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Abstract Climate change is increasing the frequency and severity of droughts globally, and grasslands are particularly vulnerable to such hydrological extremes. Drought effects at the ecosystem scale have been assessed both experimentally and through the study of naturally occurring drought, with emerging evidence that the magnitude of drought effects may vary depending on the approach used. We took advantage of a decadal study of four grasslands to directly contrast responses of aboveground net primary productivity (ANPP) to simulated vs. natural drought. The grasslands spanned a ~ threefold mean annual precipitation gradient (335–857 mm) and were all subjected to a natural 1-year drought (~ 40% reduction in precipitation from the long-term mean) and a 4 year experimental drought (~ 50% precipitation reduction). We expected that the 4 year drought would reduce ANPP more, and that post-drought recovery would be delayed, compared to the 1-year drought. We found instead that the short-term natural drought reduced ANPP more strongly than the simulated drought in all grasslands (~ 10 to ~ 50%) likely due to the co-occurrence of higher temperatures and vapor pressure deficits with reduced precipitation. Post-drought recovery was site specific and each site differed in their recovery from the natural and experimental droughts. These results align with past analyses that experiments that only manipulate soil moisture likely underestimate the magnitude of natural drought events. However, experiments can provide valuable insight into the relative sensitivity of ecosystems to reduced precipitation and soil moisture, a key aspect of drought. 

ABSTRACT Ecological stability plays a crucial role in determining the sustainability of ecosystem functioning and nature's contribution to people. Although the disruptive effects of extreme drought on ecosystem structure and functions are widely recognized, their effect on the stability of above‐ and belowground productivity remains understudied. We assessed the effects of drought on ecosystem stability using a 3‐year drought experiment established in six Eurasian steppe grasslands. The treatments imposed included ambient precipitation, chronic drought (66% reduction in precipitation throughout the growing season), and intense drought (complete exclusion of precipitation for two months during the growing season). We found that drought, irrespective of how it was imposed, reduced the stability of aboveground net primary productivity (ANPP) but had little impact on belowground net primary productivity (BNPP) stability. Reduced ANPP stability under drought was primarily attributed to changes in subordinate species stability, with mean annual precipitation (MAP) and its variability, historical drought frequency, and the aridity index (AI) also influencing responses to extreme drought. In contrast, BNPP stability was not related to any community factor investigated, but it was influenced by MAP variability and AI. Our findings that above‐ and belowground productivity stability in grasslands are differentially sensitive to multi‐year extreme drought under both common (MAP and AI) as well as unique drivers (plant community changes) highlight the complexity of predicting carbon cycle dynamics as hydrological extremes become more severe. 

Summary It has been 60 years since the discovery of C₄photosynthesis, an event that rewrote our understanding of plant adaptation, ecosystem responses to global change, and global food security. Despite six decades of research, one aspect of C₄photosynthesis that remains poorly understood is how the pathway fits into the broader context of adaptive trait spectra, which form our modern view of functional trait ecology. The C₄CO₂‐concentrating mechanism supports a general C₄plant phenotype capable of fast growth and high resource‐use efficiencies. The fast‐efficient C₄phenotype has the potential to operate at high productivity rates, while allowing for less biomass allocation to root production and nutrient acquisition, thereby providing opportunities for the evolution of novel trait covariances and the exploitation of new ecological niches. We propose the placement of the C₄fast‐efficient phenotype near the acquisitive pole of the world‐wide leaf economic spectrum, but with a pathway‐specific span of trait space, wherein selection shapes both acquisitive and conservative adaptive strategies. A trait‐based perspective of C₄photosynthesis will open new paths to crop improvement, global biogeochemical modeling, the management of invasive species, and the restoration of disturbed ecosystems, particularly in grasslands. 

ABSTRACT Extreme droughts are intensifying, yet their impact on temporal variability of grassland functioning and its drivers remains poorly understood. We imposed a 6‐year extreme drought in two semiarid grasslands to explore how drought influences the temporal variability of ANPP and identify potential stabilising mechanisms. Drought decreased ANPP while increasing its temporal variability across grasslands. In the absence of drought, ANPP variability was strongly driven by the dominant plant species (i.e., mass‐ratio effects), as captured by community‐weighted traits and species stability. However, drought decreased the dominance of perennial grasses, providing opportunities for subordinate species to alter the stability of productivity through compensatory dynamics. Specifically, under drought, species asynchrony emerged as a more important correlate of ANPP variability than community‐weighted traits or species stability. Our findings suggest that in grasslands, prolonged, extreme droughts may decrease the relative contribution of mass‐ratio effects versus compensatory dynamics to productivity stability by reducing the influence of dominant species. 

