North American tree migration paced by climate in the West, lagging in the East.

Tree fecundity and recruitment have not yet been quantified at scales needed to anticipate biogeographic shifts in response to climate change. By separating their responses, this study shows coherence across species and communities, offering the strongest support to date that migration is in progress with regional limitations on rates. The southeastern continent emerges as a fecundity hotspot, but it is situated south of population centers where high seed production could contribute to poleward population spread. By contrast, seedling success is highest in the West and North, serving to partially offset limited seed production near poleward frontiers. The evidence of fecundity and recruitment control on tree migration can inform conservation planning for the expected long-term disequilibrium between climate and forest distribution.

Crowding, climate, and the case for social distancing among trees.

In an emerging era of megadisturbance, bolstering forest resilience to wildfire, insects, and drought has become a central objective in many western forests. Climate has received considerable attention as a driver of these disturbances, but few studies have examined the complexities of climate-vegetation-disturbance interactions. Current strategies for creating resilient forests often rely on retrospective approaches, seeking to impart resilience by restoring historical conditions to contemporary landscapes, but historical conditions are becoming increasingly unattainable amidst modern bioclimatic conditions. What becomes an appropriate benchmark for resilience when we have novel forests, rapidly changing climate, and unprecedented disturbance regimes? We combined two longitudinal datasets-each representing some of the most comprehensive spatially explicit, annual tree mortality data in existence-in a post-hoc factorial design to examine the nonlinear relationships between fire, climate, forest spatial structure, and bark beetles. We found that while prefire drought elevated mortality risk, advantageous local neighborhoods could offset these effects. Surprisingly, mortality risk (Pm ) was higher in crowded local neighborhoods that burned in wet years (Pm  = 42%) compared with sparse neighborhoods that burned during drought (Pm  = 30%). Risk of beetle attack was also increased by drought, but lower conspecific crowding impeded the otherwise positive interaction between fire and beetle attack. Antecedent fire increased drought-related mortality over short timespans (<7 years) but reduced mortality over longer intervals. These results clarify interacting disturbance dynamics and provide a mechanistic underpinning for forest restoration strategies. Importantly, they demonstrate the potential for managed fire and silvicultural strategies to offset climate effects and bolster resilience to fire, beetles, and drought.

Mapping the vulnerability of giant sequoias after extreme drought in California using remote sensing.

Between 2012 and 2016, California suffered one of the most severe droughts on record. During this period Sequoiadendron giganteum (giant sequoias) in the Sequoia and Kings Canyon National Parks (SEKI), California, USA experienced canopy water content (CWC) loss, unprecedented foliage senescence, and, in a few cases, death. We present an assessment of the vulnerability of giant sequoia populations to droughts that is currently lacking and needed for management. We used a temporal trend of remotely sensed CWC obtained between 2015 and 2017, and recently georeferenced giant sequoia crowns to quantify the vulnerability of 7,408 individuals in 10 groves in the northern portion of SEKI. CWC is sensitive to changes in liquid water in tree canopies; therefore, it is a useful metric for quantifying the response of sequoia trees to drought. Temporal trends indicated that 9% of giant sequoias had a significant decline or consistently low CWC, suggesting these trees were likely operating at low photosynthetic capacity and potentially at high risk to drought stress. We also found that 20% of the giant sequoias had an increase or consistently high level of CWC, indicating these trees were at low risk to drought stress. These vulnerability categories were used in a random forest model with a combination of topographic, fire-related, and climate variables to generate high-resolution vulnerability risk maps. These maps show that higher risk is associated with lower elevation and higher climate water deficit. We also found that sequoias at higher elevations but located near meadows had higher vulnerability risk. These results and the vulnerability maps can identify vulnerable sequoias that may be difficult to save or locations of refugia to be protected, and thus may aid forest managers in preparation for future droughts.

Why is Tree Drought Mortality so Hard to Predict?

Widespread tree mortality following droughts has emerged as an environmentally and economically devastating 'ecological surprise'. It is well established that tree physiology is important in understanding drought-driven mortality; however, the accuracy of predictions based on physiology alone has been limited. We propose that complicating factors at two levels stymie predictions of drought-driven mortality: (i) organismal-level physiological and site factors that obscure understanding of drought exposure and vulnerability and (ii) community-level ecological interactions, particularly with biotic agents whose effects on tree mortality may reverse expectations based on stress physiology. We conclude with a path forward that emphasizes the need for an integrative approach to stress physiology and biotic agent dynamics when assessing forest risk to drought-driven morality in a changing climate.

