Defoliation, trampling and nutrient return differentially influence grassland productivity by modulating trait-dependent plant community composition: insights from a simulated grazing experiment.

Ungulate grazing involves multiple components, including defoliation, dung and urine return, and trampling, which supply offsetting or synergistic effects on plant community composition and productivity (ANPP), but these effects have not been fully studied. Plant functional traits may reflect the response of plants to disturbance and their impact on ecosystem functions. Species turnover and intraspecific trait variation (ITV) are important drivers of community trait composition. We conducted a simulated grazing experiment in a steppe grassland in northern China to examine the effects of defoliation, dung and urine return, and trampling on community-weighted mean (CWM), functional diversity (FD) and ANPP, and to disentangle the roles of species turnover and ITV in driving these changes. We found that defoliation had a dominant effect on CWMs and FDs of all four traits through species turnover and ITV, respectively, resulting in a convergence of traits towards as more resource-acquisitive strategy. Dung-urine return resulted in more resource-acquisitive community traits mainly through ITV, whereas there were no significant effects on FDs except for leaf C/N. Trampling increased CWM of leaf dry matter content primarily driven by ITV, and had no significant effect on FDs. Furthermore, our simulated grazing positively affected ANPP, primarily due to nutrient additions from dung and urine, and ITV largely explained the variation in ANPP. These findings highlight the multifaceted effects of grazing components on community structure and ANPP, and the significance of ITV in shaping grassland plant communities and productivity.