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The frequency, duration and magnitude of drought are projected to increase in the next few decades, with consequences for community composition and nutrient utilization of plants. Nutrient resorption, a process facilitating plant nutrient conservation, is a crucial driver for ecosystem nutrient cycling. It remains unknown how would extreme drought affect nutrient resorption of plant communities through changing community composition.We examined the contributions of different processes (richness and identity effect of species losses and species gains, as well as the context dependent effect) related with community compositional changes to the alterations of community‐level nutrient resorption efficiency after 4‐year treatments of chronic drought (66% rainfall reduction from May to August) and intense drought (100% rainfall reduction in June and July) in a temperate steppe in Hulun Buir, Inner Mongolia, China.Both chronic and intense drought significantly reduced nutrient resorption efficiency in most of the common species, but did not affect that at the community level. Drought‐induced losses of species made positive contribution to the community‐level nutrient resorption, as they generally had lower nutrient resorption efficiency. Such positive contribution balanced the negative effect of drought on the nutrient resorption of the common species.Our results indicate that drought‐induced changes in community composition, especially those species being lost, were important in mediating the community‐level responses of nutrient resorption to extreme drought in grasslands. Read the free Plain Language Summary for this article on the Journal blog. [ABSTRACT FROM AUTHOR]

Biodiversity can stabilise productivity through different mechanisms, such as asynchronous species responses to environmental variability and species stability. Global changes, like intensified drought, could negatively affect species richness, species asynchrony and species stability, but it is unclear how changes in these mechanisms will affect the stability of above‐ground primary productivity (ANPP) across ecosystems.We studied the effects of a 4‐year extreme drought on ANPP stability and the underlying mechanisms (species richness, species asynchrony and species stability) across six grasslands in Northern China. We also assessed the relative importance of these mechanisms in determining ANPP stability under extreme drought.We found that extreme drought decreased ANPP stability, species richness, species asynchrony and species stability across the six grasslands. However, structural equation modelling revealed that species asynchrony, not species richness or species stability, was the most important mechanism promoting stability of ANPP, regardless of drought across the six grasslands.Synthesis. Our results suggest that species asynchrony, not species richness and species stability, consistently buffers ecosystem stability against extreme drought across and within grasslands spanning a broad precipitation gradient. Thus, species asynchrony may be a more general mechanism for promoting stability of ANPP in grasslands in the face of intensified drought. [ABSTRACT FROM AUTHOR]

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 free Plain Language Summary for this article on the Journal blog. [ABSTRACT FROM AUTHOR]

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 FROM AUTHOR]

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 FROM AUTHOR]

Climate change is expected to increase the magnitude and frequency of extreme drought in most grassland ecosystems. Exploring the responses of below‐ground bud banks and their relationships with above‐ground plant structure and drought is need to explain how climate change will impact grassland ecosystems. However, studies on the response of community‐scale bud and shoot densities to experimental drought along an aridity gradient are rare.We experimentally removed 66% of growing season precipitation for 4 years in three temperate grasslands that spanned an aridity gradient in northern China. We quantified the legacy effects of drought on grass, forb and total community below‐ground bud density, above‐ground shoot density and the ratio of bud to shoot density 1 year following treatment.Below‐ground bud density was lowest at the highest aridity site for the entire community, while above‐ground shoot density was highest at the medium aridity site. Below‐ground bud and above‐ground shoot densities were the lowest at the high aridity site for grasses but the highest for forbs at this site. Bud:shoot ratios decreased with increasing aridity for grasses, yet remained constant for forbs along the aridity gradient. Below‐ground bud density in drought plots remained lower than controls a year following drought at each site. Experimental drought did not alter the below‐ground bud bank for grasses but decreased forb bud banks across sites. Experimental drought had little legacy effects on above‐ground shoot density and bud:shoot ratios for grasses, forbs and the total community at each site.Our results suggest that grass and forb bud banks can differ in their responses to both multi‐year drought along an aridity gradient, and that bud limitation for shoot generation may increase as grasslands get drier. Bud bank responses to climate will impact plant community functioning and resilience. Thus, incorporating bud bank dynamics will improve projections of grassland ecosystems under future climate change. Read the free Plain Language Summary for this article on the Journal blog. [ABSTRACT FROM AUTHOR]

