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Drought is intensifying globally with climate change, creating an urgency to understand ecosystem response to drought both during and after these events end to limit loss of ecosystem functioning. The literature is replete with studies of how ecosystems respond during drought, yet there are far fewer studies focused on ecosystem dynamics after drought ends. Furthermore, while the terms used to describe drought can be variable and inconsistent, so can those that describe ecosystem responses following drought. With this review, we sought to evaluate and create clear definitions of the terms that ecologists use to describe post-drought responses. We found that legacy effects, resilience and recovery were used most commonly with respect to post-drought ecosystem responses, but the definitions used to describe these terms were variable. Based on our review of the literature, we propose a framework for generalizing ecosystem responses after drought ends, which we refer to as 'the post-drought period'. We suggest that future papers need to clearly describe characteristics of the imposed drought, and we encourage authors to use the term post-drought period as a general term that encompasses responses after drought ends and use other terms as more specific descriptors of responses during the post-drought period.

Climate variability and periodic droughts have complex effects on carbon (C) fluxes, with uncertain implications for ecosystem C balance under a changing climate. Responses to climate change can be modulated by persistent effects of climate history on plant communities, soil microbial activity, and nutrient cycling (i.e., legacies). To assess how legacies of past precipitation regimes influence tallgrass prairie C cycling under new precipitation regimes, we modified a long-term irrigation experiment that simulated a wetter climate for25 years. We reversed irrigated and control (ambient precipitation) treatments in some plots and imposed an experimental drought in plots with a history of irrigation or ambient precipitation to assess how climate legacies affect aboveground net primary productivity (ANPP), soil respiration, and selected soil C pools. Legacy effects of elevated precipitation (irrigation) included higher C fluxes and altered labile soil C pools, and in some cases altered sensitivity to new climate treatments. Indeed, decades of irrigation reduced the sensitivity of both ANPP and soil respiration to drought compared with controls. Positive legacy effects of irrigation on ANPP persisted for at least 3 years following treatment reversal, were apparent in both wet and dry years, and were associated with altered plant functional composition. In contrast, legacy effects on soil respiration were comparatively short-lived and did not manifest under natural or experimentally-imposed "wet years," suggesting that legacy effects on CO

The widespread extirpation of megafauna may have destabilized ecosystems and altered biodiversity globally. Most megafauna extinctions occurred before the modern record, leaving it unclear how their loss impacts current biodiversity. We report the long-term effects of reintroducing plains bison ( Bison bison ) in a tallgrass prairie versus two land uses that commonly occur in many North American grasslands: 1) no grazing and 2) intensive growing-season grazing by domesticated cattle ( Bos taurus ). Compared to ungrazed areas, reintroducing bison increased native plant species richness by 103% at local scales (10 m 2 ) and 86% at the catchment scale. Gains in richness continued for 29 y and were resilient to the most extreme drought in four decades. These gains are now among the largest recorded increases in species richness due to grazing in grasslands globally. Grazing by domestic cattle also increased native plant species richness, but by less than half as much as bison. This study indicates that some ecosystems maintain a latent potential for increased native plant species richness following the reintroduction of native herbivores, which was unmatched by domesticated grazers. Native-grazer gains in richness were resilient to an extreme drought, a pressure likely to become more common under future global environmental change.

Climate change is altering precipitation regimes globally, with expectations of intensified precipitation patterns (for example, larger but fewer rainfall events) and more frequent and extreme drought. Both aspects of precipitation change can impact ecosystem function individually, but it is more likely that they will occur in combination. In a central US mesic grassland, we imposed an extreme 2-year drought (growing season precipitation reduced by 66%) on plots with a long-term (16-year) history of exposure to either ambient or intensified precipitation patterns (average threefold increase in event size and threefold decrease in event number during the growing season). While this intensified pattern did not alter total precipitation amount, it generally led to ecosystem responses consistent with a drier environment (for example, reduced soil moisture, aboveground net primary production (ANPP), and soil CO2 flux, but little evidence for altered root biomass). Surprisingly, this history of intensified precipitation patterns did not affect the response of ANPP to the subsequent extreme drought. In contrast, previous exposure to intensified precipitation patterns reduced root production and muted soil CO2 flux responses to rainfall events during drought. Reduced root production in plots experiencing compounded precipitation extremes was driven not by the dominant C4 grass species, Andropogon gerardii, but collectively by the subdominant species in the plant community. Overall, our results reveal that compound changes in precipitation patterns and amount affected this grassland in ways that were less apparent (that is, belowground) than responses to either change individually and significantly reduced ecosystem carbon uptake.

