Climate-induced tree-mortality pulses are obscured by broad-scale and long-term greening.

Vegetation greening has been suggested to be a dominant trend over recent decades, but severe pulses of tree mortality in forests after droughts and heatwaves have also been extensively reported. These observations raise the question of to what extent the observed severe pulses of tree mortality induced by climate could affect overall vegetation greenness across spatial grains and temporal extents. To address this issue, here we analyse three satellite-based datasets of detrended growing-season normalized difference vegetation index (NDVIGS) with spatial resolutions ranging from 30 m to 8 km for 1,303 field-documented sites experiencing severe drought- or heat-induced tree-mortality events around the globe. We find that severe tree-mortality events have distinctive but localized imprints on vegetation greenness over annual timescales, which are obscured by broad-scale and long-term greening. Specifically, although anomalies in NDVIGS (ΔNDVI) are negative during tree-mortality years, this reduction diminishes at coarser spatial resolutions (that is, 250 m and 8 km). Notably, tree-mortality-induced reductions in NDVIGS (|ΔNDVI|) at 30-m resolution are negatively related to native plant species richness and forest height, whereas topographic heterogeneity is the major factor affecting ΔNDVI differences across various spatial grain sizes. Over time periods of a decade or longer, greening consistently dominates all spatial resolutions. The findings underscore the fundamental importance of spatio-temporal scales for cohesively understanding the effects of climate change on forest productivity and tree mortality under both gradual and abrupt changes.

Abundance of Dendroctonus frontalis and D. mexicanus (Coleoptera: Scolytinae) along altitudinal transects in Mexico: Implications of climatic change for forest conservation.

Bark beetle infestations have historically been primary drivers of stand thinning in Mexican pine forests. However, bark beetle impacts have become increasingly extensive and intense, apparently associated with climate change. Our objective was to describe the possible association between abundance of bark beetle flying populations and the occurrence of given value intervals of temperature, precipitation and their balance, in order to have a better comprehension of the climatic space that might trigger larger insect abundances, an issue relevant in the context of the ongoing climatic change. Here, we monitored the abundance of two of the most important bark beetle species in Mexico, Dendroctonus frontalis and D. mexicanus. We sampled 147 sites using pheromone-baited funnel traps along 24 altitudinal transects in 11 Mexican states, from northwestern Chihuahua to southeastern Chiapas, from 2015 to 2017. Through mixed model analysis, we found that the optimum Mean Annual Temperatures were 17°C-20°C for D. frontalis in low-elevation pine-oak forest, while D. mexicanus had two optimal intervals: 11-13°C and 15-18°C. Higher atmospheric Vapor Pressure Deficit (≥ 1.0) was correlated with higher D. frontalis abundances, indicating that warming-amplified drought stress intensifies trees' vulnerability to beetle attack. As temperatures and drought stress increase further with projected future climatic changes, it is likely that these Dendroctonus species will increase tree damage at higher elevations. Pine forests in Mexico are an important source of livelihood for communities inhabiting those areas, so providing tools to tackle obstacles to forest growth and health posed by changing climate is imperative.

Topographic information improves simulated patterns of post-fire conifer regeneration in the southwest United States.

The western United States is projected to experience more frequent and severe wildfires in the future due to drier and hotter climate conditions, exacerbating destructive wildfire impacts on forest ecosystems such as tree mortality and unsuccessful post-fire regeneration. While empirical studies have revealed strong relationships between topographical information and plant regeneration, ecological processes in ecosystem models have either not fully addressed topography-mediated effects on the probability of plant regeneration, or the probability is only controlled by climate-related factors, for example, water and light stresses. In this study, we incorporated seedling survival data based on a planting experiment in the footprint of the 2011 Las Conchas Fire into the Photosynthesis and EvapoTranspiration (PnET) extension of the LANDIS-II model by adding topographic and an additional climatic variable to the probability of regeneration. The modified algorithm included topographic parameters such as heat load index and ground slope and spring precipitation. We ran simulations on the Las Conchas Fire landscape for 2012-2099 using observed and projected climate data (i.e., Representative Concentration Pathway 4.5 and 8.5). Our modification significantly reduced the number of regeneration events of three common southwestern conifer tree species (piñon, ponderosa pine, and Douglas-fir), leading to decreases in aboveground biomass, regardless of climate scenario. The modified algorithm decreased regeneration at higher elevations and increased regeneration at lower elevations relative to the original algorithm. Regenerations of three species also decreased in eastern aspects. Our findings suggest that ecosystem models may overestimate post-fire regeneration events in the southwest United States. To better represent regeneration processes following wildfire, ecosystem models need refinement to better account for the range of factors that influence tree seedling establishment. This will improve model utility for projecting the combined effects of climate and wildfire on tree species distributions.

