Unexpected patterns of sensitivity to drought in three semi-arid grasslands

Global climate models forecast an increase in the frequency and intensity of extreme weather events, including severe droughts. Based on multi-year relationships between precipitation amount and aboveground annual net primary production (ANPP), semi-arid grasslands are projected to be among the most sensitive ecosystems to changes in precipitation. To assess sensitivity to drought, as well as variability within the shortgrass steppe biome, we imposed moderate and severe rainfall reductions for two growing seasons in three undisturbed grasslands that varied in soil type and climate. We predicted strong drought-induced reductions in ANPP at all sites and greater sensitivity to drought in sites with lower average precipitation, consistent with continental-scale patterns. Identical experimental infrastructure at each site reduced growing season rainfall events by 50 or 80%, and significantly reduced average soil moisture in both years (by 21 and 46% of control levels, respectively). Despite reductions in soil moisture, ANPP responses varied unexpectedly-from no reduction in ANPP to a 51% decrease. Although sensitivity to drought was highest in the semi-arid grassland with lowest mean annual precipitation, patterns in responses to drought across these grasslands were also strongly related to rainfall event size. When growing season rainfall patterns were dominated by many smaller events, ANPP was significantly reduced by drought but not when rainfall patterns were characterized by large rain events. This interaction between drought sensitivity and rainfall event size suggests that ANPP responses to future droughts may be reduced if growing season rainfall regimes also become more extreme.     

Linking drought legacy effects across scales: From leaves to tree rings to ecosystems

Severe drought can cause lagged effects on tree physiology that negatively impact forest functioning for years. These “drought legacy effects” have been widely documented in tree‐ring records and could have important implications for our understanding of broader scale forest carbon cycling. However, legacy effects in tree‐ring increments may be decoupled from ecosystem fluxes due to (a) postdrought alterations in carbon allocation patterns; (b) temporal asynchrony between radial growth and carbon uptake; and (c) dendrochronological sampling biases. In order to link legacy effects from tree rings to whole forests, we leveraged a rich dataset from a Midwestern US forest that was severely impacted by a drought in 2012. At this site, we compiled tree‐ring records, leaf‐level gas exchange, eddy flux measurements, dendrometer band data, and satellite remote sensing estimates of greenness and leaf area before, during, and after the 2012 drought. After accounting for the relative abundance of tree species in the stand, we estimate that legacy effects led to ~10% reductions in tree‐ring width increments in the year following the severe drought. Despite this stand‐scale reduction in radial growth, we found that leaf‐level photosynthesis, gross primary productivity (GPP), and vegetation greenness were not suppressed in the year following the 2012 drought. Neither temporal asynchrony between radial growth and carbon uptake nor sampling biases could explain our observations of legacy effects in tree rings but not in GPP. Instead, elevated leaf‐level photosynthesis co‐occurred with reduced leaf area in early 2013, indicating that resources may have been allocated away from radial growth in conjunction with postdrought upregulation of photosynthesis and repair of canopy damage. Collectively, our results indicate that tree‐ring legacy effects were not observed in other canopy processes, and that postdrought canopy allocation could be an important mechanism that decouples tree‐ring signals from GPP.     

Drought timing influences the legacy of tree growth recovery

Whether and how the timing of extreme events affects the direction and magnitude of legacy effects on tree growth is poorly understood. In this study, we use a global database of Ring‐Width Index (RWI) from 2,500 sites to examine the impact and legacy effects (the departure of observed RWI from expected RWI) of extreme drought events during 1948–2008, with a particular focus on the influence of drought timing. We assessed the recovery of stem radial growth in the years following severe drought events with separate groupings designed to characterize the timing of the drought. We found that legacies from extreme droughts during the dry season (DS droughts) lasted longer and had larger impacts in each of the 3 years post drought than those from extreme droughts during the wet season (WS droughts). At the global scale, the average integrated legacy from DS droughts (0.18) was about nine times that from WS droughts (0.02). Site‐level comparisons also suggest stronger negative impacts or weaker positive impacts of DS droughts on tree growth than WS droughts. Our results, therefore, highlight that the timing of drought is a crucial factor determining drought impacts on tree recovery. Further increases in baseline aridity could therefore exacerbate the impact of punctuated droughts on terrestrial ecosystems.     