Extreme droughts generally decrease productivity in grassland ecosystems1,2,3 with negative consequences for nature’s contribution to people4,5,6,7. The extent to which this negative effect varies among grassland types and over time in response to multi-year extreme drought remains unclear. Here, using a coordinated distributed experiment that simulated four years of growing-season drought (around 66% rainfall reduction), we compared drought sensitivity within and among six representative grasslands spanning broad precipitation gradients in each of Eurasia and North America—two of the Northern Hemisphere’s largest grass-dominated regions. Aboveground plant production declined substantially with drought in the Eurasian grasslands and the effects accumulated over time, while the declines were less severe and more muted over time in the North American grasslands. Drought effects on species richness shifted from positive to negative in Eurasia, but from negative to positive in North America over time. The differing responses of plant production in these grasslands were accompanied by less common (subordinate) plant species declining in Eurasian grasslands but increasing in North American grasslands. Our findings demonstrate the high production sensitivity of Eurasian compared with North American grasslands to extreme drought (43.6% versus 25.2% reduction), and the key role of subordinate species in determining impacts of extreme drought on grassland productivity. 

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 Despite asymmetric competition and a wide array of functional similarities, two ecologically important C₄perennial grasses,Andropogon gerardiiandSorghastrum nutans, frequently codominate areas of the mesic tallgrass prairie of the US Great Plains. A subtle difference in their vegetative reproduction strategies may play a role in preventing the exclusion ofS. nutans, the presumed weaker competitor in such regions.WhileA. gerardiivegetative tiller densities peak in the early growing season and decline thereafter (determinate recruitment), those ofS. nutansmay continue to increase throughout the growing season (indeterminate recruitment), providing a potential avenue for recovery from more intensive early season competition. However, until now these patterns have only been informally observed in the field.We examined the year‐to‐year consistency of growing season vegetative tiller dynamics (measured as seasonal change in tiller densities) of each grass species from an intact tallgrass prairie in Kansas – a site within the core of both species' distributions – over a period of 8 years. Then, to investigate environmental effects on these dynamics, we examined whether they differ across a Kansas landscape varying in topography, fire management regimes, and the abundances of the study species. Finally, we expanded the investigation of environmental effects on growing season tiller dynamics by observing them at the periphery of the species' distributions in central Colorado, where climatic conditions are dryer and the study species' abundances are reduced.Synthesis. We found that the tiller densities ofA. gerardiidecline within seasons with striking consistency regardless of spatio‐temporal scale or environmental factors (topography and fire regimes). In contrast, we found the seasonal dynamics ofS. nutanstiller densities were dependent on environmental factors, with seasonal tiller density increases occurring only within the Kansas populations but not consistent between years. These observations lay the groundwork for establishing differences in tiller recruitment determinacy as a potentially important yet underappreciated mechanism for promoting coexistence and codominance among perennial plant species. 

Ecosystems are experiencing changing global patterns of mean annual precipitation (MAP) and enrichment with multiple nutrients that potentially colimit plant biomass production. In grasslands, mean aboveground plant biomass is closely related to MAP, but how this relationship changes after enrichment with multiple nutrients remains unclear. We hypothesized the global biomass–MAP relationship becomes steeper with an increasing number of added nutrients, with increases in steepness corresponding to the form of interaction among added nutrients and with increased mediation by changes in plant community diversity. We measured aboveground plant biomass production and species diversity in 71 grasslands on six continents representing the global span of grassland MAP, diversity, management, and soils. We fertilized all sites with nitrogen, phosphorus, and potassium with micronutrients in all combinations to identify which nutrients limited biomass at each site. As hypothesized, fertilizing with one, two, or three nutrients progressively steepened the global biomass–MAP relationship. The magnitude of the increase in steepness corresponded to whether sites were not limited by nitrogen or phosphorus, were limited by either one, or were colimited by both in additive, or synergistic forms. Unexpectedly, we found only weak evidence for mediation of biomass–MAP relationships by plant community diversity because relationships of species richness, evenness, and beta diversity to MAP and to biomass were weak or opposing. Site-level properties including baseline biomass production, soils, and management explained little variation in biomass–MAP relationships. These findings reveal multiple nutrient colimitation as a defining feature of the global grassland biomass–MAP relationship. 

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.