Field-based tree mortality constraint reduces estimates of model-projected forest carbon sinks.

Considerable uncertainty and debate exist in projecting the future capacity of forests to sequester atmospheric CO2. Here we estimate spatially explicit patterns of biomass loss by tree mortality (LOSS) from largely unmanaged forest plots to constrain projected (2015-2099) net primary productivity (NPP), heterotrophic respiration (HR) and net carbon sink in six dynamic global vegetation models (DGVMs) across continents. This approach relies on a strong relationship among LOSS, NPP, and HR at continental or biome scales. The DGVMs overestimated historical LOSS, particularly in tropical regions and eastern North America by as much as 5 Mg ha-1 y-1. The modeled spread of DGVM-projected NPP and HR uncertainties was substantially reduced in tropical regions after incorporating the field-based mortality constraint. The observation-constrained models show a decrease in the tropical forest carbon sink by the end of the century, particularly across South America (from 2 to 1.4 PgC y-1), and an increase in the sink in North America (from 0.8 to 1.1 PgC y-1). These results highlight the feasibility of using forest demographic data to empirically constrain forest carbon sink projections and the potential overestimation of projected tropical forest carbon sinks.

Global field observations of tree die-off reveal hotter-drought fingerprint for Earth's forests.

Earth's forests face grave challenges in the Anthropocene, including hotter droughts increasingly associated with widespread forest die-off events. But despite the vital importance of forests to global ecosystem services, their fates in a warming world remain highly uncertain. Lacking is quantitative determination of commonality in climate anomalies associated with pulses of tree mortality-from published, field-documented mortality events-required for understanding the role of extreme climate events in overall global tree die-off patterns. Here we established a geo-referenced global database documenting climate-induced mortality events spanning all tree-supporting biomes and continents, from 154 peer-reviewed studies since 1970. Our analysis quantifies a global "hotter-drought fingerprint" from these tree-mortality sites-effectively a hotter and drier climate signal for tree mortality-across 675 locations encompassing 1,303 plots. Frequency of these observed mortality-year climate conditions strongly increases nonlinearly under projected warming. Our database also provides initial footing for further community-developed, quantitative, ground-based monitoring of global tree mortality.

Climate Change Risks to Global Forest Health: Emergence of Unexpected Events of Elevated Tree Mortality Worldwide.

Recent observations of elevated tree mortality following climate extremes, like heat and drought, raise concerns about climate change risks to global forest health. We currently lack both sufficient data and understanding to identify whether these observations represent a global trend toward increasing tree mortality. Here, we document events of sudden and unexpected elevated tree mortality following heat and drought events in ecosystems that previously were considered tolerant or not at risk of exposure. These events underscore the fact that climate change may affect forests with unexpected force in the future. We use the events as examples to highlight current difficulties and challenges for realistically predicting such tree mortality events and the uncertainties about future forest condition. Advances in remote sensing technology and greater availably of high-resolution data, from both field assessments and satellites, are needed to improve both understanding and prediction of forest responses to future climate change.

Nature-based framework for sustainable afforestation in global drylands under changing climate.

Drylands cover more than 40% of Earth's land surface and occur at the margin of forest distributions due to the limited availability of water for tree growth. Recent elevated temperature and low precipitation have driven greater forest declines and pulses of tree mortality on dryland sites compared to humid sites, particularly in temperate Eurasia and North America. Afforestation of dryland areas has been widely implemented and is expected to increase in many drylands globally to enhance carbon sequestration and benefits to the human environment, but the interplay of sometimes conflicting afforestation outcomes has not been formally evaluated yet. Most previous studies point to conflicts between additional forest area and water consumption, in particular water yield and soil conservation/desalinization in drylands, but were generally confined to local and regional scales. Our global synthesis demonstrates that additional tree cover can amplify water consumption through a nonlinear increase in evapotranspiration-depending on tree species, age, and structure-which will be further intensified by future climate change. In this review we identify substantial knowledge gaps in addressing the dryland afforestation dilemma, where there are trade-offs with planted forests between increased availability of some resources and benefits to human habitats versus the depletion of other resources that are required for sustainable development of drylands. Here we propose a method of addressing comprehensive vegetation carrying capacity, based on regulating the distribution and structure of forest plantations to better deal with these trade-offs in forest multifunctionality. We also recommend new priority research topics for dryland afforestation, including: responses and feedbacks of dryland forests to climate change; shifts in the ratio of ecosystem ET to tree cover; assessing the role of scale of afforestation in influencing the trade-offs of dryland afforestation; and comprehensive modeling of the multifunctionality of dryland forests, including both ecophysiological and socioeconomic aspects, under a changing climate.