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.

Effects of postfire climate and seed availability on postfire conifer regeneration.

Large, severe fires are becoming more frequent in many forest types across the western United States and have resulted in tree mortality across tens of thousands of hectares. Conifer regeneration in these areas is limited because seeds must travel long distances to reach the interior of large burned patches and establishment is jeopardized by increasingly hot and dry conditions. To better inform postfire management in low elevation forests of California, USA, we collected 5-yr postfire recovery data from 1,234 study plots in 19 wildfires that burned from 2004-2012 and 18 yrs of seed production data from 216 seed fall traps (1999-2017). We used these data in conjunction with spatially extensive climate, topography, forest composition, and burn severity surfaces to construct taxon-specific, spatially explicit models of conifer regeneration that incorporate climate conditions and seed availability during postfire recovery windows. We found that after accounting for other predictors both postfire and historical precipitation were strong predictors of regeneration, suggesting that both direct effects of postfire moisture conditions and biological inertia from historical climate may play a role in regeneration. Alternatively, postfire regeneration may simply be driven by postfire climate and apparent relationships with historical climate could be spurious. The estimated sensitivity of regeneration to postfire seed availability was strongest in firs and all conifers combined and weaker in pines. Seed production exhibited high temporal variability with seed production varying by over two orders of magnitude among years. Our models indicate that during droughts postfire conifer regeneration declines most substantially in low-to-moderate elevation forests. These findings enhance our mechanistic understanding of forecasted and historically documented shifts in the distribution of trees.

Pre-fire drought and competition mediate post-fire conifer mortality in western U.S. National Parks.

Tree mortality is an important outcome of many forest fires. Extensive tree injuries from fire may lead directly to mortality, but environmental and biological stressors may also contribute to tree death. However, there is little evidence showing how the combined effects of two common stressors, drought and competition, influence post-fire mortality. Geographically broad observations of three common western coniferous trees subjected to prescribed fire showed the likelihood of post-fire mortality was related to intermediate-term (10 yr) pre-fire average radial growth, an important component of tree vigor. Path analysis showed that indices of competition and drought stress prior to fire can be described in terms of joint effects on growth, indirectly affecting post-fire mortality. Our results suggest that the conditions that govern the relationship between growth and mortality in unburned stands may also apply to post-fire environments. Thus, biotic and abiotic changes that affect growth negatively (e.g., drought stress) or positively (e.g., growth releases following thinning treatments) prior to fire may influence expressed fire severity, independent of fire intensity (e.g., heat flux, residence time). These relationships suggest that tree mortality may increase under stressful climatic or stand conditions even if fire behavior remains constant.

What mediates tree mortality during drought in the southern Sierra Nevada?

Severe drought has the potential to cause selective mortality within a forest, thereby inducing shifts in forest species composition. The southern Sierra Nevada foothills and mountains of California have experienced extensive forest dieback due to drought stress and insect outbreak. We used high-fidelity imaging spectroscopy (HiFIS) and light detection and ranging (LiDAR) from the Carnegie Airborne Observatory (CAO) to estimate the effect of forest dieback on species composition in response to drought stress in Sequoia National Park. Our aims were (1) to quantify site-specific conditions that mediate tree mortality along an elevation gradient in the southern Sierra Nevada Mountains, (2) to assess where mortality events have a greater probability of occurring, and (3) to estimate which tree species have a greater likelihood of mortality along the elevation gradient. A series of statistical models were generated to classify species composition and identify tree mortality, and the influences of different environmental factors were spatially quantified and analyzed to assess where mortality events have a greater likelihood of occurring. A higher probability of mortality was observed in the lower portion of the elevation gradient, on southwest- and west-facing slopes, in areas with shallow soils, on shallower slopes, and at greater distances from water. All of these factors are related to site water balance throughout the landscape. Our results also suggest that mortality is species-specific along the elevation gradient, mainly affecting Pinus ponderosa and Pinus lambertiana at lower elevations. Selective mortality within the forest may drive long-term shifts in community composition along the elevation gradient.

Why do trees die? Characterizing the drivers of background tree mortality.