Increasing nitrogen (N) deposition has altered plant communities globally, however the changes in species abundances with short-term vs. long-term N enrichment remains unclear. Stoichiometric homeostasis, quantified by the homoeostatic regulation coefficient (H) is a key trait predictive of plant species dominance and species responses to short-term global changes. It is unknown whether H changes with N enrichment over time, thereby affecting species responses to long-term N addition. Here we investigated three representative plant species how species dominance changed to short-term and long-term N addition with a field N addition experiment (2006–2013) in an Inner Mongolia grassland. Changes in species H with long-term N addition were analyzed using a sand culture experiment, and the correlation between species H and species abundances were explored to address the above research gaps. The abundance of Leymus chinensis decreased with short-term N addition, and increased with long-term N addition, while Chenopodium glaucum exhibited the opposite pattern. Cleistogenes squarrosa was only favored by 1-year N addition, and depressed by two or more years of N addition. The H values of L. chinensis and C. glaucum decreased significantly with long-term N addition, but did not change for C. squarrosa. The H values were significantly related with the abundance both in Control and long-term N addition treatments. Species abundance had opposite responses to short-term vs. long-term N addition. The decrease of H suggested the nutrients use strategy became more progressive, which mediated the responses of species abundances to short- and long-term N addition.

Aims Both extreme drought and insect herbivores can suppress plant growth in grassland communities. However, most studies have examined extreme drought and insects in isolation, and there is reason to believe that insects might alter the ability of grasslands to withstand drought. Unfortunately, few studies have tested the interactive effects of extreme drought and insect herbivores in grassland communities. Methods Here, we tested the drought–herbivore interactions using a manipulative experiment that factorially crossed extreme drought with the exclusion of insect herbivores in a temperate semiarid grassland in Inner Mongolia. Important Findings Our results demonstrated that both extreme drought and insect herbivores separately decreased total plant cover. When combined, insect herbivores reduced the impact of drought on total cover by increasing the relative abundance of drought-resistant dominant species. Our results highlight that the negative effect of extreme drought on total plant cover could be alleviated by maintaining robust insect herbivore communities.

The performance of coordinated distributed experiments designed to compare ecosystem sensitivity to global-change drivers depends on whether they cover a significant proportion of the global range of environmental variables. In the present article, we described the global distribution of climatic and soil variables and quantified main differences among continents. Then, as a test case, we assessed the representativeness of the International Drought Experiment (IDE) in parameter space. Considering the global environmental variability at this scale, the different continents harbor unique combinations of parameters. As such, coordinated experiments set up across a single continent may fail to capture the full extent of global variation in climate and soil parameter space. IDE with representation on all continents has the potential to address global scale hypotheses about ecosystem sensitivity to environmental change. Our results provide a unique vision of climate and soil variability at the global scale and highlight the need to design globally distributed networks.