Joint effects of climate, tree size, and year on annual tree growth derived from tree-ring records of ten globally distributed forests.

Tree rings provide an invaluable long-term record for understanding how climate and other drivers shape tree growth and forest productivity. However, conventional tree-ring analysis methods were not designed to simultaneously test effects of climate, tree size, and other drivers on individual growth. This has limited the potential to test ecologically relevant hypotheses on tree growth sensitivity to environmental drivers and their interactions with tree size. Here, we develop and apply a new method to simultaneously model nonlinear effects of primary climate drivers, reconstructed tree diameter at breast height (DBH), and calendar year in generalized least squares models that account for the temporal autocorrelation inherent to each individual tree's growth. We analyze data from 3811 trees representing 40 species at 10 globally distributed sites, showing that precipitation, temperature, DBH, and calendar year have additively, and often interactively, influenced annual growth over the past 120 years. Growth responses were predominantly positive to precipitation (usually over ≥3-month seasonal windows) and negative to temperature (usually maximum temperature, over ≤3-month seasonal windows), with concave-down responses in 63% of relationships. Climate sensitivity commonly varied with DBH (45% of cases tested), with larger trees usually more sensitive. Trends in ring width at small DBH were linked to the light environment under which trees established, but basal area or biomass increments consistently reached maxima at intermediate DBH. Accounting for climate and DBH, growth rate declined over time for 92% of species in secondary or disturbed stands, whereas growth trends were mixed in older forests. These trends were largely attributable to stand dynamics as cohorts and stands age, which remain challenging to disentangle from global change drivers. By providing a parsimonious approach for characterizing multiple interacting drivers of tree growth, our method reveals a more complete picture of the factors influencing growth than has previously been possible.

Mortality predispositions of conifers across western USA.

Conifer mortality rates are increasing in western North America, but the physiological mechanisms underlying this trend are not well understood. We examined tree-ring-based radial growth along with stable carbon (C) and oxygen (O) isotope composition (δ13 C and δ18 O, respectively) of dying and surviving conifers at eight old-growth forest sites across a strong moisture gradient in the western USA to retrospectively investigate mortality predispositions. Compared with surviving trees, lower growth of dying trees was detected at least one decade before mortality at seven of the eight sites. Intrinsic water-use efficiency increased over time in both dying and surviving trees, with a weaker increase in dying trees at five of the eight sites. C starvation was a strong correlate of conifer mortality based on a conceptual model incorporating growth, δ13 C, and δ18 O. However, this approach does not capture processes that occur in the final months of survival. Ultimately, C starvation may lead to increased mortality vulnerability, but hydraulic failure or biotic attack may dominate the process during the end stages of mortality in these conifers.

Pervasive shifts in forest dynamics in a changing world.

Forest dynamics arise from the interplay of environmental drivers and disturbances with the demographic processes of recruitment, growth, and mortality, subsequently driving biomass and species composition. However, forest disturbances and subsequent recovery are shifting with global changes in climate and land use, altering these dynamics. Changes in environmental drivers, land use, and disturbance regimes are forcing forests toward younger, shorter stands. Rising carbon dioxide, acclimation, adaptation, and migration can influence these impacts. Recent developments in Earth system models support increasingly realistic simulations of vegetation dynamics. In parallel, emerging remote sensing datasets promise qualitatively new and more abundant data on the underlying processes and consequences for vegetation structure. When combined, these advances hold promise for improving the scientific understanding of changes in vegetation demographics and disturbances.

Exposures to temperature beyond threshold disproportionately reduce vegetation growth in the northern hemisphere.

In recent decades, terrestrial vegetation in the northern hemisphere (NH) has been exposed to warming and more extremely high temperatures. However, the consequences of these changes for terrestrial vegetation growth remain poorly quantified and understood. By examining a satellite-based vegetation index, tree-ring measurements and land-surface model simulations, we discovered a consistent convex pattern in the responses of vegetation growth to temperature exposure (TE) for forest, shrub and grass in both the temperate (30°-50° N) and boreal (50°-70° N) NH during the period of 1982-2012. The response of vegetation growth to TE for the three vegetation types in both the temperate and boreal NH increased convergently with increasing temperature, until vegetation type-dependent temperature thresholds were reached. A TE beyond these temperature thresholds resulted in disproportionately weak positive or even strong negative responses. Vegetation growth in the boreal NH was more vulnerable to extremely high-temperature events than vegetation growth in the temporal NH. The non-linear responses discovered here provide new insights into the dynamics of northern terrestrial ecosystems in a warmer world.

