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Abstract Climate models predict more frequent, prolonged, and extreme droughts in the future. Therefore, drought experiments varying in amount and duration across a range of biogeographical scenarios provide a powerful tool for estimating how drought will affect future ecosystems. Past experimental work has been focused on the manipulation of meteorological drought: Rainout shelters are used to reduce precipitation inputs into the soil. This work has been instrumental in our ability to predict the expected effects of altered rainfall. But what about the nonrainfall components of drought? We review recent literature on the co-occurring and sometimes divergent impacts of atmospheric drying and meteorological drying. We discuss how manipulating meteorological drought or rainfall alone may not predict future changes in plant productivity, composition, or species interactions that result from climate change induced droughts. We make recommendations for how to improve these experiments using manipulations of relative humidity. 

Abstract Plant traits can be helpful for understanding grassland ecosystem responses to climate extremes, such as severe drought. However, intercontinental comparisons of how drought affects plant functional traits and ecosystem functioning are rare. The Extreme Drought in Grasslands experiment (EDGE) was established across the major grassland types in East Asia and North America (six sites on each continent) to measure variability in grassland ecosystem sensitivity to extreme, prolonged drought. At all sites, we quantified community‐weighted mean functional composition and functional diversity of two leaf economic traits, specific leaf area and leaf nitrogen content, in response to drought. We found that experimental drought significantly increased community‐weighted means of specific leaf area and leaf nitrogen content at all North American sites and at the wetter East Asian sites, but drought decreased community‐weighted means of these traits at moderate to dry East Asian sites. Drought significantly decreased functional richness but increased functional evenness and dispersion at most East Asian and North American sites. Ecosystem drought sensitivity (percentage reduction in aboveground net primary productivity) positively correlated with community‐weighted means of specific leaf area and leaf nitrogen content and negatively correlated with functional diversity (i.e., richness) on an intercontinental scale, but results differed within regions. These findings highlight both broad generalities but also unique responses to drought of community‐weighted trait means as well as their functional diversity across grassland ecosystems. 

Abstract Plant populations are limited by resource availability and exhibit physiological trade‐offs in resource acquisition strategies. These trade‐offs may constrain the ability of populations to exhibit fast growth rates under water limitation and high cover of neighbours. However, traits that confer drought tolerance may also confer resistance to competition. It remains unclear how fitness responses to these abiotic conditions and biotic interactions combine to structure grassland communities and how this relationship may change along a gradient of water availability.To address these knowledge gaps, we estimated the low‐density growth rates of populations in drought conditions with low neighbour cover and in ambient conditions with average neighbour cover for 82 species in six grassland communities across the Central Plains and Southwestern United States. We assessed the relationship between population tolerance to drought and resistance to competition and determined if this relationship was consistent across a precipitation gradient. We also tested whether population growth rates could be predicted using plant functional traits.Across six sites, we observed a positive correlation between low‐density population growth rates in drought and in the presence of interspecific neighbours. This positive relationship was particularly strong in the grasslands of the northern Great Plains but weak in the most xeric grasslands. High leaf dry matter content and a low (more negative) leaf turgor loss point were associated with high population growth rates in drought and with neighbours in most grassland communities.Synthesis: A better understanding of how both biotic and abiotic factors impact population fitness provides valuable insights into how grasslands will respond to extreme drought. Our results advance plant strategy theory by suggesting that drought tolerance increases population resistance to interspecific competition in grassland communities. However, this relationship is not evident in the driest grasslands, where above‐ground competition is likely less important. Leaf dry matter content and turgor loss point may help predict which populations will establish and persist based on local water availability and neighbour cover, and these predictions can be used to guide the conservation and restoration of biodiversity in grasslands. 

Abstract The predicted intensification of the North American Monsoon is expected to alter growing season rainfall patterns in the southwestern United States. These patterns, which have historically been characterized by frequent small rain events, are anticipated to shift towards a more extreme precipitation regime consisting of fewer, but larger rain events. Furthermore, human activities are contributing to increased atmospheric nitrogen deposition throughout this dryland region.Alterations in rainfall size and frequency, along with changes in nitrogen availability, are likely to have significant consequences for above‐ground net primary production (ANPP) and plant community dynamics in drylands. The conceptual bucket model predicts that a shift towards fewer, but larger rain events could promote greater rates of ANPP in these regions by maintaining soil moisture availability above drought stress thresholds for longer periods during the growing season. However, only a few short‐term studies have tested this hypothesis, and none have explored the interaction between altered rainfall patterns and nitrogen enrichment.To address this knowledge gap, we conducted a 14‐year rainfall addition and nitrogen fertilization experiment in a northern Chihuahuan Desert grassland to explore the long‐term impacts of changes in monsoon rainfall size and frequency, along with chronic nitrogen enrichment, on ANPP (measured as peak biomass) and plant community dynamics.Contrary to bucket model predictions, small frequent rain events promoted comparable rates of ANPP to large infrequent rain events in the absence of nitrogen enrichment. It was only when nitrogen limitation was alleviated that large infrequent rain events resulted in the greatest ANPP. Furthermore, we found that nitrogen enrichment had the greatest impact on plant community composition under the small frequent rainfall regime.Synthesis. Our long‐term field experiment highlights limitations of the bucket model by demonstrating that water and nitrogen availability sequentially limit dryland ecological processes. Specifically, our findings suggest that while water availability is the primary limiting factor for above‐ground net primary production in these ecosystems, nitrogen limitation becomes increasingly important when water is not limiting. Moreover, our findings reveal that small frequent rain events play an important but underappreciated role in driving dryland ecosystem dynamics. 