Species dominance and biodiversity in plant communities have received considerable attention and characterisation. However, species codominance, while often alleged, is seldom defined or quantified. Codominance is a common phenomenon and is likely to be an important driver of community structure, ecosystem function and the stability of both. Here we review the use of the term 'codominance' and find inconsistencies in its use, suggesting that the scientific community currently lacks a universal understanding of codominance. We address this issue by: (1) qualitatively defining codominance as mostly shared abundance that is distinctively isolated within a subset of a community, and (2) presenting a novel metric for quantifying the degree to which relative abundances are shared among a codominant subset of plant species, while also accounting for the remaining species within a plant community. Using both simulated and real-world data, we then demonstrate the process of applying the codominance metric to compare communities and to generate a quantitatively defensible subset of species to consider codominant within a community. We show that our metric effectively distinguishes the degree of codominance between four types of grassland ecosystems as well as simulated ecosystems with varying degrees of abundance sharing among community members. Overall, we make the case that increased research focusses on the conditions under which codominance occurs and the consequences for species coexistence, community structure and ecosystem function that would considerably advance the fields of community and ecosystem ecology.

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.

Drought, defined as a marked deficiency of precipitation relative to normal, occurs as periods of below-average precipitation or complete failure of precipitation inputs, and can be limited to a single season or prolonged over multiple years. Grasslands are typically quite sensitive to drought, but there can be substantial variability in the magnitude of loss of ecosystem function. We hypothesized that differences in how drought occurs may contribute to this variability. In four native Great Plains grasslands (three C4- and one C3-dominated) spanning a ~ 500-mm precipitation gradient, we imposed drought for four consecutive years by (1) reducing each rainfall event by 66% during the growing season (chronic drought) or (2) completely excluding rainfall during a shorter portion of the growing season (intense drought). The drought treatments were similar in magnitude but differed in the following characteristics: event number, event size and length of dry periods. We observed consistent drought-induced reductions (28–37%) in aboveground net primary production (ANPP) only in the C4-dominated grasslands. In general, intense drought reduced ANPP more than chronic drought, with little evidence that drought duration altered this pattern. Conversely, belowground net primary production (BNPP) was reduced by drought in all grasslands (32–64%), with BNPP reductions greater in intense vs. chronic drought treatments in the most mesic grassland. We conclude that grassland productivity responses to drought did not strongly differ between these two types of drought, but when differences existed, intense drought consistently reduced function more than chronic drought.

Plant‐associated fungi can ameliorate abiotic stress in their hosts, and changes in these fungal communities can alter plant productivity, species interactions, community structure and ecosystem processes. We investigated the response of root‐associated fungi to experimental drought (66% reduction in growing season precipitation) across six North American grassland ecosystem types to determine how extreme drought alters root‐associated fungi, and understand what abiotic factors influence root fungal community composition across grassland ecosystems. Next generation sequencing of the fungal ITS2 region demonstrated that drought primarily re‐ordered fungal species' relative abundances within host plant species, with different fungal responses depending on host identity. Grass species that declined more under drought trended toward less community re‐ordering of root fungi than species less sensitive to drought. Host identity and grassland ecosystem type defined the magnitude of drought effects on community composition, diversity and root colonization, and the most important factor affecting fungal composition was plant species identity. Fungal diversity was less responsive to drought than fungal composition. Across ecosystems, latitude and soil pH were better predictors of fungal diversity than experimental drought. Drought significantly reduced fungal diversity and evenness only in sideoats grama Bouteloua curtipendula and reduced root colonization only in blue grama B. gracilis. Our results illustrate that predicting the effects of drought on root‐associated fungi will require attention to host plant identity and ecosystem type. By comparing our findings against soil microbe responses in the same experiment, we propose the hypothesis that fungi in plant roots are less sensitive to drought than fungi inhabiting soils. Our study highlights the importance of studying community‐level re‐ordering of fungal composition for understanding plant responses to climate change.