Research frontiers for improving our understanding of drought-induced tree and forest mortality.

Accumulating evidence highlights increased mortality risks for trees during severe drought, particularly under warmer temperatures and increasing vapour pressure deficit (VPD). Resulting forest die-off events have severe consequences for ecosystem services, biophysical and biogeochemical land-atmosphere processes. Despite advances in monitoring, modelling and experimental studies of the causes and consequences of tree death from individual tree to ecosystem and global scale, a general mechanistic understanding and realistic predictions of drought mortality under future climate conditions are still lacking. We update a global tree mortality map and present a roadmap to a more holistic understanding of forest mortality across scales. We highlight priority research frontiers that promote: (1) new avenues for research on key tree ecophysiological responses to drought; (2) scaling from the tree/plot level to the ecosystem and region; (3) improvements of mortality risk predictions based on both empirical and mechanistic insights; and (4) a global monitoring network of forest mortality. In light of recent and anticipated large forest die-off events such a research agenda is timely and needed to achieve scientific understanding for realistic predictions of drought-induced tree mortality. The implementation of a sustainable network will require support by stakeholders and political authorities at the international level.

Drivers and mechanisms of tree mortality in moist tropical forests.

Tree mortality rates appear to be increasing in moist tropical forests (MTFs) with significant carbon cycle consequences. Here, we review the state of knowledge regarding MTF tree mortality, create a conceptual framework with testable hypotheses regarding the drivers, mechanisms and interactions that may underlie increasing MTF mortality rates, and identify the next steps for improved understanding and reduced prediction. Increasing mortality rates are associated with rising temperature and vapor pressure deficit, liana abundance, drought, wind events, fire and, possibly, CO2 fertilization-induced increases in stand thinning or acceleration of trees reaching larger, more vulnerable heights. The majority of these mortality drivers may kill trees in part through carbon starvation and hydraulic failure. The relative importance of each driver is unknown. High species diversity may buffer MTFs against large-scale mortality events, but recent and expected trends in mortality drivers give reason for concern regarding increasing mortality within MTFs. Models of tropical tree mortality are advancing the representation of hydraulics, carbon and demography, but require more empirical knowledge regarding the most common drivers and their subsequent mechanisms. We outline critical datasets and model developments required to test hypotheses regarding the underlying causes of increasing MTF mortality rates, and improve prediction of future mortality under climate change.

A multi-species synthesis of physiological mechanisms in drought-induced tree mortality.

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 drought-induced 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.

Tree mortality across biomes is promoted by drought intensity, lower wood density and higher specific leaf area.

Drought events are increasing globally, and reports of consequent forest mortality are widespread. However, due to a lack of a quantitative global synthesis, it is still not clear whether drought-induced mortality rates differ among global biomes and whether functional traits influence the risk of drought-induced mortality. To address these uncertainties, we performed a global meta-analysis of 58 studies of drought-induced forest mortality. Mortality rates were modelled as a function of drought, temperature, biomes, phylogenetic and functional groups and functional traits. We identified a consistent global-scale response, where mortality increased with drought severity [log mortality (trees trees-1  year-1 ) increased 0.46 (95% CI = 0.2-0.7) with one SPEI unit drought intensity]. We found no significant differences in the magnitude of the response depending on forest biomes or between angiosperms and gymnosperms or evergreen and deciduous tree species. Functional traits explained some of the variation in drought responses between species (i.e. increased from 30 to 37% when wood density and specific leaf area were included). Tree species with denser wood and lower specific leaf area showed lower mortality responses. Our results illustrate the value of functional traits for understanding patterns of drought-induced tree mortality and suggest that mortality could become increasingly widespread in the future.

Structural overshoot of tree growth with climate variability and the global spectrum of drought-induced forest dieback.

