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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 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. 

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. 

Abstract Water‐limited ecosystems are highly sensitive to not only precipitation amount, but also precipitation pattern, particularly variability in the size and timing of growing season rainfall events. Both rainfall event size and timing are expected to be altered by climate change, but the relative responses of dryland ecosystems to changes in rainfall event size versus timing have not been resolved. Here, we disentangle the effects of these different aspects of precipitation pattern on ecosystem dynamics.We experimentally assessed how these two aspects of rainfall variability impacted a semi‐arid grassland ecosystem by altering an ambient precipitation pattern to eliminate variability in (a) rainfall event size (all events were made the same size), (b) rainfall event timing (all events were uniformly spaced in time) and (c) both. Total precipitation amount was constant for all treatments. We measured responses of soil moisture, ecosystem carbon flux (e.g. net primary production and soil CO₂flux), plant community composition and physiological responses of the dominant C₄grass,Bouteloua gracilis.Removing variability in rainfall event size altered ecosystem dynamics more than a pattern of uniform event timing, but the largest impact occurred when variability in both were removed. Notably, eliminating variability in both event size and timing increased above‐ground net primary productivity by 23%, consistent with reduced water stress in the dominant C₄grass, while also reducing seasonal variability in soil CO₂flux by 35%, reflecting lower seasonal variability in soil moisture.Synthesis. Unique responses to different aspects of precipitation variability highlight the complexity of predicting how dryland ecosystems will be affected by climate change‐induced shifts in rainfall patterns. Our results provide novel support for the key roles of rainfall event size and timing, in addition to total precipitation amount, as determinants of ecosystem function. 

During the 1930s Dust Bowl drought in the central United States, species with the C₃photosynthetic pathway expanded throughout C₄-dominated grasslands. This widespread increase in C₃grasses during a decade of low rainfall and high temperatures is inconsistent with well-known traits of C₃vs. C₄pathways. Indeed, water use efficiency is generally lower, and photosynthesis is more sensitive to high temperatures in C₃than C₄species, consistent with the predominant distribution of C₃grasslands in cooler environments and at higher latitudes globally. We experimentally imposed extreme drought for 4 y in mixed C₃/C₄grasslands in Kansas and Wyoming and, similar to Dust Bowl observations, also documented three- to fivefold increases in C₃/C₄biomass ratios. To explain these paradoxical responses, we first analyzed long-term climate records to show that under nominal conditions in the central United States, C₄grasses dominate where precipitation and air temperature are strongly related (warmest months are wettest months). In contrast, C₃grasses flourish where precipitation inputs are less strongly coupled to warm temperatures. We then show that during extreme drought years, precipitation–temperature relationships weaken, and the proportion of precipitation falling during cooler months increases. This shift in precipitation seasonality provides a mechanism for C₃grasses to respond positively to multiyear drought, resolving the Dust Bowl paradox. Grasslands are globally important biomes and increasingly vulnerable to direct effects of climate extremes. Our findings highlight how extreme drought can indirectly alter precipitation seasonality and shift ecosystem phenology, affecting function in ways not predictable from key traits of C₃and C₄species.