Understanding how global change drivers (GCDs) affect aboveground net primary production (ANPP) through time is essential to predicting the reliability and maintenance of ecosystem function and services in the future. While GCDs, such as drought, warming and elevated nutrients, are known to affect mean ANPP, less is known about how they affect inter-annual variability in ANPP. We examined 27 global change experiments located in 11 different herbaceous ecosystems that varied in both abiotic and biotic conditions, to investigate changes in the mean and temporal variability of ANPP (measured as the coefficient of variation) in response to different GCD manipulations, including resource additions, warming, and irrigation. From this comprehensive data synthesis, we found that GCD treatments increased mean ANPP. However, GCD manipulations both increased and decreased temporal variability of ANPP (24% of comparisons), with no net effect overall. These inconsistent effects on temporal variation in ANPP can, in part, be attributed to site characteristics, such as mean annual precipitation and temperature as well as plant community evenness. For example, decreases in temporal variability in ANPP with the GCD treatments occurred in wetter and warmer sites with lower plant community evenness. Further, the addition of several nutrients simultaneously increased the sensitivity of ANPP to interannual variation in precipitation. Based on this analysis, we expect that GCDs will likely affect the magnitude more than the reliability over time of ecosystem production in the future.

© 2020 The Authors., Fire is a powerful ecological and evolutionary force that regulates organismal traits, population sizes, species interactions, community composition, carbon and nutrient cycling and ecosystem function. It also presents a rapidly growing societal challenge, due to both increasingly destructive wildfires and fire exclusion in fire‐dependent ecosystems. As an ecological process, fire integrates complex feedbacks among biological, social and geophysical processes, requiring coordination across several fields and scales of study. Here, we describe the diversity of ways in which fire operates as a fundamental ecological and evolutionary process on Earth. We explore research priorities in six categories of fire ecology: (a) characteristics of fire regimes, (b) changing fire regimes, (c) fire effects on above‐ground ecology, (d) fire effects on below‐ground ecology, (e) fire behaviour and (f) fire ecology modelling. We identify three emergent themes: the need to study fire across temporal scales, to assess the mechanisms underlying a variety of ecological feedbacks involving fire and to improve representation of fire in a range of modelling contexts. Synthesis: As fire regimes and our relationships with fire continue to change, prioritizing these research areas will facilitate understanding of the ecological causes and consequences of future fires and rethinking fire management alternatives., Support was provided by NSF‐DEB‐1743681 to K.K.M. and A.J.T. We thank Shalin Hai‐Jew for helpful discussion of the survey and qualitative methods.

Global change forecasts in ecosystems require knowledge of within-species diversity, particularly of dominant species within communities. We assessed site-level diversity and capacity for adaptation in Bouteloua gracilis, the dominant species in the Central US shortgrass steppe biome. We quantified genetic diversity from 17 sites across regional scales, north to south from New Mexico to South Dakota, and local scales in northern Colorado. We also quantified phenotype and plasticity within and among sites and determined the extent to which phenotypic diversity in B. gracilis was correlated with climate. Genome sequencing indicated pronounced population structure at the regional scale, and local differences indicated that gene flow and/or dispersal may also be limited. Within a common environment, we found evidence of genetic divergence in biomass-related phenotypes, plasticity, and phenotypic variance, indicating functional divergence and different adaptive potential. Phenotypes were differentiated according to climate, chiefly median Palmer Hydrological Drought Index and other aridity metrics. Our results indicate conclusive differences in genetic variation, phenotype, and plasticity in this species and suggest a mechanism explaining variation in shortgrass steppe community responses to global change. This analysis of B. gracilis intraspecific diversity across spatial scales will improve conservation and management of the shortgrass steppe ecosystem in the future.

Plants are subject to trade-offs among growth strategies such that adaptations for optimal growth in one condition can preclude optimal growth in another. Thus, we predicted that a plant species that responds positively to one global change treatment would be less likely than average to respond positively to another treatment, particularly for pairs of treatments that favor distinct traits. We examined plant species’ abundances in 39 global change experiments manipulating two or more of the following: CO2, nitrogen, phosphorus, water, temperature, or disturbance. Overall, the directional response of a species to one treatment was 13% more likely than expected to oppose its response to a another single-factor treatment. This tendency was detectable across the global data set, but held little predictive power for individual treatment combinations or within individual experiments. Although trade-offs in the ability to respond to different global change treatments exert discernible global effects, other forces obscure their influence in local communities.