High land‐use intensity exacerbates shifts in grassland vegetation composition after severe experimental drought

Climate change projections anticipate increased frequency and intensity of drought stress, but grassland responses to severe droughts and their potential to recover are poorly understood. In many grasslands, high land‐use intensity has enhanced productivity and promoted resource‐acquisitive species at the expense of resource‐conservative ones. Such changes in plant functional composition could affect the resistance to drought and the recovery after drought of grassland ecosystems with consequences for feed productivity resilience and environmental stewardship. In a 12‐site precipitation exclusion experiment in upland grassland ecosystems across Switzerland, we imposed severe edaphic drought in plots under rainout shelters and compared them with plots under ambient conditions. We used soil water potentials to scale drought stress across sites. Impacts of precipitation exclusion and drought legacy effects were examined along a gradient of land‐use intensity to determine how grasslands resisted to, and recovered after drought. In the year of precipitation exclusion, aboveground net primary productivity (ANPP) in plots under rainout shelters was ?15% to ?56% lower than in control plots. Drought effects on ANPP increased with drought severity, specified as duration of topsoil water potential ψ < ?100 kPa, irrespective of land‐use intensity. In the year after drought, ANPP had completely recovered, but total species diversity had declined by ?10%. Perennial species showed elevated mortality, but species richness of annuals showed a small increase due to enhanced recruitment. In general, the more resource‐acquisitive grasses increased at the expense of the deeper‐rooted forbs after drought, suggesting that community reorganization was driven by competition rather than plant mortality. The negative effects of precipitation exclusion on forbs increased with land‐use intensity. Our study suggests a synergistic impact of land‐use intensification and climate change on grassland vegetation composition, and implies that biomass recovery after drought may occur at the expense of biodiversity maintenance.     

Dominant tree species are at risk from exaggerated drought under climate change

Predicting the consequences of climate change on forest systems is difficult because trees may display species‐specific responses to exaggerated droughts that may not be reflected by the climatic envelope of their geographic range. Furthermore, few studies have examined the postdrought recovery potential of drought‐susceptible tree species. This study develops a robust ranking of the drought susceptibility of 21 tree species based on their mortality after two droughts (1990s and 2000s) in the savanna of north‐eastern Australia. Drought‐induced mortality was positively related to species dominance, negatively related to the ratio of postdrought seedlings to adults and had no relationship to the magnitude of extreme drought within the species current geographic ranges. These results suggest that predicting the consequences of exaggerated drought on species’ geographic ranges is difficult, but that dominant species like Eucalyptus with relatively slow rates of population recovery and dispersal are the most susceptible. The implications for savanna ecosystems are lower tree densities and basal area.     

Quantifying drylands' drought resistance and recovery: the importance of drought intensity,dominant life history and grazing regime

Projected global change will increase the level of land‐use and environmental stressors such as drought and grazing, particularly in drylands. Still, combined effects of drought and grazing on plant production are poorly understood, thus hampering adequate projections and development of mitigation strategies. We used a large, cross‐continental database consisting of 174 long‐term datasets from >30 dryland regions to quantify ecosystem responses to drought and grazing with the ultimate goal to increase functional understanding in these responses. Two key aspects of ecosystem stability, resistance to and recovery after a drought, were evaluated based on standardized and normalized aboveground net primary production (ANPP) data. Drought intensity was quantified using the standardized precipitation index. We tested effects of drought intensity, grazing regime (grazed, ungrazed), biome (grassland, shrubland, savanna) or dominant life history (annual, perennial) of the herbaceous layer to assess the relative importance of these factors for ecosystem stability, and to identify predictable relationships between drought intensity and ecosystem resistance and recovery. We found that both components of ecosystem stability were better explained by dominant herbaceous life history than by biome. Increasing drought intensity (quasi‐) linearly reduced ecosystem resistance. Even though annual and perennial systems showed the same response rate to increasing drought intensity, they differed in their general magnitude of resistance, with annual systems being ca. 27% less resistant. In contrast, systems with an herbaceous layer dominated by annuals had substantially higher postdrought recovery, particularly when grazed. Combined effects of drought and grazing were not merely additive but modulated by dominant life history of the herbaceous layer. To the best of our knowledge, our study established the first predictive, cross‐continental model between drought intensity and drought‐related relative losses in ANPP, and suggests that systems with an herbaceous layer dominated by annuals are more prone to ecosystem degradation under future global change regimes.     

