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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.

The dead foliage of scorched crowns is one of the most conspicuous signatures of wildland fires. Globally, crown scorch from fires in savannas, woodlands and forests causes tree stress and death across diverse taxa. The term crown scorch, however, is inconsistently and ambiguously defined in the literature, causing confusion and conflicting interpretation of results. Furthermore, the underlying mechanisms causing foliage death from fire are poorly understood. The consequences of crown scorch – alterations in physiological, biogeochemical and ecological processes and ecosystem recovery pathways – remain largely unexamined. Most research on the topic assumes the mechanism of leaf and bud death is exposure to lethal air temperatures, with few direct measurements of lethal heating thresholds. Notable information gaps include how energy transfer injures and kills leaves and buds, how nutrients, carbohydrates, and hormones respond, and what physiological consequences lead to mortality. We clarify definitions to encourage use of unified terminology for foliage and bud necrosis resulting from fire. We review the current understanding of the physical mechanisms driving foliar injury, discuss the physiological responses, and explore novel ecological consequences of crown injury from fire. From these elements, we propose research needs for the increasingly interdisciplinary study of fire effects.

Drought-induced tree mortality results from an interaction of several mechanisms. Plant water and carbon relations are interdependent and assessments of their individual contributions are difficult. Because drought always affects both plant hydration and carbon assimilation, it is challenging to disentangle their concomitant effects on carbon balance and carbon translocation. Here, we report results of a manipulation experiment specifically designed to separate drought effects on carbon and water relations from those on carbon translocation. In a glasshouse experiment, we manipulated the carbon balance of Norway spruce saplings exposed to either drought or carbon starvation (CO2 withdrawal), or both treatments, and compared the dynamics of carbon exchange, allocation and storage in different tissues. Drought killed trees much faster than did carbon starvation. Storage C pools were not depleted at death for droughted trees as they were for starved, well-watered trees. Hence drought has a significant detrimental effect on a plant's ability to utilize stored carbon. Unless they can be transported to where they are needed, sufficient carbon reserves alone will not assure survival of a drought except under specific conditions, such as moderate drought, or in species that maintain plant water relations required for carbon re-mobilization.

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.

Dynamic Vegetation Models (DVMs) are designed to be suitable for simulating forest succession and species range dynamics under current and future conditions based on mathematical representations of the three key processes regeneration, growth, and mortality However, mortality formulations in DVMs are typically coarse and often lack an empirical basis, which increases the uncertainty of projections of future forest dynamics and hinders their use for developing adaptation strategies to climate change. Thus, sound tree mortality models are highly needed. We developed parsimonious, species-specific mortality models for 18 European tree species using >90,000 records from inventories in Swiss and German strict forest reserves along a considerable environmental gradient. We comprehensively evaluated model performance and incorporated the new mortality functions in the dynamic forest model ForClim. Tree mortality was successfully predicted by tree size and growth. Only a few species required additional covariates in their final model to consider aspects of stand structure or climate. The relationships between mortality and its predictors reflect the indirect influences of resource availability and tree vitality, which are further shaped by species-specific attributes such as maximum longevity and shade tolerance. Considering that the behavior of the models was biologically meaningful, and that their performance was reasonably high and not impacted by changes in the sampling design, we suggest that the mortality algorithms developed here are suitable for implementation and evaluation in DVMs. In the DVM ForClim, the new mortality functions resulted in simulations of stand basal area and species composition that were generally close to historical observations. However, ForClim performance was poorer than when using the original, coarse mortality formulation. The difficulties of simulating stand structure and species composition, which were most evident for Fagus sylvatica L. and in long-term simulations, resulted from feedbacks between simulated growth and mortality as well as from extrapolation to very small and very large trees. Growth and mortality processes and their species-specific differences should thus be revisited jointly, with a particular focus on small and very large trees in relation to their shade tolerance.

Protecting and enhancing forest carbon sinks is considered a natural solution for mitigating climate change. However, the increasing frequency, intensity, and duration of droughts due to climate change can threaten the stability and growth of existing forest carbon sinks. Extreme droughts weaken plant hydraulic systems, can lead to tree mortality events, and may reduce forest diversity, making forests more vulnerable to subsequent forest disturbances, such as forest fires or pest infestations. Although early warning metrics (EWMs) derived using satellite remote sensing data are now being tested for predicting post-drought plant physiological stress and mortality, applications of unmanned aerial vehicles (UAVs) are yet to be explored extensively. Herein, we provide twenty-four prospective approaches classified into five categories: (i) physiological complexities, (ii) site-specific and confounding (abiotic) factors, (iii) interactions with biotic agents, (iv) forest carbon monitoring and optimization, and (v) technological and infrastructural developments, for adoption, future operationalization, and upscaling of UAV-based frameworks for EWM applications. These UAV considerations are paramount as they hold the potential to bridge the gap between field inventory and satellite remote sensing for assessing forest characteristics and their responses to drought conditions, identifying and prioritizing conservation needs of vulnerable and/or high-carbon-efficient tree species for efficient allocation of resources, and optimizing forest carbon management with climate change adaptation and mitigation practices in a timely and cost-effective manner.

