A B S T R A C T Greenhouse gas emissions have significantly altered global climate, and will continue to do so in the future. Increases in the frequency, duration, and/or severity of drought and heat stress associated with climate change could fundamentally alter the composition, structure, and biogeography of forests in many regions. Of particular concern are potential increases in tree mortality associated with climateinduced physiological stress and interactions with other climate-mediated processes such as insect outbreaks and wildfire. Despite this risk, existing projections of tree mortality are based on models that lack functionally realistic mortality mechanisms, and there has been no attempt to track observations of climate-driven tree mortality globally. Here we present the first global assessment of recent tree mortality attributed to drought and heat stress. Although episodic mortality occurs in the absence of climate change, studies compiled here suggest that at least some of the world's forested ecosystems already may be responding to climate change and raise concern that forests may become increasingly vulnerable to higher background tree mortality rates and die-off in response to future warming and drought, even in environments that are not normally considered water-limited. This further suggests risks to ecosystem services, including the loss of sequestered forest carbon and associated atmospheric feedbacks. Our review also identifies key information gaps and scientific uncertainties that currently hinder our ability to predict tree mortality in response to climate change and emphasizes the need for a globally coordinated observation system. Overall, our review reveals the potential for amplified tree mortality due to drought and heat in forests worldwide.Published by Elsevier B.V.

Contents Summary1I.Introduction2II. Consequences of vegetation mortality3III.Global patterns of mortality3IV.Hypotheses on mechanisms of drought‐related mortality4V.Evidence for hypothesized mechanisms5VI.Implications of future climate on hypothesized mortality mechanisms13VII.Conclusions15Acknowledgements15References15 Summary Severe droughts have been associated with regional‐scale forest mortality worldwide. Climate change is expected to exacerbate regional mortality events; however, prediction remains difficult because the physiological mechanisms underlying drought survival and mortality are poorly understood. We developed a hydraulically based theory considering carbon balance and insect resistance that allowed development and examination of hypotheses regarding survival and mortality. Multiple mechanisms may cause mortality during drought. A common mechanism for plants with isohydric regulation of water status results from avoidance of drought‐induced hydraulic failure via stomatal closure, resulting in carbon starvation and a cascade of downstream effects such as reduced resistance to biotic agents. Mortality by hydraulic failure per se may occur for isohydric seedlings or trees near their maximum height. Although anisohydric plants are relatively drought‐tolerant, they are predisposed to hydraulic failure because they operate with narrower hydraulic safety margins during drought. Elevated temperatures should exacerbate carbon starvation and hydraulic failure. Biotic agents may amplify and be amplified by drought‐induced plant stress. Wet multidecadal climate oscillations may increase plant susceptibility to drought‐induced mortality by stimulating shifts in hydraulic architecture, effectively predisposing plants to water stress. Climate warming and increased frequency of extreme events will probably cause increased regional mortality episodes. Isohydric and anisohydric water potential regulation may partition species between survival and mortality, and, as such, incorporating this hydraulic framework may be effective for modeling plant survival and mortality under future climate conditions.

Future drought is projected to occur under warmer temperature conditions as climate change progresses, referred to here as globalchange-type drought, yet quantitative assessments of the triggers and potential extent of drought-induced vegetation die-off remain pivotal uncertainties in assessing climate-change impacts. Of particular concern is regional-scale mortality of overstory trees, which rapidly alters ecosystem type, associated ecosystem properties, and land surface conditions for decades. Here, we quantify regional-scale vegetation die-off across southwestern North American woodlands in 2002-2003 in response to drought and associated bark beetle infestations. At an intensively studied site within the region, we quantified that after 15 months of depleted soil water content, >90% of the dominant, overstory tree species (Pinus edulis, a piñ on) died. The die-off was reflected in changes in a remotely sensed index of vegetation greenness (Normalized Difference Vegetation Index), not only at the intensively studied site but also across the region, extending over 12,000 km 2 or more; aerial and field surveys confirmed the general extent of the die-off. Notably, the recent drought was warmer than the previous subcontinental drought of the 1950s. The limited, available observations suggest that die-off from the recent drought was more extensive than that from the previous drought, extending into wetter sites within the tree species' distribution. Our results quantify a trigger leading to rapid, drought-induced die-off of overstory woody plants at subcontinental scale and highlight the potential for such die-off to be more severe and extensive for future global-change-type drought under warmer conditions. tree mortality ͉ vegetation dynamics ͉ climate change impacts ͉ woodlands ͉ Pinus edulis