Increased nitrogen (N) deposition is known to reduce the ecosystem stability, while the underlying mechanisms are still controversial. We conducted an 8-year multi-level N addition experiment in a temperate semi-arid grassland to identify the mechanisms (biodiversity, species asynchrony, population stability and dominant species stability) driving the N-induced loss of temporal stability of aboveground net primary productivity (ANPP). We found that N addition decreased ecosystem, population, and dominant species stability; decreased species richness and phylogenetic diversity; increased species dominance; but had nonsignificant effects on community-wide species asynchrony. Structural equation model revealed that N-induced loss of ecosystem stability was mainly driven by the loss of dominant species stability and the reduction in population stability. Moreover, species relative instability was negatively related with species relative production and the slopes increase with N addition, indicating that N addition weakened the stabilizing effect of dominant species on ecosystem function. Overall, our results highlight that the dominant species control the temporal stability of ANPP in grassland ecosystem under N addition, and support 'dominance management' as an effective strategy for conserving ecosystem functioning in grassland under N deposition.

The intensification of drought throughout the U.S. Great Plains has the potential to have large impacts on grassland functioning, as has been shown with dramatic losses of plant productivity annually. Yet, we have a poor understanding of how grassland functioning responds after drought ends. This study examined how belowground nutrient cycling responds after drought and whether legacy effects persist postdrought. We assessed the 2-year recovery of nutrient cycling processes following a 4-year experimental drought in a mesic grassland by comparing two different growing season drought treatments—chronic (each rainfall event reduced by 66%) and intense (all rain eliminated until 45% of annual rainfall was achieved)—to the control (ambient precipitation) treatment. At the beginning of the first growing season postdrought, we found that in situ soil CO2 efflux and laboratory-based soil microbial respiration were reduced by 42% and 22%, respectively, in the intense drought treatment compared to the control, but both measures had recovered by midseason (July) and remained similar to the control treatment in the second postdrought year. We also found that extractable soil ammonium and total inorganic N were elevated throughout the growing season in the first year after drought in the intense treatment. However, these differences in inorganic N pools did not persist during the growing season of the second year postdrought. The remaining measures of C and N cycling in both drought treatments showed no postdrought treatment effects. Thus, although we observed short-term legacy effects following the intense drought, C and N cycling returned to levels comparable to nondroughted grassland within a single growing season regardless of whether the drought was intense or chronic in nature. Overall, these results suggest that the key aspects of C and N cycling in mesic tallgrass prairie do not exhibit persistent legacies from 4 years of experimentally induced drought.

Plant nonstructural carbohydrates (NSC) can reflect community and ecosystem responses to environmental changes such as water availability. Climate change is predicted to increase aridity and the frequency of extreme drought events in grasslands, but it is unclear how community-scale NSC will respond to drought or how such responses may vary along aridity gradients. We experimentally imposed a 4-year drought in six grasslands along a natural aridity gradient and measured the community-weighted mean of leaf soluble sugar (SS

Climate change is predicted to increase the frequency and intensity of extreme events including droughts and large precipitation events or "deluges." While many studies have focused on the ecological impacts of individual events (e.g., a heat wave), there is growing recognition that when extreme events co-occur as compound extremes, (e.g., a heatwave during a drought), the additive effects on ecosystems are often greater than either extreme alone. In this study, we assessed a unique type of extreme-a contrasting compound extreme-where the extremes may have offsetting, rather than additive ecological effects, by examining how a deluge during a drought impacts productivity and carbon cycling in a semi-arid grassland. The experiment consisted of four treatments: a control (average precipitation), an extreme drought (5th percentile), an extreme drought interrupted by a single deluge (95th percentile), or an extreme drought interrupted by the equivalent amount of precipitation added in several smaller events. We highlight three key results. First, extreme drought resulted in early senescence, reduced carbon uptake, and a decline in net primary productivity relative to the control treatment. Second, the deluge imposed during extreme drought stimulated carbon fluxes and plant growth well above the levels of both the control and the drought treatment with several additional smaller rainfall events, emphasizing the importance of precipitation amount, event size, and timing. Third, while the deluge's positive effects on carbon fluxes and plant growth persisted for 1 month, the deluge did not completely offset the negative effects of extreme drought on end-of-season productivity. Thus, in the case of these contrasting hydroclimatic extremes, a deluge during a drought can stimulate temporally dynamic ecosystem processes (e.g., net ecosystem exchange) while only partially compensating for reductions in ecosystem functions over longer time scales (e.g., aboveground net primary productivity).

