did not differ between diversity levels, root water uptake shifted towards deeper layers (30 cm and 60 cm) in more diverse plots during periods of high vapor pressure deficit. Our results indicate that the more diverse communities were able to adjust their root water uptake, resulting in increased water uptake per root area compared to less diverse communities (52% at 20 cm, 118% at 30 cm, and 570% at 60 cm depth) and a more even distribution of water uptake over depth. Tall herbs, which had lower leaf water potential and higher stomatal conductance in more diverse mixtures, contributed disproportionately to dynamic niche partitioning in root water uptake. 5.This study underpins the role of diversity in stabilizing ecosystem function and mitigating drought stress effects during future climate change scenarios. Furthermore, the results provide evidence that root water uptake is not solely controlled by root length density distribution in communities with high plant diversity but also by spatial shifts in water acquisition. This article is protected by copyright. All rights reserved.

Numerous experiments have shown positive diversity effects on plant productivity, but little is known about related processes of carbon gain and allocation. We investigated these processes in a controlled environment (Montpellier European Ecotron) applying a continuous 13CO2 label for three weeks to 12 soil-vegetation monoliths originating from a grassland biodiversity experiment (Jena Experiment) and representing two diversity levels (4 and 16 sown species). Plant species richness did not affect community- and species-level 13C abundances neither in total biomass nor in non-structural carbohydrates (NSC). Community-level 13C excess tended to be higher in the 16-species than in the 4-species mixtures. Community-level 13C excess was positively related to canopy leaf nitrogen (N), i.e. leaf N per unit soil surface. At the species level shoot 13C abundances varied among plant functional groups and were larger in legumes and tall herbs than in grasses and small herbs and correlated positively with traits as leaf N concentrations, stomatal conductance and shoot height. The 13C abundances in NSC were larger in transport sugars (sucrose, raffinose-family oligosaccharides) than in free glucose, fructose and compounds of the storage pool (starch) suggesting that newly assimilated carbon is to a small portion allocated to storage. Our results emphasize that the functional composition of communities is key in explaining carbon assimilation in grasslands.

Little is known about the role of plant functional diversity for ecosystem-level carbon (C) fluxes. To fill this knowledge gap, we translocated monoliths hosting communities with four and 16 sown species from a long-term grassland biodiversity experiment (‘The Jena Experiment’) into a controlled environment facility for ecosystem research (Ecotron). This allowed quantifying the effects of plant diversity on ecosystem C fluxes as well as three parameters of C uptake efficiency (water and nitrogen use efficiencies and apparent quantum yield). By combining data on ecosystem C fluxes with vegetation structure and functional trait-based predictors, we found that increasing plant species and functional diversity led to higher gross and net ecosystem C uptake rates. Path analyses and light response curves unravelled the diversity of leaf nitrogen concentration in the canopy as a key functional predictor of C fluxes, either directly or indirectly via LAI and aboveground biomass.