Water limitation drives species loss in grassland communities after nitrogen addition and warming

Nutrient addition, particularly nitrogen, often increases plant aboveground biomass but causes species loss. Asymmetric competition for light is frequently assumed to explain the biomass-driven species loss. However, it remains unclear whether other factors such as water can also play a role. Increased aboveground leaf area following nitrogen addition and warming may increase transpiration and cause water limitation, leading to a decline in diversity. To test this, we conducted field measurements in a grassland community exposed to nitrogen and water addition, and warming. We found that warming and/or nitrogen addition significantly increased aboveground biomass but reduced species richness. Water addition prevented species loss in either nitrogen-enriched or warmed treatments, while it partially mitigated species loss in the treatment exposed to increases in both temperature and nitrogen. These findings thus strongly suggest that water limitation can be an important driver of species loss as biomass increases after nitrogen addition and warming when soil moisture is limiting. This result is further supported by a meta-analysis of published studies across grasslands worldwide. Our study indicates that loss of grassland species richness in the future may be greatest under a scenario of increasing temperature and nitrogen deposition, but decreasing precipitation.