Contrasting Effects of Precipitation Manipulations on Production in Two Sites within the Central Grassland Region, USA

In grassland ecosystems, where soil water most frequently controls ecosystem processes, expected changes in precipitation and temperature may have dramatic effects on ecosystem dynamics. Previous observational studies have reported that aboveground net primary production (ANPP) in grasslands is very sensitive to changes in precipitation. Yet, we lack experimentally based evidence to support these observations. Further, most of the studies have focused solely on ANPP, neglecting belowground production (BNPP). This is an important gap in our knowledge, as BNPP represents 50% or more of total net primary production (NPP) in grasslands. Here, we present results from a 3-year water manipulation experiment (2008–2010) at two sites in the central grassland region of North America, USA. We were successful in changing the soil water content in our treatments, but these changes resulted in different, but significant responses in ANPP and BNPP at our two sites. At the shortgrass steppe, we found that neither NPP nor ANPP were sensitive to treatment precipitation, and although we found BNPP was sensitive to changes in treatment precipitation, the direction of the response varied between years. In contrast, ANPP was very sensitive to treatment precipitation on the mixed-grass prairie, whereas BNPP was insensitive. Based on our finding that two grassland ecosystems showed dramatically different above and belowground production responses to soil water manipulations, we cannot assume that predicted changes in climate will cause similar above- and belowground production responses. Second, our results demonstrated that sites within the same region may differ markedly in the sensitivity of ANPP to changes in growing season precipitation.     

Asymmetric responses of primary productivity to precipitation extremes: A synthesis of grassland precipitation manipulation experiments

Climatic changes are altering Earth's hydrological cycle, resulting in altered precipitation amounts, increased interannual variability of precipitation, and more frequent extreme precipitation events. These trends will likely continue into the future, having substantial impacts on net primary productivity (NPP) and associated ecosystem services such as food production and carbon sequestration. Frequently, experimental manipulations of precipitation have linked altered precipitation regimes to changes in NPP. Yet, findings have been diverse and substantial uncertainty still surrounds generalities describing patterns of ecosystem sensitivity to altered precipitation. Additionally, we do not know whether previously observed correlations between NPP and precipitation remain accurate when precipitation changes become extreme. We synthesized results from 83 case studies of experimental precipitation manipulations in grasslands worldwide. We used meta‐analytical techniques to search for generalities and asymmetries of aboveground NPP (ANPP) and belowground NPP (BNPP) responses to both the direction and magnitude of precipitation change. Sensitivity (i.e., productivity response standardized by the amount of precipitation change) of BNPP was similar under precipitation additions and reductions, but ANPP was more sensitive to precipitation additions than reductions; this was especially evident in drier ecosystems. Additionally, overall relationships between the magnitude of productivity responses and the magnitude of precipitation change were saturating in form. The saturating form of this relationship was likely driven by ANPP responses to very extreme precipitation increases, although there were limited studies imposing extreme precipitation change, and there was considerable variation among experiments. This highlights the importance of incorporating gradients of manipulations, ranging from extreme drought to extreme precipitation increases into future climate change experiments. Additionally, policy and land management decisions related to global change scenarios should consider how ANPP and BNPP responses may differ, and that ecosystem responses to extreme events might not be predicted from relationships found under moderate environmental changes.     

Drought timing differentially affects above- and belowground productivity in a mesic grassland

Climate models forecast an intensification of the global hydrological cycle with droughts becoming more frequent and severe, and shifting to times when they have been historically uncommon. Droughts, or prolonged periods of precipitation deficiency, are characteristic of most temperate grasslands, yet few experiments have explored how variation in the seasonal timing of drought may impact ecosystem function. We investigated the response of above- and belowground net primary production (ANPP & BNPP) to altered drought timing in a mesic grassland in NE Kansas. Moderate drought treatments (25% reduction from the mean growing season precipitation [GSP]) were imposed by erecting rainout shelters in late spring (LSP), early summer (ESM), and mid-summer (MSM, n = 10 plots/treatment). These treatments were compared to two controls (long-term average GSP [LTA] and ambient GSP [AMB]) and a wet treatment (+30% of the long-term average GSP [WET]). We found that LSP drought did not significantly reduce ANPP relative to control plots while the ESM and MSM drought did despite equivalent reductions in soil moisture. In contrast, the WET treatment did not affect ANPP. Neither the WET nor the drought treatments altered BNPP as compared to the controls. Our results suggest that forecasts of ecosystem responses to climate change will be improved if both the seasonal timing of alterations in precipitation as well as differential responses of above- and belowground productivity to drought are incorporated into models.     

