Effects of climate extremes on the terrestrial carbon cycle: concepts,processes and potential future impacts

Extreme droughts, heat waves, frosts, precipitation, wind storms and other climate extremes may impact the structure, composition and functioning of terrestrial ecosystems, and thus carbon cycling and its feedbacks to the climate system. Yet, the interconnected avenues through which climate extremes drive ecological and physiological processes and alter the carbon balance are poorly understood. Here, we review the literature on carbon cycle relevant responses of ecosystems to extreme climatic events. Given that impacts of climate extremes are considered disturbances, we assume the respective general disturbance‐induced mechanisms and processes to also operate in an extreme context. The paucity of well‐defined studies currently renders a quantitative meta‐analysis impossible, but permits us to develop a deductive framework for identifying the main mechanisms (and coupling thereof) through which climate extremes may act on the carbon cycle. We find that ecosystem responses can exceed the duration of the climate impacts via lagged effects on the carbon cycle. The expected regional impacts of future climate extremes will depend on changes in the probability and severity of their occurrence, on the compound effects and timing of different climate extremes, and on the vulnerability of each land‐cover type modulated by management. Although processes and sensitivities differ among biomes, based on expert opinion, we expect forests to exhibit the largest net effect of extremes due to their large carbon pools and fluxes, potentially large indirect and lagged impacts, and long recovery time to regain previous stocks. At the global scale, we presume that droughts have the strongest and most widespread effects on terrestrial carbon cycling. Comparing impacts of climate extremes identified via remote sensing vs. ground‐based observational case studies reveals that many regions in the (sub‐)tropics are understudied. Hence, regional investigations are needed to allow a global upscaling of the impacts of climate extremes on global carbon–climate feedbacks.     

Growth and resilience responses of Scots pine to extreme droughts across Europe depend on predrought growth conditions

Global climate change is expected to further raise the frequency and severity of extreme events, such as droughts. The effects of extreme droughts on trees are difficult to disentangle given the inherent complexity of drought events (frequency, severity, duration, and timing during the growing season). Besides, drought effects might be modulated by trees’ phenotypic variability, which is, in turn, affected by long‐term local selective pressures and management legacies. Here we investigated the magnitude and the temporal changes of tree‐level resilience (i.e., resistance, recovery, and resilience) to extreme droughts. Moreover, we assessed the tree‐, site‐, and drought‐related factors and their interactions driving the tree‐level resilience to extreme droughts. We used a tree‐ring network of the widely distributed Scots pine (Pinus sylvestris) along a 2,800 km latitudinal gradient from southern Spain to northern Germany. We found that the resilience to extreme drought decreased in mid‐elevation and low productivity sites from 1980–1999 to 2000–2011 likely due to more frequent and severe droughts in the later period. Our study showed that the impact of drought on tree‐level resilience was not dependent on its latitudinal location, but rather on the type of sites trees were growing at and on their growth performances (i.e., magnitude and variability of growth) during the predrought period. We found significant interactive effects between drought duration and tree growth prior to drought, suggesting that Scots pine trees with higher magnitude and variability of growth in the long term are more vulnerable to long and severe droughts. Moreover, our results indicate that Scots pine trees that experienced more frequent droughts over the long‐term were less resistant to extreme droughts. We, therefore, conclude that the physiological resilience to extreme droughts might be constrained by their growth prior to drought, and that more frequent and longer drought periods may overstrain their potential for acclimation.     

Near‐future forest vulnerability to drought and fire varies across the western United States

