Warming delays but grazing advances leaf senescence of five plant species in an alpine meadow.

Leaf senescence is the final stage in the life cycle of leaves and is critical to plants' fitness as well as to ecosystem carbon and nutrient cycling. To date, most understanding about the responses of leaf senescence to environmental changes has derived from research in forests, but the topic has been relatively neglected, especially under grazing conditions, in natural grasslands. We conducted a 3-year manipulative asymmetric warming with moderate grazing experiment to explore the responses of leaf senescence of five main species in an alpine meadow on the Qinghai-Tibetan Plateau. We found that warming prolonged leaf longevity through earlier leaf-out and later leaf senescence, and grazing prolonged it through a greater advance in leaf-out than first leaf coloration for all plants. Warming did not affect leaf nitrogen (N) content or N resorption efficiency (NRE), but grazing increased N content in coloring leaves for P. anserine and P. nivea and decreased NRE for K. humilis, P. anserine under no-warming, and for P. nivea under warming. The interactive effects of warming and grazing on leaf phenology and leaf traits depended on species identity and year. There were positive relationships between leaf-out and leaf senescence mainly derived from grazing, and positive relationships between NRE from old leaves and leaf senescence for three out of five plant species. Therefore, our results indicated that earlier leaf-out could result in earlier leaf senescence only under grazing, but depending on plant species. Delayed leaf coloring increased NRE from old leaves for some plant species measured under warming and grazing. Our results suggested that alpine plants may develop strategies to adapt to warming and grazing to assimilate more carbon through prolonged leaf longevity rather than increased NRE through earlier leaf coloring in the alpine meadow.

Ambient climate determines the directional trend of community stability under warming and grazing.

Changes in ecological processes over time in ambient treatments are often larger than the responses to manipulative treatments in climate change experiments. However, the impacts of human-driven environmental changes on the stability of natural grasslands have been typically assessed by comparing differences between manipulative plots and reference plots. Little is known about whether or how ambient climate regulates the effects of manipulative treatments and their underlying mechanisms. We collected two datasets, one a 36-year long-term observational dataset from 1983 to 2018, and the other a 10-year manipulative asymmetric warming and grazing experiment using infrared heaters with moderate grazing from 2006 to 2015 in an alpine meadow on the Tibetan Plateau. The 36-year observational dataset shows that there was a nonlinear response of community stability to ambient temperature with a positive relationship between them due to an increase in ambient temperature in the first 25 years and then a decrease in ambient temperature thereafter. Warming and grazing decreased community stability with experiment duration through an increase in legume cover and a decrease in species asynchrony, which was due to the decreasing background temperature through time during the 10-year experiment period. Moreover, the temperature sensitivity of community stability was higher under the ambient treatment than under the manipulative treatments. Therefore, our results suggested that ambient climate may control the directional trend of community stability while manipulative treatments may determine the temperature sensitivity of the response of community stability to climate relative to the ambient treatment. Our study emphasizes the importance of the context dependency of the response of community stability to human-driven environmental changes.