Drought effects on above- and belowground production of a grazed temperate grassland ecosystem

The effect of climatic variation on terrestrial aboveground productivity (ANPP) has been well studied. However, little is known about how variable climate, including drought, may influence belowground productivity (BNPP), which constitutes most of the annual primary production of grasslands. The objectives of this study were to (1) examine how a 3-year period of declining moisture, which began as climatically wet to average across Yellowstone National Park (YNP) and ended in drought, affected ANPP and BNPP in grasslands of YNP and (2) how herds of grazing ungulates, which were shown previously to stimulate grassland shoot and root growth in YNP, may have interacted with climatic conditions to influence grassland production. ANPP and 0–20 cm BNPP, representing the bulk of the root dynamics, were measured in grazed and ungrazed (fenced) grassland at nine sites ranging widely in elevation, soil conditions and plant production during the 3-year study. Results revealed that 0–20 cm BNPP was strongly influenced by drought (P = 0.0005) and declined from 1999 to 2001 among ungrazed and grazed grasslands by 39 and 49%, respectively. The greater reduction in 0–20 cm BNPP among grazed grasslands was due, in part, to a decline (P = 0.07) in the stimulatory effect of grazing, i.e., the ratio g BNPP stimulated: g shoot consumed. In contrast, ANPP was unaffected by drought in either type of grassland. Thus, the effect of this drought in YNP was a large reduction in BNPP, which was a function of (1) a direct negative influence of increased moisture stress on root growth and (2) a weak interaction between drought and herbivory that led to a decline in the positive feedback from grazers to BNPP. These findings highlight the need to better understand factors that control root growth and to study the effects of climatic variation on grasslands within an ecosystem framework to include potentially important climate–consumer interactions.     

Regional grassland productivity responses to precipitation during multiyear above‐ and below‐average rainfall periods

There is considerable uncertainty in the magnitude and direction of changes in precipitation associated with climate change, and ecosystem responses are also uncertain. Multiyear periods of above‐ and below‐average rainfall may foretell consequences of changes in rainfall regime. We compiled long‐term aboveground net primary productivity (ANPP) and precipitation (PPT) data for eight North American grasslands, and quantified relationships between ANPP and PPT at each site, and in 1–3 year periods of above‐ and below‐average rainfall for mesic, semiarid cool, and semiarid warm grassland types. Our objective was to improve understanding of ANPP dynamics associated with changing climatic conditions by contrasting PPT–ANPP relationships in above‐ and below‐average PPT years to those that occurred during sequences of multiple above‐ and below‐average years. We found differences in PPT–ANPP relationships in above‐ and below‐average years compared to long‐term site averages, and variation in ANPP not explained by PPT totals that likely are attributed to legacy effects. The correlation between ANPP and current‐ and prior‐year conditions changed from year to year throughout multiyear periods, with some legacy effects declining, and new responses emerging. Thus, ANPP in a given year was influenced by sequences of conditions that varied across grassland types and climates. Most importantly, the influence of prior‐year ANPP often increased with the length of multiyear periods, whereas the influence of the amount of current‐year PPT declined. Although the mechanisms by which a directional change in the frequency of above‐ and below‐average years imposes a persistent change in grassland ANPP require further investigation, our results emphasize the importance of legacy effects on productivity for sequences of above‐ vs. below‐average years, and illustrate the utility of long‐term data to examine these patterns.     

