Chris M. Clark

In a 10-year (1996-2005) biodiversity experiment, the mechanisms underlying the increasingly positive effect of biodiversity on plant biomass production shifted from sampling to complementarity over time. The effect of diversity on plant biomass was associated primarily with the accumulation of higher total plant nitrogen pools (N g m-2) and secondarily with more efficient N use at higher diversity. The accumulation of N in living plant biomass was significantly increased by the presence of legumes, C4 grasses, and their combined presence. Thus, these results provide clear evidence for the increasing effects of complementarity through time and suggest a mechanism whereby diversity increases complementarity through the increased input and retention of N, a commonly limiting nutrient.

Summary The Eurasian steppe has long been subject to grazing by domestic ungulates at high levels, resulting in widespread deterioration of biodiversity and ecosystem services. While abundant evidence demonstrates that heavy grazing alters the ecosystem structure and function of grasslands, research on how grazing specifically affects ecosystem functioning and stoichiometry on broad scales is scarce because of a lack of adequate ungrazed reference sites. We examined the effects of grazing on ecosystem functioning and C : N : P stoichiometry across a precipitation gradient along the 700 km C hina– M ongolia transect ( CMT ), covering three community types: meadow steppe, typical steppe and desert steppe. Long‐term grazing has dramatically altered the C , N and P pools and stoichiometry of steppe ecosystems along the CMT . Grazing reduced the C , N and P pools in above‐ground biomass and litter, while the responses in below‐ground biomass and soil C , N and P pools to grazing differed substantially among community types. Grazing increased N content and decreased C : N ratios in all plant compartments, suggesting accelerated N cycling. The altered C : N : P stoichiometry may be explained by changes in the composition of species and functional groups as well as increased foliar N and P contents for the same species in grazed communities. Synthesis and applications . Plant stoichiometric responses to grazing ranged from large in the meadow steppe to small in the typical steppe to generally insignificant in the desert steppe, implying that different underlying mechanisms operated along the regional precipitation gradient. Our findings suggest that reducing the stocking rate and restoring the vastly degraded steppes are essential to sustain native steppe biodiversity, ecosystem functioning and biological capacity for mitigating the impact of climate change in the Inner M ongolia grassland.

Global energy use and food production have increased nitrogen inputs to ecosystems worldwide, impacting plant community diversity, composition, and function. Previous studies show considerable variation across terrestrial herbaceous ecosystems in the magnitude of species loss following nitrogen (N) enrichment. What controls this variation remains unknown. We present results from 23 N-addition experiments across North America, representing a range of climatic, soil and plant community properties, to determine conditions that lead to greater diversity decline. Species loss in these communities ranged from 0 to 65% of control richness. Using hierarchical structural equation modelling, we found greater species loss in communities with a lower soil cation exchange capacity, colder regional temperature, and larger production increase following N addition, independent of initial species richness, plant productivity, and the relative abundance of most plant functional groups. Our results indicate sensitivity to N addition is co-determined by environmental conditions and production responsiveness, which overwhelm the effects of initial community structure and composition.

A number of experiments have addressed how increases in nitrogen availability increase the productivity and decrease the diversity of plant communities. We lack, however, a rigorous mechanistic understanding of how changes in abundance of particular species combine to produce changes in community productivity and diversity. Single experiments cannot provide insight into this issue because each species occurs only once per experiment, and each experiment is conducted in only one location; thus, it is impossible from single experiments to determine whether responses of particular species are consistent across environments or dependent on the particular environmental context in which the experiment was conducted. To address this issue, we assembled a dataset of 20 herbaceous species that were each represented in at least 6 different fertilization experiments and tested whether responses were general across experiments. Of the 20 species, one consistently increased in relative abundance and five consistently decreased across replicate experiments. A partially‐overlapping group of 8 species displayed responses to nitrogen that varied predictably among experiments as a function of geographic location, neighboring species, or a handful of other community characteristics (ANPP, precipitation, species richness, relative abundance of focal species in control plots, and community composition). Thus, despite modest replication and a limited number of predictor variables, we were able to identify consistent patterns in response of 10 out of 20 species across multiple experiments. We conclude that the responses of individual species to nitrogen addition are often predictable, but that in most cases these responses are functions of the abiotic or biotic environment. Thus, a rigorous understanding of how plant species respond to nitrogen addition will have to consider not only the traits of individual plant species, but also aspects of the communities in which those plants live.