Growth, drought response, and climate-associated genomic structure in whitebark pine in the Sierra Nevada of California.

Whitebark pine (Pinus albicaulis Engelm.) has experienced rapid population declines and is listed as threatened under the Endangered Species Act in the United States. Whitebark pine in the Sierra Nevada of California represents the southernmost end of the species' distribution and, like other portions of its range, faces threats from an introduced pathogen, native bark beetles, and a rapidly warming climate. Beyond these chronic stressors, there is also concern about how this species will respond to acute stressors, such as drought. We present patterns of stem growth from 766 large (average diameter at breast height >25 cm), disease-free whitebark pine across the Sierra Nevada before and during a recent period of drought. We contextualize growth patterns using population genomic diversity and structure from a subset of 327 trees. Sampled whitebark pine generally had positive to neutral stem growth trends from 1970 to 2011, which was positively correlated with minimum temperature and precipitation. Indices of stem growth during drought years (2012 to 2015) relative to a predrought interval were mostly positive to neutral at our sampled sites. Individual tree growth response phenotypes appeared to be linked to genotypic variation in climate-associated loci, suggesting that some genotypes can take better advantage of local climatic conditions than others. We speculate that reduced snowpack during the 2012 to 2015 drought years may have lengthened the growing season while retaining sufficient moisture to maintain growth at most study sites. Growth responses may differ under future warming, however, particularly if drought severity increases and modifies interactions with pests and pathogens.

Predictive accuracy of post-fire conifer death declines over time in models based on crown and bole injury.

A key uncertainty of empirical models of post-fire tree mortality is understanding the drivers of elevated post-fire mortality several years following fire, known as delayed mortality. Delayed mortality can represent a substantial fraction of mortality, particularly for large trees that are a conservation focus in western US coniferous forests. Current post-fire tree mortality models have undergone limited evaluation of how injury level and time since fire interact to influence model accuracy and predictor variable importance. Less severe injuries potentially serve as an indicator for vulnerability to additional stressors such as bark beetle attack or moisture stress. We used a collection of 164,293 individual tree records to examine post-fire tree mortality in eight western USA conifers: Abies concolor, Abies grandis, Calocedrus decurrens, Larix occidentalis, Pinus contorta, Pinus lambertiana, Pinus ponderosa, and Pseudotsuga menziesii. We evaluated the importance of fire injury predictors on discriminating between surviving trees versus immediate and delayed post-fire mortality. We fit balanced random forest models for each species using cumulative tree mortality from 1 to 5-years post-fire. We compared these results to multi-class random forest models using first-year mortality, 2-5-year mortality, and survival 5-years post-fire as a response variable. Crown volume scorched, diameter at breast height, and relative bark char height, were used as predictor variables. The cumulative mortality models all predicted trees that died within 1-year of fire with high accuracy but failed to predict 2-5-year mortality. The multi-class models were an improvement but had lower accuracy for predicting 2-5-year mortality. Multi-class model accuracies ranged from 85% to 95% across all species for predicting 1-year post-fire mortality, 42%-71% for predicting 2-5-year mortality, and 64%-85% for predicting trees that lived past 5-years. Our study highlights the differences in tree species tolerance to fire injury and suggests that including second-order predictors such as beetle attack or climatic water stress before and after fire will be critical to improve accuracy and better understand the mechanisms and patterns of fire-caused tree death. Random forest models have potential for management applications such as post-fire harvesting and simulating future stand dynamics.

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.

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.

The Fire and Tree Mortality Database, for empirical modeling of individual tree mortality after fire.

Wildland fires have a multitude of ecological effects in forests, woodlands, and savannas across the globe. A major focus of past research has been on tree mortality from fire, as trees provide a vast range of biological services. We assembled a database of individual-tree records from prescribed fires and wildfires in the United States. The Fire and Tree Mortality (FTM) database includes records from 164,293 individual trees with records of fire injury (crown scorch, bole char, etc.), tree diameter, and either mortality or top-kill up to ten years post-fire. Data span 142 species and 62 genera, from 409 fires occurring from 1981-2016. Additional variables such as insect attack are included when available. The FTM database can be used to evaluate individual fire-caused mortality models for pre-fire planning and post-fire decision support, to develop improved models, and to explore general patterns of individual fire-induced tree death. The database can also be used to identify knowledge gaps that could be addressed in future research.

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.

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.

Climatic stress increases forest fire severity across the western United States.

Pervasive warming can lead to chronic stress on forest trees, which may contribute to mortality resulting from fire-caused injuries. Longitudinal analyses of forest plots from across the western US show that high pre-fire climatic water deficit was related to increased post-fire tree mortality probabilities. This relationship between climate and fire was present after accounting for fire defences and injuries, and appeared to influence the effects of crown and stem injuries. Climate and fire interactions did not vary substantially across geographical regions, major genera and tree sizes. Our findings support recent physiological evidence showing that both drought and heating from fire can impair xylem conductivity. Warming trends have been linked to increasing probabilities of severe fire weather and fire spread; our results suggest that warming may also increase forest fire severity (the number of trees killed) independent of fire intensity (the amount of heat released during a fire).

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.