Shift in community structure in an early‐successional Mediterranean shrubland driven by long‐term experimental warming and drought and natural extreme droughts

Global warming and recurring drought are expected to accelerate water limitation for plant communities in semiarid Mediterranean ecosystems and produce directional shifts in structure and composition that are not easily detected, and supporting evidence is scarce. We conducted a long‐term (17 years) nocturnal‐warming (+0.6°C) and drought (?40% rainfall) experiments in an early‐successional Mediterranean shrubland to study the changes in community structure and composition, contrasting functional groups and dominant species, and the superimposed effects of natural extreme drought. Species richness decreased in both the warming and drought treatments. Responses to the moderate warming were associated with decreases in herb abundance, and responses to the drought were associated with decreases in both herb and shrub abundances. The drought also significantly decreased community diversity and evenness. Changes in abundance differed between herbs (decreases) and shrubs (increases or no changes). Both warming and drought, especially drought, increased the relative species richness and abundance of shrubs, favoring the establishment of shrubs. Both warming and drought produced significant shifts in plant community composition. Experimental warming shifted the community composition from Erica multiflora toward Rosmarinus officinalis, and drought consistently shifted the composition toward Globularia alypum. The responses in biodiversity (e.g., community biodiversity, changes of functional groups and compositional shifts) were also strongly correlated with atmospheric drought (SPEI) in winter–spring and/or summer, indicating sensitivity to water limitation in this early‐successional Mediterranean ecosystem, especially to natural extreme droughts. Our results suggest that the shifts in species assembles and community diversity and composition are accelerated by the long‐term nocturnal‐warming and drought, combined with natural severe droughts, and that the magnitude of the impacts of climate change is also correlated with the successional status of ecosystem. The results thus highlight the necessity for assessing the impacts on ecosystemic functioning and services and developing effective measures for conserving biodiversity.     

Contingent productivity responses to more extreme rainfall regimes across a grassland biome

Climate models predict, and empirical evidence confirms, that more extreme precipitation regimes are occurring in tandem with warmer atmospheric temperatures. These more extreme rainfall patterns are characterized by increased event size separated by longer within season drought periods and represent novel climatic conditions whose consequences for different ecosystem types are largely unknown. Here, we present results from an experiment in which more extreme rainfall patterns were imposed in three native grassland sites in the Central Plains Region of North America, USA. Along this 600 km precipitation–productivity gradient, there was strong sensitivity of temperate grasslands to more extreme growing season rainfall regimes, with responses of aboveground net primary productivity (ANPP) contingent on mean soil water levels for different grassland types. At the mesic end of the gradient (tallgrass prairie), longer dry intervals between events led to extended periods of below-average soil water content, increased plant water stress and reduced ANPP by 18%. The opposite response occurred at the dry end (semiarid steppe), where a shift to fewer, but larger, events increased periods of above-average soil water content, reduced seasonal plant water stress and resulted in a 30% increase in ANPP. At an intermediate mixed grass prairie site with high plant species richness, ANPP was most sensitive to more extreme rainfall regimes (70% increase). These results highlight the inherent complexity in predicting how terrestrial ecosystems will respond to forecast novel climate conditions as well as the difficulties in extending inferences from single site experiments across biomes. Even with no change in annual precipitation amount, ANPP responses in a relatively uniform physiographic region differed in both magnitude and direction in response to within season changes in rainfall event size/frequency.     

Biological invasions and climate change amplify each other’s effects on dryland degradation

Climate models predict that, in the coming decades, many arid regions will experience increasingly hot conditions and will be affected more frequently by drought. These regions are also experiencing rapid vegetation change, notably invasion by exotic grasses. Invasive grasses spread rapidly into native desert ecosystems due, in particular, to interannual variability in precipitation and periodic fires. The resultant destruction of non-fire-adapted native shrub and grass communities and of the inherent soil resource heterogeneity can yield invader-dominated grasslands. Moreover, recurrent droughts are expected to cause widespread physiological stress and mortality of both invasive and native plants, as well as the loss of soil resources. However, the magnitude of these effects may differ between invasive and native grasses, especially under warmer conditions, rendering the trajectory of vegetated communities uncertain. Using the Biosphere 2 facility in the Sonoran Desert, we evaluated the viability of these hypothesized relationships by simulating combinations of drought and elevated temperature (+5°C) and assessing the ecophysiological and mortality responses of both a dominant invasive grass (Pennisetum ciliare or buffelgrass) and a dominant native grass (Heteropogan contortus or tanglehead). While both grasses survived protracted drought at ambient temperatures by inducing dormancy, drought under warmed conditions exceeded the tolerance limits of the native species, resulting in greater and more rapid mortality than exhibited by the invasive. Thus, two major drivers of global environmental change, biological invasion and climate change, can be expected to synergistically accelerate ecosystem degradation unless large-scale interventions are enacted.     