Recent prolonged droughts and catastrophic wildfires in the western United States have raised concerns about the potential for forest mortality to impact forest structure, forest ecosystem services, and the economic vitality of communities in the coming decades. We used the Community Land Model (CLM) to determine forest vulnerability to mortality from drought and fire by the year 2049. We modified CLM to represent 13 major forest types in the western United States and ran simulations at a 4‐km grid resolution, driven with climate projections from two general circulation models under one emissions scenario (RCP 8.5). We developed metrics of vulnerability to short‐term extreme and prolonged drought based on annual allocation to stem growth and net primary productivity. We calculated fire vulnerability based on changes in simulated future area burned relative to historical area burned. Simulated historical drought vulnerability was medium to high in areas with observations of recent drought‐related mortality. Comparisons of observed and simulated historical area burned indicate simulated future fire vulnerability could be underestimated by 3% in the Sierra Nevada and overestimated by 3% in the Rocky Mountains. Projections show that water‐limited forests in the Rocky Mountains, Southwest, and Great Basin regions will be the most vulnerable to future drought‐related mortality, and vulnerability to future fire will be highest in the Sierra Nevada and portions of the Rocky Mountains. High carbon‐density forests in the Pacific coast and western Cascades regions are projected to be the least vulnerable to either drought or fire. Importantly, differences in climate projections lead to only 1% of the domain with conflicting low and high vulnerability to fire and no area with conflicting drought vulnerability. Our drought vulnerability metrics could be incorporated as probabilistic mortality rates in earth system models, enabling more robust estimates of the feedbacks between the land and atmosphere over the 21st century.     

极端降水和极端干旱事件对草原生态系统的影响

当前人类活动的加剧显著地影响着全球大气循环的格局。大气循环的多个模型均预测未来全球气候变化的显著特征是极端降水事件和极端干旱事件发生的频率会显著增加。水分是干旱、半干旱区草原植物生长发育的限制性资源, 而草原生态系统是陆地生态系统中对降水格局变化非常敏感的系统。但是, 关于极端降水事件和极端干旱事件对草原生态系统结构和功能的影响还是以分散的个案研究为主, 甚至关于极端气候事件的定义迄今也不尽相同。为此, 该文在分析极端气候事件定义及其研究方法的基础上, 总结了极端降水事件和极端干旱事件对草原生态系统土壤水分和养分状况、植物生长发育和生理特性、群落结构、生产力和碳循环过程的影响, 并提出了未来极端气候事件研究中应重点关注的5个重要方向, 以及控制试验研究的2个关键科学问题, 对开展全球变化背景下草原生态系统对极端气候事件响应机制的研究具有指导意义。     

Carbon and Nitrogen Decoupling Under an 11-Year Drought in the Shortgrass Steppe

The frequency and magnitude of drought is expected to increase in the US Great Plains under future climate regimes. Although semiarid systems are considered highly resistant to water limitation, novel drought events could alter linkages among biogeochemical processes, and result in new feedbacks that influence the timescale of ecosystem recovery. We examined changes in carbon and nitrogen cycling in the last 2 years of an 11-year drought manipulation in the shortgrass steppe, and under the first 2 years of recovery from drought. We measured plant production, plant tissue chemistry, soil trace gas flux, and soil inorganic nitrogen dynamics to test the extent that this magnitude of drought altered carbon and nitrogen fluxes and how these changes affected post-drought dynamics. We found that soil inorganic nitrogen was up to five times higher under severe drought than under control conditions, but that this nitrogen may not have been accessible to plants and microbial communities during drought due to diffusion limitations. Drought plots had higher N₂O flux when they received equal rainfall pulses, showing that this accumulated N may be vulnerable to loss. In addition, plants in drought plots had higher tissue nitrogen for 2 years following drought. These results show that decadal-length droughts that may occur under future precipitation regimes are likely to alter ecosystem properties through interactions among precipitation, vegetation, and N cycling. Shifts in plant N, vulnerability of nitrogen to loss, and rainfall use efficiency that we observed are likely to affect the recovery time of semiarid systems subject to droughts of this magnitude.     

Drought and aquatic ecosystems: an introduction

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Forest biogeochemistry in response to drought

Trees alter their use and allocation of nutrients in response to drought, and changes in soil nutrient cycling and trace gas flux (N₂O and CH₄) are observed when experimental drought is imposed on forests. In extreme droughts, trees are increasingly susceptible to attack by pests and pathogens, which can lead to major changes in nutrient flux to the soil. Extreme droughts often lead to more common and more intense forest fires, causing dramatic changes in the nutrient storage and loss from forest ecosystems. Changes in the future manifestation of drought will affect carbon uptake and storage in forests, leading to feedbacks to the Earth's climate system. We must improve the recognition of drought in nature, our ability to manage our forests in the face of drought, and the parameterization of drought in earth system models for improved predictions of carbon uptake and storage in the world's forests.     