It remains unclear how competitive exclusion is avoided between two ecologically, economically, and culturally important codominant grass species in the tallgrass prairie of the Great Plains, Andropogon gerardii and Sorghastrum nutans. These functionally similar C4 grasses appear to coexist despite considerable niche overlap, and asymmetric competition and drought tolerance in favor of A. gerardii. According to the stress gradient hypothesis, it may be that the sum of interactions between these species, which is typically negative (competitive) due to similar resource requirements, shifts to positive (facilitative) as abiotic stress increases. For instance, if the canopy cover of the stronger competitor reduces losses of subcanopy humidity or shallow soil moisture, recruitment of S. nutans tillers may be extended further into the drought event than would occur in the absence of A. gerardii. As later months of the growing season are drier on average where these species are codominant, such a mechanism may enable S. nutans to recover from early season asymmetric competition and stabilize their codominance. We tested this hypothesis in a greenhouse experiment in which we manipulated community composition and water availability in the latter half of the growing season. We found no evidence that a shift from a negative to positive interaction occurs, with each species performing similarly in mixed communities and monocultures. The similarities of the two species in their functional traits and responses to water limitations may limit such a shift in interaction net effects and suggests that other mechanisms are determining coexistence of these co-occuring C4 grasses.

Globally, extreme droughts are likely to become more frequent, more intense, and longer in duration with climate change. Understanding the impacts of drought on ecosystem function (i.e., resistance) and its recovery after drought (i.e., resilience) is critical for long-term sustainability of ecosystem services under climate change. We tested the effects of drought duration on ecosystem resistance by imposing four years of extreme drought (

Ongoing intensification of the hydrological cycle is altering rainfall regimes by increasing the frequency of extreme wet and dry years and the size of individual rainfall events. Despite long-standing recognition of the importance of precipitation amount and variability for most terrestrial ecosystem processes, we lack understanding of their interactive effects on ecosystem functioning. We quantified this interaction in native grassland by experimentally eliminating temporal variability in growing season rainfall over a wide range of precipitation amounts, from extreme wet to dry conditions. We contrasted the rain use efficiency (RUE) of above-ground net primary productivity (ANPP) under conditions of experimentally reduced versus naturally high rainfall variability using a 32-year precipitation-ANPP dataset from the same site as our experiment. We found that increased growing season rainfall variability can reduce RUE and thus ecosystem functioning by as much as 42% during dry years, but that such impacts weaken as years become wetter. During low precipitation years, RUE is lowest when rainfall event sizes are relatively large, and when a larger proportion of total rainfall is derived from large events. Thus, a shift towards precipitation regimes dominated by fewer but larger rainfall events, already documented over much of the globe, can be expected to reduce the functioning of mesic ecosystems primarily during drought, when ecosystem processes are already compromised by low water availability.