Drought legacies are dependent on water table depth,wood anatomy and drought timing across the eastern US

Severe droughts can impart long‐lasting legacies on forest ecosystems through lagged effects that hinder tree recovery and suppress whole‐forest carbon uptake. However, the local climatic and edaphic factors that interact to affect drought legacies in temperate forests remain unknown. Here, we pair a dataset of 143 tree ring chronologies across the mesic forests of the eastern US with historical climate and local soil properties. We found legacy effects to be widespread, the magnitude of which increased markedly in diffuse porous species, sites with deep water tables, and in response to late‐season droughts (August–September). Using an ensemble of downscaled climate projections, we additionally show that our sites are projected to drastically increase in water deficit and drought frequency by the end of the century, potentially increasing the size of legacy effects by up to 65% and acting as a significant process shaping forest composition, carbon uptake and mortality.     

Hyposensitive canopy conductance renders ecosystems vulnerable to meteorological droughts

Increased meteorological drought intensity with rising atmospheric demand for water (hereafter vapor pressure deficit [VPD]) increases the risk of tree mortality and ecosystem dysfunction worldwide. Ecosystem-scale water-use strategy is increasingly recognized as a key factor in regulating drought-related ecosystem responses. However, the link between water-use strategy and ecosystem vulnerability to meteorological droughts is poorly established. Using the global flux observations, historic hydroclimatic data, remote-sensing products, and plant functional-trait archive, we identified potentially vulnerable ecosystems, examining how ecosystem water-use strategy, quantified by the percentage bias (δ) of the empirical canopy conductance sensitivity to VPD relative to the theoretical value, mediated ecosystem responses to droughts. We found that prevailing soil water availability substantially impacted δ in dryland regions where ecosystems with insufficient soil moisture usually showed conservative water-use strategy, while ecosystems in humid regions exhibited more pronounced climatic adaptability. Hyposensitive and hypersensitive ecosystems, classified based on δ falling below or above the theoretical sensitivity, respectively, achieved similar net ecosystem productivity during droughts, employing different structural and functional strategies. However, hyposensitive ecosystems, risking their hydraulic system with a permissive water-use strategy, were unable to recover from droughts as quickly as hypersensitive ones. Our findings highlight that processed-based models predicting current functions and future performance of vegetation should account for the greater vulnerability of hyposensitive ecosystems to intensifying atmospheric and soil droughts.     

Drought effects on litterfall, wood production and belowground carbon cycling in an Amazon forest: results of a throughfall reduction experiment

The Amazon Basin experiences severe droughts that may become more common in the future. Little is known of the effects of such droughts on Amazon forest productivity and carbon allocation. We tested the prediction that severe drought decreases litterfall and wood production but potentially has multiple cancelling effects on belowground production within a 7-year partial throughfall exclusion experiment. We simulated an approximately 35-41% reduction in effective rainfall from 2000 through 2004 in a 1ha plot and compared forest response with a similar control plot. Wood production was the most sensitive component of above-ground net primary productivity (ANPP) to drought, declining by 13% the first year and up to 62% thereafter. Litterfall declined only in the third year of drought, with a maximum difference of 23% below the control plot. Soil CO2 efflux and its 14C signature showed no significant treatment response, suggesting similar amounts and sources of belowground production. ANPP was similar between plots in 2000 and declined to a low of 41% below the control plot during the subsequent treatment years, rebounding to only a 10% difference during the first post-treatment year. Live aboveground carbon declined by 32.5Mgha-1 through the effects of drought on ANPP and tree mortality. Results of this unreplicated, long-term, large-scale ecosystem manipulation experiment demonstrate that multi-year severe drought can substantially reduce Amazon forest carbon stocks.     

