Abstract. Insurance effects of biodiversity can stabilize the functioning of multispecies ecosystems against environmental variability when differential species' responses lead to asynchronous population dynamics. When responses are not perfectly positively correlated, declines in some populations are compensated by increases in others, smoothing variability in ecosystem productivity. This variance reduction effect of biodiversity is analogous to the riskspreading benefits of diverse investment portfolios in financial markets.We use data from the BIODEPTH network of grassland biodiversity experiments to perform a general test for stabilizing effects of plant diversity on the temporal variability of individual species, functional groups, and aggregate communities. We tested three potential mechanisms: reduction of temporal variability through population asynchrony; enhancement of long-term average performance through positive selection effects; and increases in the temporal mean due to overyielding.Our results support a stabilizing effect of diversity on the temporal variability of grassland aboveground annual net primary production through two mechanisms. Two-species communities with greater population asynchrony were more stable in their average production over time due to compensatory fluctuations. Overyielding also stabilized productivity by increasing levels of average biomass production relative to temporal variability. However, there was no evidence for a performance-enhancing effect on the temporal mean through positive selection effects. In combination with previous work, our results suggest that stabilizing effects of diversity on community productivity through population asynchrony and overyielding appear to be general in grassland ecosystems.

Ecosystem effects of biodiversity manipulations in European grasslands AbstractWe present a multisite analysis of the relationship between plant diversity and ecosystem functioning within the European BIODEPTH network of plant-diversity manipulation experiments. We report results of the analysis of 11 variables addressing several aspects of key ecosystem processes like biomass production, resource use (space, light, and nitrogen), and decomposition, measured across three years in plots of varying plant species richness at eight different European grassland field sites. Differences among sites explained substantial and significant amounts of the variation of most of the ecosystem processes examined. However, against this background of geographic variation, all the aspects of plant diversity and composition we examined (i.e., both numbers and types of species and functional groups) produced significant, mostly positive impacts on ecosystem processes.Analyses using the additive partitioning method revealed that complementarity effects (greater net yields than predicted from monocultures due to resource partitioning, positive interactions, etc.) were stronger and more consistent than selection effects (the covariance between monoculture yield and change in yield in mixtures) caused by dominance of species with particular traits. In general, communities with a higher diversity of species and functional groups were more productive and utilized resources more completely by intercepting more light, taking up more nitrogen, and occupying more of the available space. Diversity had significant effects through both increased vegetation cover and greater nitrogen retention by plants when this resource was more abundant through N2 fixation by legumes. However, additional positive diversity effects remained even after controlling for differences in vegetation cover and for the presence of legumes in communities. Diversity effects were stronger on above-than belowground processes. In particular, clear diversity effects on decomposition were only observed at one of the eight sites.The ecosystem effects of plant diversity also varied between sites and years. In general, diversity effects were lowest in the first year and stronger later in the experiment, indicating that they were not transitional due to community establishment. These analyses of our complete ecosystem process data set largely reinforce our previous results, and those from comparable biodiversity experiments, and extend the generality of diversity-ecosystem functioning relationships to multiple sites, years, and processes. Abstract. We present a multisite analysis of the relationship between plant diversity and ecosystem functioning within the European BIODEPTH network of plant-diversity manipulation experiments. We report results of the analysis of 11 variables addressing several aspects of key ecosystem processes like biomass production, resource use (space, light, and nitrogen), and decomposition, measured across three years in plots of varying plant species ri...

The relationship between plant diversity and nitrate leaching into groundwater was investigated in a mid‐European semi‐natural grassland ecosystem. An experimental approach was used to directly manipulate plant diversity in the field, while holding other environmental factors constant. Species loss was simulated by establishing grassland communities of 16, 8, 4, 2, 1, and 0 plant species, composed of 3, 2, or 1 functional groups (grasses, legumes, and non‐legume herbs). Every diversity treatment was replicated with several different species mixtures. Nitrate leaching was determined by continuous extraction of soil solution below the rooting zone and modeling of seepage rates. The concentration of nitrate in the soil solution was highly variable within each level of diversity. In bare ground plots and several low‐diversity mixtures containing legumes, nitrate concentrations were higher than the official European Union threshold value for drinking water of 50 mg/L, with maximum values of up to 350 mg/L measured in Trifolium pratense monocultures. Total annual loss of nitrate was unaffected by the number of plant species or functional groups, but it was highly dependent on the specific species composition of the communities, and plots with legumes lost significantly more nitrate than plots without them. Aboveground biomass had no influence on nitrate loss, whereas leaching was negatively correlated with increasing root biomass. The abundance of legumes within a community, litter decomposition rates, and net nitrification were all positively correlated with total nitrate loss. However, in those communities containing legumes, leaching decreased with increasing diversity, because higher species richness led to a reduction in legume dominance, to a reduced nitrate supply through nitrification, and to a complementary uptake of nitrate by grasses and non‐leguminous herbs. Based on these results, we expect that increasing the diversity of non‐leguminous species or functional groups would reduce the risk of nitrate leaching in low‐diversity grass–clover mixtures of ley‐farming systems, while allowing for a more efficient exploitation of the beneficial fertilization effect provided by legumes. Corresponding Editor: M. Loreau.

Global change drivers are rapidly altering resource availability and biodiversity. While there is consensus that greater biodiversity increases the functioning of ecosystems, the extent to which biodiversity buffers ecosystem productivity in response to changes in resource availability remains unclear. We use data from 16 grassland experiments across North America and Europe that manipulated plant species richness and one of two essential resources—soil nutrients or water—to assess the direction and strength of the interaction between plant diversity and resource alteration on above-ground productivity and net biodiversity, complementarity, and selection effects. Despite strong increases in productivity with nutrient addition and decreases in productivity with drought, we found that resource alterations did not alter biodiversity–ecosystem functioning relationships. Our results suggest that these relationships are largely determined by increases in complementarity effects along plant species richness gradients. Although nutrient addition reduced complementarity effects at high diversity, this appears to be due to high biomass in monocultures under nutrient enrichment. Our results indicate that diversity and the complementarity of species are important regulators of grassland ecosystem productivity, regardless of changes in other drivers of ecosystem function.

The effects of biodiversity on ecosystem functioning generally increase over time but the underlying processes remain unclear. Using 26 long-term grassland and forest experimental ecosystems we demonstrate that biodiversity-ecosystem functioning relationships strengthen mainly by greater increases in functioning in high-diversity communities in grasslands and forests. In grasslands, biodiversity effects also strengthen due to decreases in functioning in low-diversity communities. Contrasting trends across grasslands are associated with differences in soil characteristics.