Extreme drought impacts have been underestimated in grasslands and shrublands globally.

Climate change is increasing the frequency and severity of short-term (~1 y) drought events-the most common duration of drought-globally. Yet the impact of this intensification of drought on ecosystem functioning remains poorly resolved. This is due in part to the widely disparate approaches ecologists have employed to study drought, variation in the severity and duration of drought studied, and differences among ecosystems in vegetation, edaphic and climatic attributes that can mediate drought impacts. To overcome these problems and better identify the factors that modulate drought responses, we used a coordinated distributed experiment to quantify the impact of short-term drought on grassland and shrubland ecosystems. With a standardized approach, we imposed ~a single year of drought at 100 sites on six continents. Here we show that loss of a foundational ecosystem function-aboveground net primary production (ANPP)-was 60% greater at sites that experienced statistically extreme drought (1-in-100-y event) vs. those sites where drought was nominal (historically more common) in magnitude (35% vs. 21%, respectively). This reduction in a key carbon cycle process with a single year of extreme drought greatly exceeds previously reported losses for grasslands and shrublands. Our global experiment also revealed high variability in drought response but that relative reductions in ANPP were greater in drier ecosystems and those with fewer plant species. Overall, our results demonstrate with unprecedented rigor that the global impacts of projected increases in drought severity have been significantly underestimated and that drier and less diverse sites are likely to be most vulnerable to extreme drought.

Reintroducing bison results in long-running and resilient increases in grassland diversity.

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 m2) 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.

Grazing herbivores reduce herbaceous biomass and fire activity across African savannas.

Fire and herbivory interact to alter ecosystems and carbon cycling. In savannas, herbivores can reduce fire activity by removing grass biomass, but the size of these effects and what regulates them remain uncertain. To examine grazing effects on fuels and fire regimes across African savannas, we combined data from herbivore exclosure experiments with remotely sensed data on fire activity and herbivore density. We show that, broadly across African savannas, grazing herbivores substantially reduce both herbaceous biomass and fire activity. The size of these effects was strongly associated with grazing herbivore densities, and surprisingly, was mostly consistent across different environments. A one-zebra increase in herbivore biomass density (~100 kg/km2 of metabolic biomass) resulted in a ~53 kg/ha reduction in standing herbaceous biomass and a ~0.43 percentage point reduction in burned area. Our results indicate that fire models can be improved by incorporating grazing effects on grass biomass.

Accounting for herbaceous communities in process-based models will advance our understanding of "grassy" ecosystems.

Grassland and other herbaceous communities cover significant portions of Earth's terrestrial surface and provide many critical services, such as carbon sequestration, wildlife habitat, and food production. Forecasts of global change impacts on these services will require predictive tools, such as process-based dynamic vegetation models. Yet, model representation of herbaceous communities and ecosystems lags substantially behind that of tree communities and forests. The limited representation of herbaceous communities within models arises from two important knowledge gaps: first, our empirical understanding of the principles governing herbaceous vegetation dynamics is either incomplete or does not provide mechanistic information necessary to drive herbaceous community processes with models; second, current model structure and parameterization of grass and other herbaceous plant functional types limits the ability of models to predict outcomes of competition and growth for herbaceous vegetation. In this review, we provide direction for addressing these gaps by: (1) presenting a brief history of how vegetation dynamics have been developed and incorporated into earth system models, (2) reporting on a model simulation activity to evaluate current model capability to represent herbaceous vegetation dynamics and ecosystem function, and (3) detailing several ecological properties and phenomena that should be a focus for both empiricists and modelers to improve representation of herbaceous vegetation in models. Together, empiricists and modelers can improve representation of herbaceous ecosystem processes within models. In so doing, we will greatly enhance our ability to forecast future states of the earth system, which is of high importance given the rapid rate of environmental change on our planet.

Traits that distinguish dominant species across aridity gradients differ from those that respond to soil moisture.

Many plant traits respond to changes in water availability and might be useful for understanding ecosystem properties such as net primary production (NPP). This is especially evident in grasslands where NPP is water-limited and primarily determined by the traits of dominant species. We measured root and shoot morphology, leaf hydraulic traits, and NPP of four dominant North American prairie grasses in response to four levels of soil moisture in a greenhouse experiment. We expected that traits of species from drier regions would be more responsive to reduced water availability and that this would make these species more resistant to low soil moisture than species from wetter regions. All four species grew taller, produced more biomass, and increased total root length in wetter treatments. Each species reduced its leaf turgor loss point (TLP) in drier conditions, but only two species (one xeric, one mesic) maintained leaf water potential above TLP. We identified a suite of traits that clearly distinguished species from one another, but, surprisingly, these traits were relatively unresponsive to reduced soil moisture. Specifically, more xeric species produced thinner roots with higher specific root length and had a lower root mass fraction. This suggest that root traits are critical for distinguishing species from one another but might not respond strongly to changing water availability, though this warrants further investigation in the field. Overall, we found that NPP of these dominant grass species responded similarly to varying levels of soil moisture despite differences in species morphology, physiology, and habitat of origin.