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.     

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.     

Critical transitions in rainfall manipulation experiments on grasslands

As a result of climate and land‐use changes, grasslands have been subjected to intensifying drought regimes. Extreme droughts could interfere in the positive feedbacks between grasses and soil water content, pushing grasslands across critical thresholds of productivity and leading them to collapse. If this happens, systems may show hysteresis and costly management interventions might be necessary to restore predrought productivity. Thus, neglecting critical transitions may lead to mismanagement of grasslands and to irreversible loss of ecosystem services. Rainfall manipulation experiments constitute a powerful approach to investigate the risk of such critical transitions. However, experiments performed to date have rarely applied extreme droughts and have used resilience indices that disregard the existence of hysteresis. Here, we suggest how to incorporate critical transitions when designing rainfall manipulation experiments on grasslands and when measuring their resilience to drought. The ideas presented here have the potential to trigger a perspective shift among experimental researchers, into a new state where the existence of critical transitions will be discussed, experimentally tested, and largely considered when assessing and managing vegetation resilience to global changes.     

Effects of Extreme Weather Events on Plant Productivity and Tissue Die-Back are Modified by Community Composition

Extreme weather events are expected to increase in frequency and magnitude due to climate change. Their effects on vegetation are widely unknown. Here, experimental grassland and heath communities in Central Europe were exposed either to a simulated single drought or to a prolonged heavy rainfall event. The magnitude of manipulations imitated the local 100-year weather extreme according to extreme value statistics. Overall productivity of both plant communities remained stable in the face of drought and heavy rainfall, despite significant effects on tissue die-back. Grassland communities were more resistant against the extreme weather events than heath communities. Furthermore, effects of extreme weather events on community tissue die-back were modified by functional diversity, even though conclusiveness in this part is limited by the fact that only one species composition was available per diversity level within this case study. More diverse grassland communities exhibited less tissue die-back than less complex grassland communities. On the other side, more diverse heath communities were more vulnerable to extreme weather events compared to less complex heath communities. Furthermore, legumes did not effectively contribute to the buffering against extreme weather events in both vegetation types. Tissue die-back proved a strong stress response in plant communities exposed to 100-year extreme weather events, even though one important ecosystem function, namely productivity, remained surprisingly stable in this experiment. Theories and concepts on biodiversity and ecosystem functioning (insurance hypothesis, redundancy hypothesis) may have to be revisited when extreme weather conditions are considered.     

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.     

Drought survival is a threshold function of habitat size and population density in a fish metapopulation

Because smaller habitats dry more frequently and severely during droughts, habitat size is likely a key driver of survival in populations during climate change and associated increased extreme drought frequency. Here, we show that survival in populations during droughts is a threshold function of habitat size driven by an interaction with population density in metapopulations of the forest pool dwelling fish, Neochanna apoda. A mark–recapture study involving 830 N. apoda individuals during a one‐in‐seventy‐year extreme drought revealed that survival during droughts was high for populations occupying pools deeper than 139 mm, but declined steeply in shallower pools. This threshold was caused by an interaction between increasing population density and drought magnitude associated with decreasing habitat size, which acted synergistically to increase physiological stress and mortality. This confirmed two long‐held hypotheses, firstly concerning the interactive role of population density and physiological stress, herein driven by habitat size, and secondly, the occurrence of drought survival thresholds. Our results demonstrate how survival in populations during droughts will depend strongly on habitat size and highlight that minimum habitat size thresholds will likely be required to maximize survival as the frequency and intensity of droughts are projected to increase as a result of global climate change.     

