Uncertainty about the effects of warming and grazing on soil nitrogen (N) availability, species composition, and aboveground net primary production (ANPP) limits our ability to predict how global carbon sequestration will vary under future warming with grazing in alpine regions. Through a controlled asymmetrical warming (1.2/1.7 degrees C during daytime/nighttime) with a grazing experiment from 2006 to 2010 in an alpine meadow, we found that warming alone and moderate grazing did not significantly affect soil net N mineralization. Although plant species richness significantly decreased by 10% due to warming after 2008, we caution that this may be due to the transient occurrence or disappearance of some rare plant species in all treatments. Warming significantly increased graminoid cover, except in 2009, and legume cover after 2008, but reduced non-legume forb cover in the community. Grazing significantly decreased cover of graminoids and legumes before 2009 but increased forb cover in 2010. Warming significantly increased ANPP regardless of grazing, whereas grazing reduced the response of ANPP to warming. N addition did not affect ANPP in both warming and grazing treatments. Our findings suggest that soil N availability does not determine ANPP under simulated warming and that heavy grazing rather than warming causes degradation of the alpine meadows.

Knowledge about the role of litter and dung decomposition in nutrient cycling and response to climate change and grazing in alpine ecosystems is still rudimentary. We conducted two separate studies to assess the relative role of warming and grazing on litter mass loss and on the temperature sensitivity of litter and dung mass loss. Experiments were conducted for 1-2 years under a controlled warming-grazing system and along an elevation gradient from 3200 to 3800 m. A free-air temperature enhancement system (FATE) using infrared heaters and grazing significantly increased soil temperatures (average 0.5-1.6 1C) from 0 to 40 cm depth, but neither warming nor grazing affected soil moisture except early in the growing seasons at 30 cm soil depth. Heaters caused greater soil warming at night-time compared with daytime, but grazing resulted in greater soil warming during daytime compared with night-time. Annual average values of the soil temperature at 5 cm were 3.2, 2.4 and 0.3 1C at 3200, 3600 and 3800 m, respectively. Neither warming nor grazing caused changes of litter quality for the first year of the controlled warming-grazing experiment. The effects of warming and grazing on litter mass losses were additive, increasing litter mass losses by about 19.3% and 8.3%, respectively, for the 2-year decomposition periods. The temperature sensitivity of litter mass losses was approximately 11% 1C À1 based on the controlled warming-grazing experiment. The annual cumulative litter mass loss was approximately 2.5 times that of dung along the elevation gradient. However, the temperature sensitivity (about 18% 1C À1 ) of the dung mass loss was about three times that of the litter mass loss. These results suggest greater warming at night-time compared with daytime may accelerate litter mass loss, and grazing will enhance carbon loss to atmosphere in the region through a decrease of litter biomass and an increase of dung production with an increase of stocking rate in future warmer conditions.

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Understanding how flowering phenology responds to warming and cooling (i.e., symmetric or asymmetric response) is needed to predict the response of flowering phenology to future climate change that will happen with the occurrence of warm and cold years superimposed upon a long-term trend. A three-year reciprocal translocation experiment was performed along an elevation gradient from 3200 m to 3800 m in the Tibetan Plateau for six alpine plants. Transplanting to lower elevation (warming) advanced the first flowering date (FFD) and transplanting to higher elevation (cooling) had the opposite effect. The FFD of early spring flowering plants (ESF) was four times less sensitive to warming than to cooling (by À2.1 d/8C and 8.4 d/8C, respectively), while midsummer flowering plants (MSF) were about twice as sensitive to warming than to cooling (À8.0 d/8C and 4.9 d/8C, respectively). Compared with pooled warming and cooling data, warming alone significantly underpredicted 3.1 d/8C for ESF and overestimated 1.7 d/8C for MSF. These results suggest that future empirical and experimental studies should consider nonlinear temperature responses that can cause such warming-cooling asymmetries as well as differing life strategies (ESF vs. MSF) among plant species.

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