1. Heat waves and droughts are expected to increase in frequency and severity in many regions with future climate change, threatening the survival of a number of forest ecosystems. However, our understanding of the physiological processes and mechanisms underlying drought-induced tree mortality is incomplete. Here, we present results on the physiological response of young Norway spruce trees exposed to lethal drought stress. 2. We applied three levels of drought treatment (control, drying—rewetting, complete drought) and monitored relevant physiological functions and processes of carbon and water relations at high temporal resolution until tree death occurred. 3. Only trees subjected to continuous drought died in our experiment. Trees subjected to drying—rewetting cycles consistently recovered in their ability to transport water, indicating that these trees do not suffer permanent damage to the hydraulic system. In all cases, drought reduced carbon assimilation, caused changes in carbon allocation and appeared to have severely reduced phloem functioning and carbon translocation. Structural growth was sacrificed for carbon investment in maintenance respiration and osmoprotection. Severe drought caused trees to rely on stored carbon reserves but, in contrast to above-ground tissues, only root carbon pools were strongly reduced when trees died. 4. Our results indicate that drought-induced changes in carbon allocation, use and transport differ between above- and below-ground tissues in trees. While root death may have been caused by carbon depletion, this was definitely not the case in above-ground tissues. Our findings indicate that mortality mechanisms are not defined at the organism level but rather within tree compartments.

BackgroundRecent increases in forest tree mortality should increase the abundance coarse woody detritus (CWD) and ultimately lead to increased atmospheric carbon dioxide. However, the time course of carbon release from CWD is not well understood. We compiled CWD decomposition rate-constants (i.e., k) to examine how tree species, piece diameter, position (i.e., standing versus downed), canopy openness, and macroclimate influenced k. To illustrate their implications we modeled the effect of species and position on estimates of decomposition-related carbon flux. We examined a subset of currently used models to determine if their structure accounted for these factors.ResultsGlobally k of downed CWD varied at least 244-fold with interspecies variation at individual sites up to 76-fold. While k generally decreased with increasing piece diameter, under open canopies the opposite occurred. Standing CWD sometimes exhibited little decomposition, but sometimes had k values up to 3 times faster than downed CWD. There was a clear response of k to mean annual temperature of ≈ 2.6 times per 10 ℃; however, there was considerable variation for a given mean annual temperature related to species, diameter, and position. A key feature of carbon release from CWD after disturbance was the “evolution” of the ecosystem-level k value as positions and species mixtures of the remaining CWD changed. Variations in decomposition caused by disturbance (e.g., changes in species, positions, sizes, and microclimate) had the potential to cause net carbon fluxes to the atmosphere to be highly nonlinear. While several models currently being used for carbon accounting and assessing land-use/climate change would potentially capture some of these post disturbance changes in fluxes and carbon balances, many would not.ConclusionsWhile much has been learned in the last 5 decades about CWD decomposition, to fully understand the time course of carbon release from increased mortality and other aspects of global change a new phase of global CWD research that is more systematic, experimental, and replicated needs to be initiated. If our findings are to be fully applied in modeling, an approach acknowledging how the rate of carbon release evolves over time should be implemented.

• Photosynthesis in woody tissues (P wt) is less sensitive to water shortage than in leaves, hence, P wt might be a crucial carbon source to alleviate drought stress. To evaluate the impact of P wt on tree drought tolerance, woody tissues of 4-m-tall drought-stressed Populus tremula trees were subjected to a light-exclusion treatment across the entire plant to inhibit Pwt. • Xylem water potential (Ψ xylem), sap flow (F H₂O), leaf net photosynthesis (P n,l), stem diameter variations (ΔD), in vivo acoustic emissions in stems (AEs) and nonstructural carbohydrate concentrations ([NSC]) were monitored to comprehensively assess water and carbon relations at whole-tree level. • Under well-watered conditions, P wt kept Ψ xylem at a higher level, lowered FH₂O and had no effect on [NSC]. Under drought, Ψ xylem, F H₂O and P n,l in light-excluded trees rapidly decreased in concert with reductions in branch xylem starch concentration. Moreover, sub-daily patterns of ΔD, FH₂O and AEs were strongly related, suggesting that in vivo AEs may inform not only about embolism events, but also about capacitive release and replenishment of stem water pools. • Results highlight the importance of P wt in maintaining xylem hydraulic integrity under drought conditions and in sustaining NSC pools to potentially limit increases in xylem tension.

AIM: To investigate the role of clay in limiting the development of woody biomass in drylands. LOCATION: Australia. METHODS: Spatial data representing woody biomass, soil clay content and climate for the Australian continent were interpreted to explore the relationship between a moisture index and woody biomass on clay and non‐clay substrates. Moisture and soil water potential were investigated through time for clay and sand plains. Sub‐soil clay and other soil characteristics were compared between areas with and without high tree mortality after drought. RESULTS: The substantially lower woody biomass on clay soils than non‐clay soils in the drylands of the Australian continent verifies the inverse texture effect. A clay floodplain consistently held more water than an adjacent sandplain, but soil moisture in the clay was effectively inaccessible to plants for much longer periods because of more negative values of water potential in the clay soil. Small increases in sub‐soil clay were associated with high tree mortality. MAIN CONCLUSIONS: Our findings suggest that the ‘inverse texture effect’, which describes the negative association of woody biomass with soil clay content in drylands, may mostly relate to the moisture‐retentive properties of clay rather than being caused by reduced infiltration and enhanced evaporation from clay‐rich soils. Our evidence suggests that clay soils constrain woodiness in drylands by exacerbating water stress, and confirm the fundamental role of aridity in determining woody biomass and the global distribution of grassland.