Short-term climatic trends affect the temporal variability of macroinvertebrates in California 'Mediterranean' streams

1. Long‐term studies in ecology are essential for understanding natural variability and in interpreting responses to disturbances and human perturbations. We assessed the long‐term variability, stability and persistence of macroinvertebrate communities by analysing data from three regions in northern California with a mediterranean‐climate. During the study period, precipitation either increased or decreased, and extreme drought events occurred in each region. 2. Temporal trends in precipitation resulted in shifts from ‘dry‐year’ communities, dominated by taxa adapted to no or low flow, to ‘wet‐year’ communities dominated by taxa adapted to high flows. The abundance of chironomid larvae was an important driver of community change. Directional change in community composition occurred at all sites and was correlated with precipitation patterns, with more dramatic change occurring in smaller streams. 3. All communities exhibited high to moderate persistence (defined by the presence/absence of a species) and moderate to low stability (defined by changes in abundance) over the study period. Stability and persistence were correlated with climatic variation (precipitation and El Niño Southern Oscillation) and stream size. Stability and persistence increased as a result of drought in small streams (first‐order) but decreased in larger streams (second‐ and third‐order). Communities from the dry season were less stable than those from the wet‐season. 4. This study demonstrates the importance of long‐term studies in capturing the effects of and recovery from rare events, such as the prolonged and extreme droughts considered here.     

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.     

Climate change and variability impacts on grazing herds: Insights from a system dynamics approach for semi‐arid Australian rangelands

Grazing livestock are an important source of food and income for millions of people worldwide. Changes in mean climate and increasing climate variability are affecting grasslands' carrying capacity, thus threatening the livelihood of millions of people as well as the health of grassland ecosystems. Compared with cropping systems, relatively little is known about the impact of such climatic changes on grasslands and livestock productivity and the adaptation responses available to farmers. In this study, we analysed the relationship between changes in mean precipitation, precipitation variability, farming practices and grazing cattle using a system dynamics approach for a semi‐arid Australian rangeland system. We found that forage production and animal stocking rates were significantly affected by drought intensities and durations as well as by long‐term climate trends. After a drought event, herd size recovery times ranged from years to decades in the absence of proactive restocking through animal purchases. Decreases in the annual precipitation means or increases in the interannual (year‐to‐year) and intra‐annual (month‐to‐month) precipitation variability, all reduced herd sizes. The contribution of farming practices versus climate effect on herd dynamics varied depending on the herd characteristics considered. Climate contributed the most to the variance in stocking rates, followed by forage productivity levels and feeding supplementation practices (with or without urea and molasses). While intensification strategies and favourable climates increased long‐term herd sizes, they also resulted in larger reductions in animal numbers during droughts and raised total enteric methane emissions. In the face of future climate trends, the grazing sector will need to increase its adaptability. Understanding which farming strategies can be beneficial, where, and when, as well as the enabling mechanisms required to implement them, will be critical for effectively improving rangelands and the livelihoods of pastoralists worldwide.     

Amazon drought and forest response: Largely reduced forest photosynthesis but slightly increased canopy greenness during the extreme drought of 2015/2016

Amazon droughts have impacted regional ecosystem functioning as well as global carbon cycling. The severe dry‐season droughts in 2005 and 2010, driven by Atlantic sea surface temperature (SST) anomaly, have been widely investigated in terms of drought severity and impacts on ecosystems. Although the influence of Pacific SST anomaly on wet‐season precipitation has been well recognized, it remains uncertain to what extent the droughts driven by Pacific SST anomaly could affect forest greenness and photosynthesis in the Amazon. Here, we examined the monthly and annual dynamics of forest greenness and photosynthetic capacity when Amazon ecosystems experienced an extreme drought in 2015/2016 driven by a strong El Niño event. We found that the drought during August 2015–July 2016 was one of the two most severe meteorological droughts since 1901. Due to the enhanced solar radiation during this drought, overall forest greenness showed a small increase, and 21.6% of forests even greened up (greenness index anomaly ≥1 standard deviation). In contrast, solar‐induced chlorophyll fluorescence (SIF), an indicator of vegetation photosynthetic capacity, showed a significant decrease. Responses of forest greenness and photosynthesis decoupled during this drought, indicating that forest photosynthesis could still be suppressed regardless of the variation in canopy greenness. If future El Niño frequency increases as projected by earth system models, droughts would result in persistent reduction in Amazon forest productivity, substantial changes in tree composition, and considerable carbon emissions from Amazon.     