The drivers of background tree mortality rates-the typical low rates of tree mortality found in forests in the absence of acute stresses like drought-are central to our understanding of forest dynamics, the effects of ongoing environmental changes on forests, and the causes and consequences of geographical gradients in the nature and strength of biotic interactions. To shed light on factors contributing to background tree mortality, we analyzed detailed pathological data from 200,668 tree-years of observation and 3,729 individual tree deaths, recorded over a 13-yr period in a network of old-growth forest plots in California's Sierra Nevada mountain range. We found that: (1) Biotic mortality factors (mostly insects and pathogens) dominated (58%), particularly in larger trees (86%). Bark beetles were the most prevalent (40%), even though there were no outbreaks during the study period; in contrast, the contribution of defoliators was negligible. (2) Relative occurrences of broad classes of mortality factors (biotic, 58%; suppression, 51%; and mechanical, 25%) are similar among tree taxa, but may vary with tree size and growth rate. (3) We found little evidence of distinct groups of mortality factors that predictably occur together on trees. Our results have at least three sets of implications. First, rather than being driven by abiotic factors such as lightning or windstorms, the "ambient" or "random" background mortality that many forest models presume to be independent of tree growth rate is instead dominated by biotic agents of tree mortality, with potentially critical implications for forecasting future mortality. Mechanistic models of background mortality, even for healthy, rapidly growing trees, must therefore include the insects and pathogens that kill trees. Second, the biotic agents of tree mortality, instead of occurring in a few predictable combinations, may generally act opportunistically and with a relatively large degree of independence from one another. Finally, beyond the current emphasis on folivory and leaf defenses, studies of broad-scale gradients in the nature and strength of biotic interactions should also include biotic attacks on, and defenses of, tree stems and roots.

Temperate forest health in an era of emerging megadisturbance.

Although disturbances such as fire and native insects can contribute to natural dynamics of forest health, exceptional droughts, directly and in combination with other disturbance factors, are pushing some temperate forests beyond thresholds of sustainability. Interactions from increasing temperatures, drought, native insects and pathogens, and uncharacteristically severe wildfire are resulting in forest mortality beyond the levels of 20th-century experience. Additional anthropogenic stressors, such as atmospheric pollution and invasive species, further weaken trees in some regions. Although continuing climate change will likely drive many areas of temperate forest toward large-scale transformations, management actions can help ease transitions and minimize losses of socially valued ecosystem services.

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.

Climatic correlates of tree mortality in water- and energy-limited forests.

Recent increases in tree mortality rates across the western USA are correlated with increasing temperatures, but mechanisms remain unresolved. Specifically, increasing mortality could predominantly be a consequence of temperature-induced increases in either (1) drought stress, or (2) the effectiveness of tree-killing insects and pathogens. Using long-term data from California's Sierra Nevada mountain range, we found that in water-limited (low-elevation) forests mortality was unambiguously best modeled by climatic water deficit, consistent with the first mechanism. In energy-limited (high-elevation) forests deficit models were only equivocally better than temperature models, suggesting that the second mechanism is increasingly important in these forests. We could not distinguish between models predicting mortality using absolute versus relative changes in water deficit, and these two model types led to different forecasts of mortality vulnerability under future climate scenarios. Our results provide evidence for differing climatic controls of tree mortality in water- and energy-limited forests, while highlighting the need for an improved understanding of tree mortality processes.

Comment on "Changes in climatic water balance drive downhill shifts in plant species' optimum elevations".

Crimmins et al. (Reports, 21 January 2011, p. 324) attributed an apparent downward elevational shift of California plant species to a precipitation-induced decline in climatic water deficit. We show that the authors miscalculated deficit, that the apparent decline in species' elevations is likely a consequence of geographic biases, and that unlike temperature changes, precipitation changes should not be expected to cause coordinated directional shifts in species' elevations.

Widespread increase of tree mortality rates in the western United States.

Persistent changes in tree mortality rates can alter forest structure, composition, and ecosystem services such as carbon sequestration. Our analyses of longitudinal data from unmanaged old forests in the western United States showed that background (noncatastrophic) mortality rates have increased rapidly in recent decades, with doubling periods ranging from 17 to 29 years among regions. Increases were also pervasive across elevations, tree sizes, dominant genera, and past fire histories. Forest density and basal area declined slightly, which suggests that increasing mortality was not caused by endogenous increases in competition. Because mortality increased in small trees, the overall increase in mortality rates cannot be attributed solely to aging of large trees. Regional warming and consequent increases in water deficits are likely contributors to the increases in tree mortality rates.

Spatial elements of mortality risk in old-growth forests.

