Density-dependent plant growth drives grazer stimulation of aboveground net primary production in Yellowstone grasslands

The mechanisms by which grazing animals influence aboveground net primary production (ANPP) in grasslands have long been an area of active research. The prevailing wisdom is that grazing can increase ANPP by increasing the availability of growth-limiting resources such as nitrogen and water, but recent theory suggests that the density-dependent growth of grassland vegetation can lead to grazer-stimulation of ANPP simply by removing shoot biomass and increasing relative growth rate (RGR). We compared the relative roles of resource availability and density-dependent growth in driving positive responses of ANPP to grazing in Yellowstone National Park. We measured the effects of clipping (50% simulated grazing intensity) and natural grazing on soil nitrogen availability, soil moisture, and shoot growth over 2 months in two grassland plant communities (mesic and dry) grazed primarily by bison. Clipping increased RGR by over 100% in both grassland types but had no effect on N availability or soil moisture during the same growth periods. Clipping stimulated ANPP only at mesic grassland, and the magnitude of this effect was strongly related to the initial plant biomass at the time of clipping relative to estimated peak biomass, supporting the density-dependent framework. Bison grazing had qualitatively similar effects on ANPP and RGR to clipping with no accompanying effects on N availability or soil moisture. Our results show how grazing can stimulate ANPP independent of a direct influence on resource availability simply by exploiting the dynamics of density-dependent plant growth.