Multiple global change drivers show independent, not interactive effects: a long-term case study in tallgrass prairie.

Ecosystems are faced with an onslaught of co-occurring global change drivers. While frequently studied independently, the effects of multiple global change drivers have the potential to be additive, antagonistic, or synergistic. Global warming, for example, may intensify the effects of more variable precipitation regimes with warmer temperatures increasing evapotranspiration and thereby amplifying the effect of already dry soils. Here, we present the long-term effects (11 years) of altered precipitation patterns (increased intra-annual variability in the growing season) and warming (1 °C year-round) on plant community composition and aboveground net primary productivity (ANPP), a key measure of ecosystem functioning in mesic tallgrass prairie. Based on past results, we expected that increased precipitation variability and warming would have additive effects on both community composition and ANPP. Increased precipitation variability altered plant community composition and increased richness, with no effect on ANPP. In contrast, warming decreased ANPP via reduction in grass stems and biomass but had no effect on the plant community. Contrary to expectations, across all measured variables, precipitation and warming treatments had no interactive effects. While treatment interactions did not occur, each treatment did individually impact a different component of the ecosystem (i.e., community vs. function). Thus, different aspects of the ecosystem may be sensitive to different global change drivers in mesic grassland ecosystems.

Resolving the Dust Bowl paradox of grassland responses to extreme drought.

During the 1930s Dust Bowl drought in the central United States, species with the C3 photosynthetic pathway expanded throughout C4-dominated grasslands. This widespread increase in C3 grasses during a decade of low rainfall and high temperatures is inconsistent with well-known traits of C3 vs. C4 pathways. Indeed, water use efficiency is generally lower, and photosynthesis is more sensitive to high temperatures in C3 than C4 species, consistent with the predominant distribution of C3 grasslands in cooler environments and at higher latitudes globally. We experimentally imposed extreme drought for 4 y in mixed C3/C4 grasslands in Kansas and Wyoming and, similar to Dust Bowl observations, also documented three- to fivefold increases in C3/C4 biomass ratios. To explain these paradoxical responses, we first analyzed long-term climate records to show that under nominal conditions in the central United States, C4 grasses dominate where precipitation and air temperature are strongly related (warmest months are wettest months). In contrast, C3 grasses 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 C3 grasses 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 C3 and C4 species.

What happens after drought ends: synthesizing terms and definitions.

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 legacies determine grassland responses to future rainfall regimes.

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 for >25 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 CO2 efflux are contingent on current conditions. Although total soil C remained similar across treatments, long-term irrigation increased labile soil C and the sensitivity of microbial biomass C to drought. Importantly, the magnitude of legacy effects for all response variables varied with topography, suggesting that landscape can modulate the strength and direction of climate legacies. Our results demonstrate the role of climate history as an important determinant of terrestrial C cycling responses to future climate changes.

Compound hydroclimatic extremes in a semi-arid grassland: Drought, deluge, and the carbon cycle.

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).

Richness, not evenness, varies across water availability gradients in grassy biomes on five continents.

We sought to understand the role that water availability (expressed as an aridity index) plays in determining regional and global patterns of richness and evenness, and in turn how these water availability-diversity relationships may result in different richness-evenness relationships at regional and global scales. We examined relationships between water availability, richness and evenness for eight grassy biomes spanning broad water availability gradients on five continents. Our study found that relationships between richness and water availability switched from positive for drier (South Africa, Tibet and USA) vs. negative for wetter (India) biomes, though were not significant for the remaining biomes. In contrast, only the India biome showed a significant relationship between water availability and evenness, which was negative. Globally, the richness-water availability relationship was hump-shaped, however, not significant for evenness. At the regional scale, a positive richness-evenness relationship was found for grassy biomes in India and Inner Mongolia, China. In contrast, this relationship was weakly concave-up globally. These results suggest that different, independent factors are determining patterns of species richness and evenness in grassy biomes, resulting in differing richness-evenness relationships at regional and global scales. As a consequence, richness and evenness may respond very differently across spatial gradients to anthropogenic changes, such as climate change.

Integrative modelling reveals mechanisms linking productivity and plant species richness.

How ecosystem productivity and species richness are interrelated is one of the most debated subjects in the history of ecology. Decades of intensive study have yet to discern the actual mechanisms behind observed global patterns. Here, by integrating the predictions from multiple theories into a single model and using data from 1,126 grassland plots spanning five continents, we detect the clear signals of numerous underlying mechanisms linking productivity and richness. We find that an integrative model has substantially higher explanatory power than traditional bivariate analyses. In addition, the specific results unveil several surprising findings that conflict with classical models. These include the isolation of a strong and consistent enhancement of productivity by richness, an effect in striking contrast with superficial data patterns. Also revealed is a consistent importance of competition across the full range of productivity values, in direct conflict with some (but not all) proposed models. The promotion of local richness by macroecological gradients in climatic favourability, generally seen as a competing hypothesis, is also found to be important in our analysis. The results demonstrate that an integrative modelling approach leads to a major advance in our ability to discern the underlying processes operating in ecological systems.