For many species of long-lived organisms, such as trees, survival appears to be the most critical vital rate affecting population persistence. However, methods commonly used to quantify tree death, such as relating tree mortality risk solely to diameter growth, almost certainly do not account for important spatial processes. Our goal in this study was to detect and, if present, to quantify the relevance of such processes. For this purpose, we examined purely spatial aspects of mortality for four species, Abies concolor, Abies magnifica, Calocedrus decurrens, and Pinus lambertiana, in an old-growth conifer forest in the Sierra Nevada of California, USA. The analysis was performed using data from nine fully mapped long-term monitoring plots. In three cases, the results unequivocally supported the inclusion of spatial information in models used to predict mortality. For Abies concolor, our results suggested that growth rate may not always adequately capture increased mortality risk due to competition. We also found evidence of a facilitative effect for this species, with mortality risk decreasing with proximity to conspecific neighbors. For Pinus lambertiana, mortality risk increased with density of conspecific neighbors, in keeping with a mechanism of increased pathogen or insect pressure (i.e., a Janzen-Connell type effect). Finally, we found that models estimating risk of being crushed were strongly improved by the inclusion of a simple index of spatial proximity. Not only did spatial indices improve models, those improvements were relevant for mortality prediction. For P. lambertiana, spatial factors were important for estimation of mortality risk regardless of growth rate. For A. concolor, although most of the population fell within spatial conditions in which mortality risk was well described by growth, trees that died occurred outside those conditions in a disproportionate fashion. Furthermore, as stands of A. concolor become increasingly dense, such spatial factors are likely to become increasingly important. In general, models that fail to account for spatial pattern are at risk of failure as conditions change.

Apparent climatically induced increase of tree mortality rates in a temperate forest.

We provide a first detailed analysis of long-term, annual-resolution demographic trends in a temperate forest. After tracking the fates of 21,338 trees in a network of old-growth forest plots in the Sierra Nevada of California, we found that mortality rate, but not the recruitment rate, increased significantly over the 22 years of measurement (1983-2004). Mortality rates increased in both of two dominant taxonomic groups (Abies and Pinus) and in different forest types (different elevational zones). The increase in overall mortality rate resulted from an increase in tree deaths attributed to stress and biotic causes, and coincided with a temperature-driven increase in an index of drought. Our findings suggest that these forests (and by implication, other water-limited forests) may be sensitive to temperature-driven drought stress, and may be poised for die-back if future climates continue to feature rising temperatures without compensating increases in precipitation.

Climate change and forests of the future: managing in the face of uncertainty.

We offer a conceptual framework for managing forested ecosystems under an assumption that future environments will be different from present but that we cannot be certain about the specifics of change. We encourage flexible approaches that promote reversible and incremental steps, and that favor ongoing learning and capacity to modify direction as situations change. We suggest that no single solution fits all future challenges, especially in the context of changing climates, and that the best strategy is to mix different approaches for different situations. Resources managers will be challenged to integrate adaptation strategies (actions that help ecosystems accommodate changes adaptively) and mitigation strategies (actions that enable ecosystems to reduce anthropogenic influences on global climate) into overall plans. Adaptive strategies include resistance options (forestall impacts and protect highly valued resources), resilience options (improve the capacity of ecosystems to return to desired conditions after disturbance), and response options (facilitate transition of ecosystems from current to new conditions). Mitigation strategies include options to sequester carbon and reduce overall greenhouse gas emissions. Priority-setting approaches (e.g., triage), appropriate for rapidly changing conditions and for situations where needs are greater than available capacity to respond, will become increasingly important in the future.

Forest turnover rates follow global and regional patterns of productivity.

Using a global database, we found that forest turnover rates (the average of tree mortality and recruitment rates) parallel broad-scale patterns of net primary productivity. First, forest turnover was higher in tropical than in temperate forests. Second, as recently demonstrated by others, Amazonian forest turnover was higher on fertile than infertile soils. Third, within temperate latitudes, turnover was highest in angiosperm forests, intermediate in mixed forests, and lowest in gymnosperm forests. Finally, within a single forest physiognomic type, turnover declined sharply with elevation (hence with temperature). These patterns of turnover in populations of trees are broadly similar to the patterns of turnover in populations of plant organs (leaves and roots) found in other studies. Our findings suggest a link between forest mass balance and the population dynamics of trees, and have implications for understanding and predicting the effects of environmental changes on forest structure and terrestrial carbon dynamics.