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 free Plain Language Summary for this article on the Journal blog. [ABSTRACT FROM AUTHOR]

Recurrent droughts are an inevitable consequence of climate change, yet how grasslands respond to such events is unclear. We conducted a 6‐year rainfall manipulation experiment in a semiarid grassland that consisted of an initial 2‐year drought (2015–2016), followed by a recovery period (2017–2018) and, finally, a second 2‐year drought (2019–2020). In each year, we estimated aboveground net primary productivity (ANPP), species richness, community‐weighted mean (CWM) plant traits, and several indices of functional diversity. The initial drought led to reduced ANPP, which was primarily driven by limited growth of forbs in the first year and grasses in the second year. Total ANPP completely recovered as the rapid recovery of grass productivity compensated for the slow recovery of forb productivity. The subsequent drought led to a greater reduction in total ANPP than the initial drought due to the greater decline of both grass and forb productivity. The structural equation models revealed that soil moisture influenced ANPP responses directly during the initial drought, and indirectly during the subsequent drought by lowering functional diversity, which resulted in reduced total ANPP. Additionally, ANPP was positively influenced by CWM plant height and leaf nitrogen during the recovery period and recurrent drought, respectively. Overall, the greater impact of the second drought on ecosystem function than the initial drought, as well as the underlying differential mechanism, underscores the need for an understanding of how increased drought frequency may alter semiarid grassland functioning. [ABSTRACT FROM AUTHOR]

Drought events induced by global climate change strongly modify ecosystem structure and function in grasslands. Higher plant biodiversity can enhance ecosystem resistance to drought; however, the potential effects of drought on the relationships between above‐ground productivity (ANPP) and biodiversity along a natural aridity gradient remain poorly understood.Here, we studied the effects of experimental drought on plant community structure and ANPP during the two different dry years (2015 and 2017) spanning three temperate steppe sites with low‐, medium‐ and high‐aridity levels. We also examined the effects of drought and aridity on the associations of plant species (richness and abundance), functional (community‐weighted trait means and functional dispersion) and phylogenetic diversity (evolutionary lineages) with ANPP. A structural equation modelling (SEM) was used to elucidate how the drought affected ANPP by altering biodiversity attributes.We found that experimental drought shaped the biodiversity–ANPP relationships, and the biodiversity–ANPP relationships differed among three sites. At the low‐aridity site, ANPP was correlated with plant functional traits, whereas ANPP was correlated with functional dispersion, species and phylogenetic diversity at the medium‐aridity site. At the high‐aridity site, ANPP was correlated with species richness, functional traits and phylogenetic diversity. Compared with 2015, natural drought in 2017 strengthened the associations of species diversity, functional dispersion, leaf dry matter content and leaf nitrogen content with ANPP across three sites. SEM further showed that both experimental and natural drought directly decreased ANPP and indirectly decreased ANPP by reducing community‐weighted mean of plant height.Synthesis. These results suggest that the aridity increase or drought occurrence is likely to induce the shifts in the associations of ANPP with species and functional diversity, highlighting the important role of water availability or precipitation conditions in regulating the biodiversity–ANPP relationships in temperate steppes. Overall, functional traits related to plant size play a crucial role in modulating the response of productivity to drought across three steppe sites. Thus, to buffer the negative effects of drought on steppe ANPP should pay attention to protect the species with large plant size. Read the free Plain Language Summary for this article on the Journal blog. [ABSTRACT FROM AUTHOR]

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 free Plain Language Summary for this article on the Journal blog. [ABSTRACT FROM AUTHOR]