Abstract. Patterns, mechanisms, projections, and consequences of tree mortality and associated broadscale forest die-off due to drought accompanied by warmer temperatures-''hotter drought'', an emerging characteristic of the Anthropocene-are the focus of rapidly expanding literature. Despite recent observational, experimental, and modeling studies suggesting increased vulnerability of trees to hotter drought and associated pests and pathogens, substantial debate remains among research, management and policy-making communities regarding future tree mortality risks. We summarize key mortalityrelevant findings, differentiating between those implying lesser versus greater levels of vulnerability. Evidence suggesting lesser vulnerability includes forest benefits of elevated [CO 2 ] and increased water-use efficiency; observed and modeled increases in forest growth and canopy greening; widespread increases in woody-plant biomass, density, and extent; compensatory physiological, morphological, and genetic mechanisms; dampening ecological feedbacks; and potential mitigation by forest management. In contrast, recent studies document more rapid mortality under hotter drought due to negative tree physiological responses and accelerated biotic attacks. Additional evidence suggesting greater vulnerability includes rising background mortality rates; projected increases in drought frequency, intensity, and duration; limitations of vegetation models such as inadequately represented mortality processes; warming feedbacks from die-off; and wildfire synergies. Grouping these findings we identify ten contrasting perspectives that shape the vulnerability debate but have not been discussed collectively. We also present a set of global vulnerability drivers that are known with high confidence: (1) droughts eventually occur everywhere; (2) warming produces hotter droughts; (3) atmospheric moisture demand increases nonlinearly with temperature during drought; (4) mortality can occur faster in hotter drought, consistent with fundamental physiology; (5) shorter droughts occur more frequently than longer droughts and can become lethal under warming, increasing the frequency of lethal drought nonlinearly; and (6) mortality happens rapidly relative to growth intervals needed for forest recovery. These high-confidence drivers, in concert with research supporting greater vulnerability perspectives, support an overall viewpoint of greater forest vulnerability globally. We surmise that mortality vulnerability is being discounted in part due to difficulties in predicting threshold responses to extreme climate events. Given the profound ecological and societal implications of underestimating global vulnerability to hotter drought, we highlight urgent challenges for research, management, and policy-making communities.

As the climate changes, drought may reduce tree productivity and survival across many forest ecosystems; however, the relative influence of specific climate parameters on forest decline is poorly understood. We derive a forest drought-stress index (FDSI) for the southwestern United States using a comprehensive tree-ring data set representing AD 1000-2007. The FDSI is approximately equally influenced by the warm-season vapour-pressure deficit (largely controlled by temperature) and cold-season precipitation, together explaining 82% of the FDSI variability. Correspondence between the FDSI and measures of forest productivity, mortality, bark-beetle outbreak and wildfire validate the FDSI as a holistic forest-vigour indicator. If the vapour-pressure deficit continues increasing as projected by climate models, the mean forest drought-stress by the 2050s will exceed that of the most severe droughts in the past 1,000 years. Collectively, the results foreshadow twenty-first-century changes in forest structures and compositions, with transition of forests in the southwestern United States, and perhaps water-limited forests globally, towards distributions unfamiliar to modern civilization.

Ecological memory is central to how ecosystems respond to disturbance and is maintained by two types of legacies – information and material. Species life‐history traits represent an adaptive response to disturbance and are an information legacy; in contrast, the abiotic and biotic structures (such as seeds or nutrients) produced by single disturbance events are material legacies. Disturbance characteristics that support or maintain these legacies enhance ecological resilience and maintain a “safe operating space” for ecosystem recovery. However, legacies can be lost or diminished as disturbance regimes and environmental conditions change, generating a “resilience debt” that manifests only after the system is disturbed. Strong effects of ecological memory on post‐disturbance dynamics imply that contingencies (effects that cannot be predicted with certainty) of individual disturbances, interactions among disturbances, and climate variability combine to affect ecosystem resilience. We illustrate these concepts and introduce a novel ecosystem resilience framework with examples of forest disturbances, primarily from North America. Identifying legacies that support resilience in a particular ecosystem can help scientists and resource managers anticipate when disturbances may trigger abrupt shifts in forest ecosystems, and when forests are likely to be resilient.

In coming decades, global climate changes are expected to produce large shifts in vegetation distributions at unprecedented rates. These shifts are expected to be most rapid and extreme at ecotones, the boundaries between ecosystems, particularly those in semiarid landscapes. However, current models do not adequately provide for such rapid effects-particularly those caused by mortality-largely because of the lack of data from field studies. Here we report the most rapid landscape-scale shift of a woody ecotone ever documented: in northern New Mexico in the 1950s, the ecotone between semiarid ponderosa pine forest and piñon-juniper woodland shifted extensively (2 km or more) and rapidly (<5 years) through mortality of ponderosa pines in response to a severe drought. This shift has persisted for 40 years. Forest patches within the shift zone became much more fragmented, and soil erosion greatly accelerated. The rapidity and the complex dynamics of the persistent shift point to the need to represent more accurately these dynamics, especially the mortality factor, in assessments of the effects of climate change.

Widespread tree mortality associated with drought has been observed on all forested continents and global change is expected to exacerbate vegetation vulnerability. Forest mortality has implications for future biosphere–atmosphere interactions of carbon, water and energy balance, and is poorly represented in dynamic vegetation models. Reducing uncertainty requires improved mortality projections founded on robust physiological processes. However, the proposed mechanisms of droughtinduced mortality, including hydraulic failure and carbon starvation, are unresolved. A growing number of empirical studies have investigated these mechanisms, but data have not been consistently analysed across species and biomes using a standardized physiological framework. Here, we show that xylem hydraulic failure was ubiquitous across multiple tree taxa at drought-induced mortality. All species assessed had 60% or higher loss of xylem hydraulic conductivity, consistent with proposed theoretical and modelled survival thresholds. We found diverse responses in non-structural carbohydrate reserves at mortality, indicating that evidence supporting carbon starvation was not universal. Reduced non-structural carbohydrates were more common for gymnosperms than angiosperms, associated with xylem hydraulic vulnerability, and may have a role in reducing hydraulic function. Our finding that hydraulic failure at drought-induced mortality was persistent across species indicates that substantial improvement in vegetation modelling can be achieved using thresholds in hydraulic function