Not all droughts are created equal: the impacts of interannual drought pattern and magnitude on grassland carbon cycling

Climate extremes, such as drought, may have immediate and potentially prolonged effects on carbon cycling. Grasslands store approximately one‐third of all terrestrial carbon and may become carbon sources during droughts. However, the magnitude and duration of drought‐induced disruptions to the carbon cycle, as well as the mechanisms responsible, remain poorly understood. Over the next century, global climate models predict an increase in two types of drought: chronic but subtle ‘press‐droughts’, and shorter term but extreme ‘pulse‐droughts’. Much of our current understanding of the ecological impacts of drought comes from experimental rainfall manipulations. These studies have been highly valuable, but are often short term and rarely quantify carbon feedbacks. To address this knowledge gap, we used the Community Land Model 4.0 to examine the individual and interactive effects of pulse‐ and press‐droughts on carbon cycling in a mesic grassland of the US Great Plains. A series of modeling experiments were imposed by varying drought magnitude (precipitation amount) and interannual pattern (press‐ vs. pulse‐droughts) to examine the effects on carbon storage and cycling at annual to century timescales. We present three main findings. First, a single‐year pulse‐drought had immediate and prolonged effects on carbon storage due to differential sensitivities of ecosystem respiration and gross primary production. Second, short‐term pulse‐droughts caused greater carbon loss than chronic press‐droughts when total precipitation reductions over a 20‐year period were equivalent. Third, combining pulse‐ and press‐droughts had intermediate effects on carbon loss compared to the independent drought types, except at high drought levels. Overall, these results suggest that interannual drought pattern may be as important for carbon dynamics as drought magnitude and that extreme droughts may have long‐lasting carbon feedbacks in grassland ecosystems.     

天山北坡不同海拔梯度山地草原生态系统地上净初级生产力对气候变化及放牧的响应

以天山北坡三工河流域为例,利用改进后的Biome-BGC模型分别模拟了仅气候变化（Clm）、气候变化与放牧联合作用（ClmGra）下研究区不同海拔梯度3种山地草原生态系统（低山干旱草原,LAG;森林草甸草原,FMG;高寒草甸草原,AMG）1959—2009年地上净初级生产力（Aboveground Net Primary Production,ANPP）的动态,并通过假设27种放牧强度情景（0—8 羊/ha）模拟了其ANPP随放牧强度增加的变化趋势。近50年气候变化致使研究区各海拔梯度草原生态系统ANPP整体均呈上升趋势,但在放牧联合作用下,不同草原类型ANPP变化趋势差异显著;放牧导致FMG和AMG的ANPP呈下降态势,分别减少30.0%和33.2%,对比之下,由于1980前较低放牧强度促进了LAG的ANPP,放牧导致其ANPP整体增加1.3%。随着放牧强度增加,LAG的ANPP呈先增后减趋势,且在干旱年份最为显著;而FMG和AMG的ANPP呈显著非线性递减趋势。这些结果表明,近50年气候波动可能有利于中亚干旱区山地草原生态系统生产力的提高,但日益增强的放牧活动导致其净初级生产力显著降低;放牧对FMG与AMG生产力的负面效应随放牧强度增加而增强,但适度放牧可能促进LAG净初级生产力,尤其在干旱年份。     

High ecosystem stability of evergreen broadleaf forests under severe droughts

Global increase in drought occurrences threatens the stability of terrestrial ecosystem functioning. Evergreen broadleaf forests (EBFs) keep leaves throughout the year, and therefore could experience higher drought risks than other biomes. However, the recent temporal variability of global vegetation productivity or land carbon sink is mainly driven by non‐evergreen ecosystems, such as semiarid grasslands, croplands, and boreal forests. Thus, we hypothesize that EBFs have higher stability than other biomes under the increasingly extreme droughts. Here we use long‐term Standardized Precipitation and Evaporation Index (SPEI) data and satellite‐derived Enhanced Vegetation Index (EVI) products to quantify the temporal stability (ratio of mean annual EVI to its SD), resistance (ability to maintain its original levels during droughts), and resilience (rate of EVI recovering to pre‐drought levels) at biome and global scales. We identified significantly increasing trends of annual drought severity (SPEI range: ?0.08 to ?1.80), area (areal fraction range: 2%–19%), and duration (month range: 7.9–9.1) in the EBF biome over 2000–2014. However, EBFs showed the highest resistance of EVI to droughts, but no significant differences in resilience of EVI to droughts were found among biomes (forests, grasslands, savannas, and shrublands). Global resistance and resilience of EVI to droughts were largely affected by temperature and solar radiation. These findings suggest that EBFs have higher stability than other biomes despite the greater drought exposure. Thus, the conservation of EBFs is critical for stabilizing global vegetation productivity and land carbon sink under more‐intense climate extremes in the future.