Plant traits can provide unique insights into plant performance at the community scale. Functional composition, defined by both functional diversity and community‐weighted trait means (CWMs), can affect the stability of above‐ground net primary production (ANPP) in response to climate extremes. Further complexity arises, however, when functional composition itself responds to environmental change. The duration of climate extremes, such as drought, is expected to increase with rising global temperatures; thus, understanding the impacts of long‐term drought on functional composition and the corresponding effect that has on ecosystem function could improve predictions of ecosystem sensitivity to climate change. We experimentally reduced growing season precipitation by 66% across six temperate grasslands for 4 years and measured changes in three indices of functional diversity (functional dispersion, richness and evenness), community‐weighted trait means and phylogenetic diversity (PD). Specific leaf area (SLA), leaf nitrogen content (LNC) and (at most sites) leaf turgor loss point (πTLP) were measured for species cumulatively representing ~90% plant cover at each site. Long‐term drought led to increased community functional dispersion in three sites, with negligible effects on the remaining sites. Species re‐ordering following the mortality/senescence of dominant species was the main driver of increased functional dispersion. The response of functional diversity was not consistently matched by changes in phylogenetic diversity. Community‐level drought strategies (assessed as CWMs) largely shifted from drought tolerance to drought avoidance and/or escape strategies, as evidenced by higher community‐weighted πTLP, SLA and LNC. Lastly, ecosystem drought sensitivity (i.e. relative reduction in ANPP in drought plots) was positively correlated with community‐weighted SLA and negatively correlated with functional diversity. Synthesis. Increased functional diversity following long‐term drought may stabilize ecosystem functioning in response to future drought. However, shifts in community‐scale drought strategies may increase ecosystem drought sensitivity, depending on the nature and timing of drought. Thus, our results highlight the importance of considering both functional diversity and abundance‐weighted traits means of plant communities as their collective effect may either stabilize or enhance ecosystem sensitivity to drought.

Direct quantification of terrestrial biosphere responses to global change is crucial for projections of future climate change in Earth system models. Here, we synthesized ecosystem carbon-cycling data from 1,119 experiments performed over the past four decades concerning changes in temperature, precipitation, CO2 and nitrogen across major terrestrial vegetation types of the world. Most experiments manipulated single rather than multiple global change drivers in temperate ecosystems of the USA, Europe and China. The magnitudes of warming and elevated CO2 treatments were consistent with the ranges of future projections, whereas those of precipitation changes and nitrogen inputs often exceeded the projected ranges. Increases in global change drivers consistently accelerated, but decreased precipitation slowed down carbon-cycle processes. Nonlinear (including synergistic and antagonistic) effects among global change drivers were rare. Belowground carbon allocation responded negatively to increased precipitation and nitrogen addition and positively to decreased precipitation and elevated CO2. The sensitivities of carbon variables to multiple global change drivers depended on the background climate and ecosystem condition, suggesting that Earth system models should be evaluated using site-specific conditions for best uses of this large dataset. Together, this synthesis underscores an urgent need to explore the interactions among multiple global change drivers in under-represented regions such as semi-arid ecosystems, forests in the tropics and subtropics, and Arctic tundra when forecasting future terrestrial carbon-climate feedback. National Natural Science Foundation of ChinaNational Natural Science Foundation of China [31430015, 31830012]; US NSFNational Science Foundation (NSF) [DEB-0955771]; ClimMani COST actionEuropean Cooperation in Science and Technology (COST) [ES1308] We thank J. Wang (Hebei University), S. Yang (Institute of Botany, Chinese Academy of Sciences), L. Zhou (East China Normal University), C. Qiao (Xinyang Normal University) and H. Li (Henan University) for their help in meta-analyses and interaction analyses, and H. Li, Y. Liu (Institute of Tibetan Plateau Research, Chinese Academy of Sciences) and Y. He (Peking University) for their help in plotting figures. This work was financially supported by the National Natural Science Foundation of China (grant nos. 31430015 and 31830012). This study emerged from the INTERFACE Workshop in Beijing, China (https://www.bio.purdue.edu/INTERFACE/) supported by the US NSF DEB-0955771. We also acknowledge support from the ClimMani COST action (ES1308). Public domain – authored by a U.S. government employee