Summary: Climate extremes pose enormous threats to natural ecosystems. Arbuscular mycorrhizal (AM) fungi are key plant symbionts that can affect plant community dynamics and ecosystem stability. However, knowledge about how AM fungal communities respond to climate extremes in natural ecosystems remains elusive.Based on a grassland extreme drought experiment in Inner Mongolia, we investigated the response of AM fungal communities to extreme drought in association with plant communities. The experiment simulated two types of extreme drought (chronic/intense) of once‐in‐20‐year occurrence.AM fungal richness and community composition exhibited high sensitivity to extreme drought and were more sensitive to intense drought than chronic drought. This community sensitivity (i.e. decline in richness and shifts in community composition) of AM fungi can be jointly explained by soil moisture, plant richness, and aboveground productivity. Notably, the robustness of the plant–AM fungal community co‐response increased with drought intensity.Our results indicate that AM fungal communities are sensitive to climate extremes, and we propose that the plant community mediates AM fungal community responses. Given the ubiquitous nature of AM associations, their climate sensitivity may have profound consequences on plant communities and ecosystem stability under climate change. [ABSTRACT FROM AUTHOR]

Extreme drought decreases aboveground net primary production (ANPP) in most grasslands, but the magnitude of ANPP reductions varies especially in C3‐dominated grasslands. Because the mechanisms underlying such differential ecosystem responses to drought are not well resolved, we experimentally imposed an extreme 4‐yr drought (2015–2018) in two C3 grasslands that differed in aridity. These sites had similar annual precipitation and dominant grass species (Leymus chinensis) but different annual temperatures and thus water availability. Drought treatments differentially affected these two semiarid grasslands, with ANPP of the drier site reduced more than at the wetter site. Structural equation modeling revealed that community‐weighted means for some traits modified relationships between soil moisture and ANPP, often due to intraspecific variation. Specifically, drought reduced community mean plant height at both sites, resulting in a reduction in ANPP beyond that attributable to reduced soil moisture alone. Higher community mean leaf carbon content enhanced the negative effects of drought on ANPP at the drier site, and ANPP–soil‐moisture relationships were influenced by soil C:N ratio at the wetter site. Importantly, neither species richness nor functional dispersion were significantly correlated with ANPP at either site. Overall, as expected, soil moisture was a dominant, direct driver of ANPP response to drought, but differential sensitivity to drought in these two grasslands was also related to soil fertility and plant traits. [ABSTRACT FROM AUTHOR]

Drought is viewed as a naturally recurring phenomenon in many Australian agricultural systems. Identifying regional changes in frequency and severity of drought induced by climate change is required to develop regionally specific adaptation strategies. In this study, we provided a first look at the impacts of climate change on 21st century drought characteristics over the New South Wales wheat belt of southeastern Australia. These impacts were assessed from an ensemble of 28 statistical downscaled global climate models under representative concentration pathway (RCP8.5). A modified relative standardized precipitation and evapotranspiration index (rSPEI) at the seasonal scale (3 months) was used to analyse temporal and spatial changes in drought. Results indicated that there was a tendency towards more frequent and severe winter–spring droughts over the study area. Moreover, winter–spring drought prone areas were expected to expand from west to east. Until the end of the 21st century, more than half the wheat belt would be vulnerable to winter–spring drought. The combined effects of reduced precipitation and increased temperature during future winter and spring seasons were the main reasons causing these changes of drought. In addition, summer and autumn droughts would have both slight temporal and spatial changes across the study region. This study also revealed that traditionally dry areas would likely experience an increased frequency of drought compared to wetter areas when subjected to a same increase in temperature or decrease in precipitation. Furthermore, the western part of the wheat belt might be unsuitable for winter crops in the future, or at least exposed to an increased risk of variable yield and would require a gradual transformation which might include summer crops or pastures. Investments in cropping land should be focused on the east part of the wheat belt to achieve more consistent financial returns. Drought prone areas were identified in the study region using the modified drought index, rSPEI. Spring and winter drought prone areas were likely to expand remarkably. By the end of the 21st century, over half of the study area was at a high risk of suffering spring and winter drought. Summer and autumn drought prone areas were primarily in the southwest and northwest and might only experience slight changes. [ABSTRACT FROM AUTHOR]