Determinants of community compositional change are equally affected by global change.

Global change is impacting plant community composition, but the mechanisms underlying these changes are unclear. Using a dataset of 58 global change experiments, we tested the five fundamental mechanisms of community change: changes in evenness and richness, reordering, species gains and losses. We found 71% of communities were impacted by global change treatments, and 88% of communities that were exposed to two or more global change drivers were impacted. Further, all mechanisms of change were equally likely to be affected by global change treatments-species losses and changes in richness were just as common as species gains and reordering. We also found no evidence of a progression of community changes, for example, reordering and changes in evenness did not precede species gains and losses. We demonstrate that all processes underlying plant community composition changes are equally affected by treatments and often occur simultaneously, necessitating a wholistic approach to quantifying community changes.

Defining codominance in plant communities.

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.

Precipitation-productivity relationships and the duration of precipitation anomalies: An underappreciated dimension of climate change.

In terrestrial ecosystems, climate change forecasts of increased frequencies and magnitudes of wet and dry precipitation anomalies are expected to shift precipitation-net primary productivity (PPT-NPP) relationships from linear to nonlinear. Less understood, however, is how future changes in the duration of PPT anomalies will alter PPT-NPP relationships. A review of the literature shows strong potential for the duration of wet and dry PPT anomalies to impact NPP and to interact with the magnitude of anomalies. Within semi-arid and mesic grassland ecosystems, PPT gradient experiments indicate that short-duration (1 year) PPT anomalies are often insufficient to drive nonlinear aboveground NPP responses. But long-term studies, within desert to forest ecosystems, demonstrate how multi-year PPT anomalies may result in increasing impacts on NPP through time, and thus alter PPT-NPP relationships. We present a conceptual model detailing how NPP responses to PPT anomalies may amplify with the duration of an event, how responses may vary in xeric vs. mesic ecosystems, and how these differences are most likely due to demographic mechanisms. Experiments that can unravel the independent and interactive impacts of the magnitude and duration of wet and dry PPT anomalies are needed, with multi-site long-term PPT gradient experiments particularly well-suited for this task.

Is a drought a drought in grasslands? Productivity responses to different types of 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.

Biodiversity increases the resistance of ecosystem productivity to climate extremes.

It remains unclear whether biodiversity buffers ecosystems against climate extremes, which are becoming increasingly frequent worldwide. Early results suggested that the ecosystem productivity of diverse grassland plant communities was more resistant, changing less during drought, and more resilient, recovering more quickly after drought, than that of depauperate communities. However, subsequent experimental tests produced mixed results. Here we use data from 46 experiments that manipulated grassland plant diversity to test whether biodiversity provides resistance during and resilience after climate events. We show that biodiversity increased ecosystem resistance for a broad range of climate events, including wet or dry, moderate or extreme, and brief or prolonged events. Across all studies and climate events, the productivity of low-diversity communities with one or two species changed by approximately 50% during climate events, whereas that of high-diversity communities with 16-32 species was more resistant, changing by only approximately 25%. By a year after each climate event, ecosystem productivity had often fully recovered, or overshot, normal levels of productivity in both high- and low-diversity communities, leading to no detectable dependence of ecosystem resilience on biodiversity. Our results suggest that biodiversity mainly stabilizes ecosystem productivity, and productivity-dependent ecosystem services, by increasing resistance to climate events. Anthropogenic environmental changes that drive biodiversity loss thus seem likely to decrease ecosystem stability, and restoration of biodiversity to increase it, mainly by changing the resistance of ecosystem productivity to climate events.

Genetic and functional variation across regional and local scales is associated with climate in a foundational prairie grass.

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.

Lineage-based functional types: characterising functional diversity to enhance the representation of ecological behaviour in Land Surface Models.

Process-based vegetation models attempt to represent the wide range of trait variation in biomes by grouping ecologically similar species into plant functional types (PFTs). This approach has been successful in representing many aspects of plant physiology and biophysics but struggles to capture biogeographic history and ecological dynamics that determine biome boundaries and plant distributions. Grass-dominated ecosystems are broadly distributed across all vegetated continents and harbour large functional diversity, yet most Land Surface Models (LSMs) summarise grasses into two generic PFTs based primarily on differences between temperate C3 grasses and (sub)tropical C4 grasses. Incorporation of species-level trait variation is an active area of research to enhance the ecological realism of PFTs, which form the basis for vegetation processes and dynamics in LSMs. Using reported measurements, we developed grass functional trait values (physiological, structural, biochemical, anatomical, phenological, and disturbance-related) of dominant lineages to improve LSM representations. Our method is fundamentally different from previous efforts, as it uses phylogenetic relatedness to create lineage-based functional types (LFTs), situated between species-level trait data and PFT-level abstractions, thus providing a realistic representation of functional diversity and opening the door to the development of new vegetation models.

Precipitation amount and event size interact to reduce ecosystem functioning during dry years in a mesic grassland.

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.