Ongoing climate change poses significant threats to plant function and distribution. Increased temperatures and altered precipitation regimes amplify drought frequency and intensity, elevating plant stress and mortality. Large-scale forest mortality events will have far-reaching impacts on carbon and hydrological cycling, biodiversity, and ecosystem services. However, biogeographical theory and global vegetation models poorly represent recent forest die-off patterns. Furthermore, as trees are sessile and long-lived, their responses to climate extremes are substantially dependent on historical factors. We show that periods of favourable climatic and management conditions that facilitate abundant tree growth can lead to structural overshoot of aboveground tree biomass due to a subsequent temporal mismatch between water demand and availability. When environmental favourability declines, increases in water and temperature stress that are protracted, rapid, or both, drive a gradient of tree structural responses that can modify forest self-thinning relationships. Responses ranging from premature leaf senescence and partial canopy dieback to whole-tree mortality reduce canopy leaf area during the stress period and for a lagged recovery window thereafter. Such temporal mismatches of water requirements from availability can occur at local to regional scales throughout a species geographical range. As climate change projections predict large future fluctuations in both wet and dry conditions, we expect forests to become increasingly structurally mismatched to water availability and thus overbuilt during more stressful episodes. By accounting for the historical context of biomass development, our approach can explain previously problematic aspects of large-scale forest mortality, such as why it can occur throughout the range of a species and yet still be locally highly variable, and why some events seem readily attributable to an ongoing drought while others do not. This refined understanding can facilitate better projections of structural overshoot responses, enabling improved prediction of changes in forest distribution and function from regional to global scales.

Multiscale perspectives of fire, climate and humans in western North America and the Jemez Mountains, USA.

Interannual climate variations have been important drivers of wildfire occurrence in ponderosa pine forests across western North America for at least 400 years, but at finer scales of mountain ranges and landscapes human land uses sometimes over-rode climate influences. We reconstruct and analyse effects of high human population densities in forests of the Jemez Mountains, New Mexico from ca 1300 CE to Present. Prior to the 1680 Pueblo Revolt, human land uses reduced the occurrence of widespread fires while simultaneously adding more ignitions resulting in many small-extent fires. During the 18th and 19th centuries, wet/dry oscillations and their effects on fuels dynamics controlled widespread fire occurrence. In the late 19th century, intensive livestock grazing disrupted fuels continuity and fire spread and then active fire suppression maintained the absence of widespread surface fires during most of the 20th century. The abundance and continuity of fuels is the most important controlling variable in fire regimes of these semi-arid forests. Reduction of widespread fires owing to reduction of fuel continuity emerges as a hallmark of extensive human impacts on past forests and fire regimes.This article is part of the themed issue 'The interaction of fire and mankind'.

The impacts of increasing drought on forest dynamics, structure, and biodiversity in the United States.

We synthesize insights from current understanding of drought impacts at stand-to-biogeographic scales, including management options, and we identify challenges to be addressed with new research. Large stand-level shifts underway in western forests already are showing the importance of interactions involving drought, insects, and fire. Diebacks, changes in composition and structure, and shifting range limits are widely observed. In the eastern US, the effects of increasing drought are becoming better understood at the level of individual trees, but this knowledge cannot yet be confidently translated to predictions of changing structure and diversity of forest stands. While eastern forests have not experienced the types of changes seen in western forests in recent decades, they too are vulnerable to drought and could experience significant changes with increased severity, frequency, or duration in drought. Throughout the continental United States, the combination of projected large climate-induced shifts in suitable habitat from modeling studies and limited potential for the rapid migration of tree populations suggests that changing tree and forest biogeography could substantially lag habitat shifts already underway. Forest management practices can partially ameliorate drought impacts through reductions in stand density, selection of drought-tolerant species and genotypes, artificial regeneration, and the development of multistructured stands. However, silvicultural treatments also could exacerbate drought impacts unless implemented with careful attention to site and stand characteristics. Gaps in our understanding should motivate new research on the effects of interactions involving climate and other species at the stand scale and how interactions and multiple responses are represented in models. This assessment indicates that, without a stronger empirical basis for drought impacts at the stand scale, more complex models may provide limited guidance.

Tree mortality from drought, insects, and their interactions in a changing climate.

Climate change is expected to drive increased tree mortality through drought, heat stress, and insect attacks, with manifold impacts on forest ecosystems. Yet, climate-induced tree mortality and biotic disturbance agents are largely absent from process-based ecosystem models. Using data sets from the western USA and associated studies, we present a framework for determining the relative contribution of drought stress, insect attack, and their interactions, which is critical for modeling mortality in future climates. We outline a simple approach that identifies the mechanisms associated with two guilds of insects - bark beetles and defoliators - which are responsible for substantial tree mortality. We then discuss cross-biome patterns of insect-driven tree mortality and draw upon available evidence contrasting the prevalence of insect outbreaks in temperate and tropical regions. We conclude with an overview of tools and promising avenues to address major challenges. Ultimately, a multitrophic approach that captures tree physiology, insect populations, and tree-insect interactions will better inform projections of forest ecosystem responses to climate change.