Drought, widely studied as an important driver of ecosystem dynamics, is predicted to increase in frequency and severity globally. To study drought, ecologists must define or at least operationalize what constitutes a drought. How this is accomplished in practice is unclear, particularly given that climatologists have long struggled to agree on definitions of drought, beyond general variants of "an abnormal deficiency of water." We conducted a literature review of ecological drought studies (564 papers) to assess how ecologists describe and study drought. We found that ecologists characterize drought in a wide variety of ways (reduced precipitation, low soil moisture, reduced streamflow, etc.), but relatively few publications (~32%) explicitly define what are, and are not, drought conditions. More troubling, a surprising number of papers (~30%) simply equated "dry conditions" with "drought" and provided little characterization of the drought conditions studied. For a subset of these, we calculated Standardized Precipitation Evapotranspiration Index values for the reported drought periods. We found that while almost 90% of the studies were conducted under conditions quantifiable as slightly to extremely drier than average, ~50% were within the range of normal climatic variability. We conclude that the current state of the ecological drought literature hinders synthesis and our ability to draw broad ecological inferences because drought is often declared but is not explicitly defined or well characterized. We suggest that future drought publications provide at least one of the following: (a) the climatic context of the drought period based on long-term records; (b) standardized climatic index values; (c) published metrics from drought-monitoring organizations; (d) a quantitative definition of what the authors consider to be drought conditions for their system. With more detailed and consistent quantification of drought conditions, comparisons among studies can be more rigorous, increasing our understanding of the ecological effects of drought.

Desertification of sandy grasslands is an increasing problem, with serious negative impacts on ecosystem functions. Sandy grasslands are fragile with low ecosystem productivity mainly because of the sandy soil structure with low water and nutrient holding capacities and especially low levels of nitrogen. Here, we evaluate the long‐term impacts of sediment addition from a local reservoir, and grass and legume cultivation (artificial grasslands) on a sandy grassland in eastern Inner Mongolia, China. The results showed that even after 32 years, sediment addition had improved soil structure significantly, that is, increasing of silt and clay contents, soil bulk density, and water holding capacity. As the result of improved soil structure, ecosystem functions, including aboveground net primary productivity (ANPP) and soil carbon, nitrogen (N), and phosphorus storage, increased significantly. Net C, N, and P sequestration increased even after accounting for the sediment addition, due, at least partially, to the greater plant biomass trapping large quantities of wind‐blown dust. Plant cultivation, especially the addition of a legume, further increased ANPP significantly, that is, the cultivation of Leymus chinensis and the legume Medicago sativa increased ANPP 6.99 and 44.62 times, respectively. Our study highlights that improvements in soil structure and cultivation with legume species can increased substantially the productivity of sandy grasslands and that the initial increases in grass biomass promoted the sequestration of wind‐blown dust, which helped sustain the increases in productivity.

Nitrogen (N) enrichment often increases aboveground net primary productivity (ANPP) of the ecosystem, but it is unclear if belowground net primary productivity (BNPP) track responses of ANPP. Moreover, the frequency of N inputs may affect primary productivity but is rarely studied. To assess the response patterns of above- and belowground productivity to rates of N addition under different addition frequencies, we manipulated the rate (0-50 g N m-2 year-1 ) and frequency (twice vs. monthly additions per year) of NH4 NO3 inputs for six consecutive years in a temperate grassland in northern China and measured ANPP and BNPP from 2012 to 2014. In the low range of N addition rates, BNPP showed the greatest negative response and ANPP showed the greatest positive responses with increases in N addition (

The pattern of a few abundant species and many rarer species is a defining characteristic of communities worldwide. These abundant species are often referred to as dominant species. Yet, despite their importance, the term dominant species is poorly defined and often used to convey different information by different authors. Based on a review of historical and contemporary definitions we develop a synthetic definition of dominant species. This definition incorporates the relative local abundance of a species, its ubiquity across the landscape, and its impact on community and ecosystem properties. A meta-analysis of removal studies shows that the loss of species identified as dominant by authors can significantly impact ecosystem functioning and community structure. We recommend two metrics that can be used jointly to identify dominant species in a given community and provide a roadmap for future avenues of research on dominant species. In our review, we make the case that the identity and effects of dominant species on their environments are key to linking patterns of diversity to ecosystem function, including predicting impacts of species loss and other aspects of global change on ecosystems.