Abstract Asexual reproduction plays a fundamental role in the structure, dynamics and persistence of perennial grasslands. Thus, assessing how asexual reproductive traits of plant communities respond to drought may be key for understanding grassland resistance to drought and recovery following drought.Here, we quantified three asexual reproductive traits (i.e. above‐ground tiller abundance, below‐ground bud abundance and the ratio of tillers to buds) during a 4‐year severe drought and a 2‐year drought recovery period in four grasslands that spanned an aridity gradient in northern China. We also assessed the relationship between these traits and the resistance and recovery of above‐ground net primary productivity (ANPP).We found that drought had limited and largely inconsistent effects on asexual reproduction among drought and recovery years and grasslands overall. Drought increased tiller abundance in the first treatment year and reduced bud banks by the fourth treatment year across grasslands. However, neither of the three asexual reproductive traits were correlated with drought resistance of ANPP. Drought legacies differed among the four grasslands with positive, negative and no legacies evident for the three asexual reproductive traits, and no clear relationship with aridity. Bud banks and tiller to bud ratio decreased and increased, respectively, in the first recovery year, but not in the second recovery year. In contrast to drought resistance, community bud abundance was strongly related to recovery, such that communities with higher bud abundance had greater ANPP recovery following drought.Synthesis. These results suggest that asexual reproductive traits may be important drivers of ecosystem recovery after drought, but that variable responses of these asexual reproduction traits during drought complicates predictions of overall grassland responses. 

Abstract Grasslands are subject to climate change, such as severe drought, and an important aspect of their functioning is temporal stability in response to extreme climate events. Previous research has explored the impacts of extreme drought and post‐drought periods on grassland stability, yet the mechanistic pathways behind these changes have rarely been studied.Here, we implemented an experiment with 4 years of drought and 3 years of recovery to assess the effects of drought and post‐drought on the temporal stability of above‐ground net primary productivity (ANPP) and its underlying mechanisms. To do so, we measured community‐weighted mean (CWM) of six plant growth and nine seed traits, functional diversity, population stability and species asynchrony across two cold, semiarid grasslands in northern China. We also performed piecewise structural equation models (SEMs) to assess the relationships between ANPP stability and its underlying mechanisms and how drought and post‐drought periods alter the relative contribution of these mechanisms to ANPP stability.We found that temporal stability of ANPP was not reduced during drought due to grasses maintaining productivity, which compensated for increased variation of forb productivity. Moreover, ANPP recovered rapidly after drought, and both grasses and forbs contributed to community stability during the post‐drought period. Overall, ANPP stability decreased during the combined drought and post‐drought periods because of rapid changes in ANPP from drought to post‐drought. SEMs revealed that the temporal stability of ANPP during drought and post‐drought periods was modulated by functional diversity and community‐weighted mean traits directly and indirectly by altering species asynchrony and population stability. Specifically, the temporal stability of ANPP was positively correlated with functional divergence of plant communities. CWMs of seed traits (e.g. seed width and thickness), rather than plant growth traits (e.g. specific leaf area and leaf nutrient content), stabilized grassland ANPP. Productivity of plant communities with large and thick seeds was less sensitive to precipitation changes over time.These results emphasize the importance of considering both the functional trait distribution among species and seed traits of dominant species since their combined effects can stabilize ecosystem functions under global climate change scenarios. Read the freePlain Language Summaryfor this article on the Journal blog. 

Abstract Plant traits are useful proxies of plant strategies and can influence community and ecosystem responses to climate extremes, such as severe drought. Few studies, however, have investigated both the immediate and lagged effects of drought on community‐weighted mean (CWM) plant traits, with even less research on the relative roles of interspecific vs. intraspecific trait variability in such responses.We experimentally reduced growing season precipitation by 66% in two cold‐semi‐arid grassland sites in northern China for four consecutive years to explore the drought resistance of CWM traits as well as their recovery 2 years following the drought. In addition, we isolated the effects of both interspecific and intraspecific trait variability on shifts in CWM traits.At both sites, we observed significant effects of drought on interspecific and intraspecific trait variability which, in some cases, led to significant changes in CWM traits. For example, drought led to reduced CWM plant height and leaf phosphorous content, but increased leaf carbon content at both sites, with responses primarily due to intraspecific trait shifts. Surprisingly, these CWM traits recovered completely 2 years after the extreme drought. Intraspecific trait variability influenced CWM traits via both positive and negative covariation with interspecific trait variability during drought and recovery phases.These findings highlight the important role of interspecific and intraspecific trait variability in driving the response and recovery of CWM traits following extreme, prolonged drought. Read the freePlain Language Summaryfor this article on the Journal blog. 

