Increasing native, but not exotic, biodiversity increases aboveground productivity in ungrazed and intensely grazed grasslands.

Species-rich native grasslands are frequently converted to species-poor exotic grasslands or pastures; however, the consequences of these changes for ecosystem functioning remain unclear. Cattle grazing (ungrazed or intensely grazed once), plant species origin (native or exotic), and species richness (4-species mixture or monoculture) treatments were fully crossed and randomly assigned to plots of grassland plants. We tested whether (1) native and exotic plots exhibited different responses to grazing for six ecosystem functions (i.e., aboveground productivity, light interception, fine root biomass, tracer nitrogen uptake, biomass consumption, and aboveground biomass recovery), and (2) biodiversity-ecosystem functioning relationships depended on grazing or species origin. We found that native and exotic species exhibited different responses to grazing for three of the ecosystem functions we considered. Intense grazing decreased fine root biomass by 53% in exotic plots, but had no effect on fine root biomass in native plots. The proportion of standing biomass consumed by cattle was 16% less in exotic than in native grazed plots. Aboveground biomass recovery was 30% less in native than in exotic plots. Intense grazing decreased aboveground productivity by 25%, light interception by 14%, and tracer nitrogen uptake by 54%, and these effects were similar in native and exotic plots. Increasing species richness from one to four species increased aboveground productivity by 42%, and light interception by 44%, in both ungrazed and intensely grazed native plots. In contrast, increasing species richness did not influence biomass production or resource uptake in ungrazed or intensely grazed exotic plots. These results suggest that converting native grasslands to exotic grasslands or pastures changes ecosystem structure and processes, and the relationship between biodiversity and ecosystem functioning.

Biodiversity, productivity and the temporal stability of productivity: patterns and processes.

Theory predicts that the temporal stability of productivity, measured as the ratio of the mean to the standard deviation of community biomass, increases with species richness and evenness. We used experimental species mixtures of grassland plants to test this hypothesis and identified the mechanisms involved. Additionally, we tested whether biodiversity, productivity and temporal stability were similarly influenced by particular types of species interactions. We found that productivity was less variable among years in plots planted with more species. Temporal stability did not depend on whether the species were planted equally abundant (high evenness) or not (realistically low evenness). Greater richness increased temporal stability by increasing overyielding, asynchrony of species fluctuations and statistical averaging. Species interactions that favoured unproductive species increased both biodiversity and temporal stability. Species interactions that resulted in niche partitioning or facilitation increased both productivity and temporal stability. Thus, species interactions can promote biodiversity and ecosystem services.

Biodiversity maintenance mechanisms differ between native and novel exotic-dominated communities.

In many systems, native communities are being replaced by novel exotic-dominated ones. We experimentally compared species diversity decline between nine-species grassland communities under field conditions to test whether diversity maintenance mechanisms differed between communities containing all exotic or all native species using a pool of 40 species. Aboveground biomass was greater in exotic than native plots, and this difference was larger in mixtures than in monocultures. Species diversity declined more in exotic than native communities and declines were explained by different mechanisms. In exotic communities, overyielding species had high biomass in monoculture and diversity declined linearly as this selection effect increased. In native communities, however, overyielding species had low biomass in monoculture and there was no relationship between the selection effect and diversity decline. This suggests that, for this system, yielding behaviour is fundamentally different between presumably co-evolved natives and coevolutionarily naive exotic species, and that native-exotic status is important to consider.

Species interaction mechanisms maintain grassland plant species diversity.

Development of theory has outpaced experimental tests for most maintenance of diversity mechanisms. Here we demonstrate how data from biodiversity-ecosystem functioning experiments can be used to determine the mechanisms that maintain plant species diversity. We hypothesized that grassland plant diversity is maintained by two classes of mechanisms: (1) equalizing mechanisms, which reduce asymmetric competition by reducing differences in monoculture biomass production among species in mixture, and (2) species interaction mechanisms, which increase overyielding by increasing niche partitioning and facilitation among species in mixture. Specifically, equalizing mechanisms reduce the coefficient of variation in monoculture biomass production among species in mixture. Species interaction mechanisms increase species overyielding in mixture, especially for low-biomass species. We tested these predictions with a seven-year data set from an experiment that varied grassland plant species evenness and richness. We used path analysis to model effects of these mechanisms on annual and multiyear changes in diversity. We found that diversity was frequently maintained by species interaction mechanisms and was infrequently maintained by equalizing mechanisms. Species interaction mechanisms maintained diversity by allowing the species that produced the least biomass in monoculture to benefit the most from species interactions in mixture. Equalizing mechanisms infrequently maintained diversity because asymmetric competition infrequently resulted in competitive exclusion. We propose that this mechanistic framework be used to better understand the specific processes that influence diversity.