Drought timing and local climate determine the sensitivity of eastern temperate forests to drought

Projected changes in temperature and drought regime are likely to reduce carbon (C) storage in forests, thereby amplifying rates of climate change. While such reductions are often presumed to be greatest in semi‐arid forests that experience widespread tree mortality, the consequences of drought may also be important in temperate mesic forests of Eastern North America (ENA) if tree growth is significantly curtailed by drought. Investigations of the environmental conditions that determine drought sensitivity are critically needed to accurately predict ecosystem feedbacks to climate change. We matched site factors with the growth responses to drought of 10,753 trees across mesic forests of ENA, representing 24 species and 346 stands, to determine the broad‐scale drivers of drought sensitivity for the dominant trees in ENA. Here we show that two factors—the timing of drought, and the atmospheric demand for water (i.e., local potential evapotranspiration; PET)—are stronger drivers of drought sensitivity than soil and stand characteristics. Drought‐induced reductions in tree growth were greatest when the droughts occurred during early‐season peaks in radial growth, especially for trees growing in the warmest, driest regions (i.e., highest PET). Further, mean species trait values (rooting depth and ψ₅₀) were poor predictors of drought sensitivity, as intraspecific variation in sensitivity was equal to or greater than interspecific variation in 17 of 24 species. From a general circulation model ensemble, we find that future increases in early‐season PET may exacerbate these effects, and potentially offset gains in C uptake and storage in ENA owing to other global change factors.     

Negative feedback processes following drainage slow down permafrost degradation

The sustainability of the vast Arctic permafrost carbon pool under climate change is of paramount importance for global climate trajectories. Accurate climate change forecasts, therefore, depend on a reliable representation of mechanisms governing Arctic carbon cycle processes, but this task is complicated by the complex interaction of multiple controls on Arctic ecosystem changes, linked through both positive and negative feedbacks. As a primary example, predicted Arctic warming can be substantially influenced by shifts in hydrologic regimes, linked to, for example, altered precipitation patterns or changes in topography following permafrost degradation. This study presents observational evidence how severe drainage, a scenario that may affect large Arctic areas with ice‐rich permafrost soils under future climate change, affects biogeochemical and biogeophysical processes within an Arctic floodplain. Our in situ data demonstrate reduced carbon losses and transfer of sensible heat to the atmosphere, and effects linked to drainage‐induced long‐term shifts in vegetation communities and soil thermal regimes largely counterbalanced the immediate drainage impact. Moreover, higher surface albedo in combination with low thermal conductivity cooled the permafrost soils. Accordingly, long‐term drainage effects linked to warming‐induced permafrost degradation hold the potential to alleviate positive feedbacks between permafrost carbon and Arctic warming, and to slow down permafrost degradation. Self‐stabilizing effects associated with ecosystem disturbance such as these drainage impacts are a key factor for predicting future feedbacks between Arctic permafrost and climate change, and, thus, neglect of these mechanisms will exaggerate the impacts of Arctic change on future global climate projections.     

Climate‐driven diversity change in annual grasslands: Drought plus deluge does not equal normal

Climate forecasts agree that increased variability and extremes will tend to reduce the availability of water in many terrestrial ecosystems. Increasingly severe droughts may be exacerbated both by warmer temperatures and by the relative unavailability of water that arrives in more sporadic and intense rainfall events. Using long‐term data and an experimental water manipulation, we examined the resilience of a heterogeneous annual grassland community to a prolonged series of dry winters that led to a decline in plant species richness (2000–2014), followed by a near‐record wet winter (2016–2017), a climatic sequence that broadly resembles the predicted future in its high variability. In our 80, 5‐m² observational plots, species richness did not recover in response to the wet winter, and the positive relationship of richness to annual winter rainfall thus showed a significant weakening trend over the 18‐year time period. In experiments on 100, 1‐m² plots, wintertime water supplementation increased and drought shelters decreased the seedling survival and final individual biomass of native annual forbs, the main functional group contributing to the observed long‐term decline in richness. Water supplementation also increased the total cover of native annual forbs, but only increased richness within nested subplots to which seeds were also added. We conclude that prolonged dry winters, by increasing seedling mortality and reducing growth of native forbs, may have diminished the seedbank and thus the recovery potential of diversity in this community. However, the wet winter and the watering treatment did cause recovery of the community mean values of a key functional trait (specific leaf area, an indicator of drought intolerance), suggesting that some aggregate community properties may be stabilized by functional redundancy among species.