Abstract Grasslands are expected to experience droughts of unprecedented frequency and magnitude in the future. Characterizing grassland responses and recovery from drought is therefore critical to predict the vulnerability of grassland ecosystems to climate change. Most previous studies have focused on ecosystem responses during drought while investigations of post‐drought recovery are rare. Few studies have used functional traits, and in particular bud or clonal traits, to explore the mechanisms underlying grassland responses to and recovery from drought.To address this issue, we experimentally imposed a four‐year drought in a C₃‐dominated grassland in northeastern China and monitored recovery for 3 years post‐drought. We investigated the immediate and legacy effects of drought on total above‐ground net primary productivity (ANPP), ANPP of functional groups (rhizomatous grasses, bunch grasses and forbs), and how the legacy effects were driven by plant species diversity, clonal traits and vegetative traits.We found that drought progressively reduced total ANPP over the 4‐year period. The reductions in total ANPP in the first and third drought years were caused by the decrease in ANPP of bunch grasses only, while that of the second year was caused by declines in ANPP of bunch grasses and forbs, and the fourth year decline was linked to all three functional groups. The post‐drought recovery of ANPP, which occurred despite the continued loss of plant species diversity, was mainly driven by rapid recovery of rhizomatous and bunch grasses, which compensated for the slow response by forbs. The rapid post‐drought recovery of these grasses can be attributed to their relatively large, intact bud and shoot densities post‐drought, as well as the recovery of plant height and specific leaf area. The rapid recovery of grasses possibly restricted the growth and distribution of forbs, resulting in reduced forb ANPP and, consequently, lower species diversity during the recovery period.Synthesis. These results highlight the potential for positive legacy effects of drought on ANPP as well as the important and complementary roles of plant reproductive and vegetative traits in mediating ecosystem recovery from drought in a C₃‐dominated grassland. 

Abstract Seeds provide the basis of genetic diversity in perennial grassland communities and their traits may influence ecosystem resistance to extreme drought. However, we know little about how drought effects the community functional composition of seed traits and the corresponding implications for ecosystem resistance to drought.We experimentally removed 66% of growing season precipitation for 4 years across five arid and semi‐arid grasslands in northern China and assessed how this multi‐year drought impacted community‐weighted means (CWMs) of seed traits, seed trait functional diversity and above‐ground net primary productivity (ANPP).Experimental drought had limited effects on CWM traits and the few effects that did occur varied by site and year. For three separate sites, and in different years, drought reduced seed length and phosphorus content but increased both seed and seed‐coat thickness. Additionally, drought led to increased seed functional evenness, divergence, dispersion and richness, but only in some sites, and mostly in later years following cumulative effects of water limitation. However, we observed a strong negative relationship between drought‐induced reductions in ANPP and CWMs of seed‐coat thickness, indicating that a high abundance of dominant species with thick seeds may increase ecosystem resistance to drought. Seed trait functional diversity was not significantly predictive of ANPP, providing little evidence for a diversity effect.Our results suggest that monitoring community composition with a focus on seed traits may provide a valuable indicator of ecosystem resistance to future droughts despite inconsistent responses of seed trait composition overall. This highlights the importance of developing a comprehensive seed and reproductive traits database for arid and semi‐arid grassland biomes. Read the freePlain Language Summaryfor this article on the Journal blog. 

Root production influences carbon and nutrient cycles and subsidizes soil biodiversity. However, the long‐term dynamics and drivers of belowground production are poorly understood for most ecosystems. In drylands, fire, eutrophication, and precipitation regimes could affect not only root production but also how roots track interannual variability in climate. We manipulated the intra‐annual precipitation regime, soil nitrogen, and fire in four common Chihuahuan Desert ecosystem types (three grasslands and one shrubland) in New Mexico, USA, where the 100‐year record indicates both long‐term drying and increasing interannual variability in aridity. First, we evaluated how root production tracked aridity over 10–17 years using climate sensitivity functions, which quantify long‐term, nonlinear relationships between biological processes and climate. Next, we determined the degree to which perturbations by fire, nitrogen addition or intra‐annual rainfall altered the sensitivity of root production to both mean and interannual variability in aridity. All ecosystems had nonlinear climate sensitivities that predicted declines in production with increases in the interannual variance of aridity. However, root production was the most sensitive to aridity in Chihuahuan Desert shrubland, with reduced production under drier and more variable aridity. Among the perturbations, only fire altered the sensitivity of root production to aridity. Root production was more than twice as sensitive to declines with aridity following prescribed fire than in unburned conditions. Neither the intra‐annual seasonal rainfall regime nor chronic nitrogen fertilization altered the sensitivity of roots to aridity. Our results yield new insight into how dryland plant roots respond to climate change. Our comparison of dryland ecosystems of the northern Chihuahuan Desert predicted that root production in shrublands would be more sensitive to future climates that are drier and more variable than root production in dry grasslands. Field manipulations revealed that fire could amplify the climate sensitivity of dry grassland root production, but in contrast, the climate sensitivity of root production was largely resistant to changes in the seasonal rainfall regime or increased soil fertilization.