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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.more » « less
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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.more » « less
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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.more » « less
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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.more » « less
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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.more » « less
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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.more » « less
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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.more » « less
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Global climate change is expected to cause more frequent extreme droughts in many parts of the world. Despite the crucial role of roots in water acquisition and plant survival, our understanding of ecosystem vulnerability to drought is primarily based on aboveground impacts. As return intervals between droughts decrease, root responses to one drought might alter responses to subsequent droughts, but this remains unresolved. We conducted a seven‐year experiment that imposed extreme drought (growing season precipitation reduced 66%) in a mesic grassland. Plots were droughted during years 1–2 (‘Drought 1'), or years 5–6 (‘Drought 2') or both. We quantified root production during year 6 (final year of Drought 2) and year 7 (first year after Drought 2), when all plots received ambient precipitation. We found that repeated drought decreased root mass production more than twice as much as a single drought (−63% versus −27%, respectively, relative to ambient precipitation). Root mass production of the dominant C4grassAndropogon gerardiidid not decrease significantly with either one or two droughts.A. gerardiiroot traits differed from subdominant species on average across all treatments, but drought did not alter root traits of eitherA. gerardiior the subdominant species (collectively). In year 6, root production in plots droughted 4 years ago had not recovered (−21% versus control), but root production recovered in all formerly droughted plots in year 7, when precipitation was above average. Our results highlight the complexity of root responses to drought. Drought‐induced reductions in root production can persist for years after drought and repeated drought can reduce production even further, but this does not preclude rapid recovery of root production in a wet year.more » « less
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Abstract We review results from field experiments that simulate drought, an ecologically impactful global change threat that is predicted to increase in magnitude, extent, duration and frequency. Our goal is to address, from primarily an ecosystem perspective, the questions ‘What have we learned from drought experiments?’ and ‘Where do we go from here?’.Drought experiments are among the most numerous climate change manipulations and have been deployed across a wide range of biomes, although most are conducted in short‐statured, water‐limited ecosystems. Collectively, these experiments have enabled ecologists to quantify the negative responses to drought that occur for most aspects of ecosystem structure and function. Multiple meta‐analyses of responses have also enabled comparisons of relative effect sizes of drought across hundreds of sites, particularly for carbon cycle metrics. Overall, drought experiments have provided strong evidence that ecosystem sensitivity to drought increases with aridity, but that plant traits associated with aridity are not necessarily predictive of drought resistance. There is also intriguing evidence that as drought magnitude or duration increases to extreme levels, plant strategies may shift from drought tolerance to drought escape/avoidance.We highlight three areas where more drought experiments are needed to advance our understanding. First, because drought is intensifying in multiple ways, experiments are needed that address alterations in drought magnitude versus duration, timing and/or frequency (individually and interactively). Second, drivers of drought may be shifting—from precipitation deficits to rising atmospheric demand for water—and disentangling how ecosystems respond to changes in hydrological ‘supply versus demand’ is critical for understanding drought impacts in the future. Finally, more attention should be focussed on post‐drought recovery periods since legacies of drought can affect ecosystem functioning much longer than the drought itself.We conclude with a call for a fundamental shift in the focus of drought experiments from those designed primarily as ‘response experiments’, quantifying the magnitude of change in ecosystem structure and function, to more ‘mechanistic experiments’—those that explicitly manipulate ecological processes or attributes thought to underpin drought responses. Read the freePlain Language Summaryfor this article on the Journal blog.more » « less
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