Fire frequency in low-elevation coniferous forests in western North America has greatly declined since the late 1800s. In many areas, this has increased tree density and the proportion of ...shade-tolerant species, reduced resource availability, and increased forest susceptibility to forest insect pests and high-severity wildfire. In response, treatments are often implemented with the goal of increasing ecosystem resilience by increasing resistance to disturbance. We capitalized on an existing replicated study of fire and stand density treatments in a ponderosa pine (Pinus ponderosa)–Douglas-fir (Pseudotsuga menziesii) forest in western Montana, USA, that experienced a naturally occurring mountain pine beetle (MPB; Dendroctonus ponderosae) outbreak 5 yr after implementation of fuels treatments. We explored whether treatment effects on tree-level defense and stand structure affected resistance to MPB. Mortality from MPB was highest in the denser, untreated control and burn-only treatments, with approximately 50% and 39%, respectively, of ponderosa pine killed during the outbreak, compared to almost no mortality in the thin-only and thin-burn treatments. Thinning treatments, with or without fire, dramatically increased tree growth and resin ducts relative to control and burn-only treatments. Prescribed burning did not increase resin ducts but did cause changes in resin chemistry that may have affected MPB communication and lowered attack success. While ponderosa pine remained dominant in the thin and thin-burn treatments after the outbreak, the high pine mortality in the control and burn-only treatment caused a shift in species dominance to Douglas-fir. The high Douglas-fir component in the control and burn-only treatments due to 20th century fire exclusion, coupled with high pine mortality from MPB, has likely reduced resilience of this forest beyond the ability to return to a ponderosa pine-dominated system in the absence of further fire or mechanical treatment. Our results show treatments designed to increase resistance to high-severity fire in ponderosa pine-dominated forests in the Northern Rockies can also increase resistance to MPB, even during an outbreak. This study suggests that fuel and restoration treatments in fire-dependent ponderosa pine forests that reduce tree density increase ecosystem resilience in the short term, while the reintroduction of fire is important for long-term resilience.
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Each year wildland fires kill and injure trees on millions of forested hectares globally, affecting plant and animal biodiversity, carbon storage, hydrologic processes, and ecosystem services. The ...underlying mechanisms of fire-caused tree mortality remain poorly understood, however, limiting the ability to accurately predict mortality and develop robust modeling applications, especially under novel future climates. Virtually all post-fire tree mortality prediction systems are based on the same underlying empirical model described in Ryan and Reinhardt (1988 Can. J. For. Res. 18 1291-7), which was developed from a limited number of species, stretching model assumptions beyond intended limits. We review the current understanding of the mechanisms of fire-induced tree mortality, provide recommended standardized terminology, describe model applications and limitations, and conclude with key knowledge gaps and future directions for research. We suggest a two-pronged approach to future research: (1) continued improvements and evaluations of empirical models to quantify uncertainty and incorporate new regions and species and (2) acceleration of basic, physiological research on the proximate and ultimate causes of fire-induced tree mortality to incorporate processes of tree death into models. Advances in both empirical and process fire-induced tree modeling will allow creation of hybrid models that could advance understanding of how fire injures and kills trees, while improving prediction accuracy of fire-driven feedbacks on ecosystems and landscapes, particularly under novel future conditions.
Tree mortality associated with drought and concurrent bark beetle outbreaks is expected to increase with further climate change. When these two types of disturbance occur in concert it complicates ...our ability to accurately predict future forest mortality. The recent extreme California USA drought and bark beetle outbreaks resulted in extensive tree mortality and provides a unique opportunity to examine questions of why some trees die while others survive these co-occurring disturbances. We use plot-level data combined with a three-proxy tree-level approach using radial growth, carbon isotopes, and resin duct metrics to evaluate 1) whether variability in stand structure, tree growth or size, carbon isotope discrimination, or defenses precede mortality, 2) how relationships between these proxies differ for surviving and now-dead trees, and 3) whether generalizable risk factors for tree mortality exist across pinyon pine (Pinus monophylla), ponderosa pine (P. ponderosa), white fir (Abies concolor), and incense cedar (Calocedrus decurrens) affected by the combination of drought and beetle outbreaks. We find that risk factors associated with mortality differ between species, and that few generalizable patterns exist when bark beetle outbreaks occur in concert with a particularly long, hot drought. We see evidence that both long-term differences in physiology and shorter-term beetle-related selection and variability in defenses influence mortality susceptibility for ponderosa pine, whereas beetle dynamics may play a more prominent role in mortality patterns for white fir and pinyon pine. In contrast, incense cedar mortality appears to be attributable to long-term effects of growth suppression. Risk factors that predispose some trees to drought and beetle-related mortality likely reflect species-specific strategies for dealing with these particular disturbance types. The combined influence of beetles and drought necessitates the consideration of multiple, species-specific risk factors to more accurately model forest mortality in the face of similar extreme events more likely under future climates.
Identified within-species risk factors for tree-level mortality due to the combination of severe drought and concurrent bark beetle outbreaks associated with the 2012–2016 California, USA drought. Risk factors are categorized by species and category of metric measured. While we do not attempt to disentangle the effects of drought and bark beetles, host-specific beetles were identified as contributing to mortality for ponderosa pine, pinyon pine, and white fir, while incense cedar showed no clear evidence of bark beetle attacks. BAI refers to basal area increment (i.e. radial tree growth), Δ13C refers to discrimination of the heavier (13C) carbon isotope measured from tree rings, and PDSI refers to the Palmer Drought Severity Index. Display omitted
•Recent drought and bark beetles caused extensive tree mortality in California, USA.•A three-proxy approach is used to elucidate risk factors for tree-level mortality.•Living and dead trees exhibit some differences in growth, isotopes (Δ13C) and defenses.•Differences between living and dead trees are not consistent across species.•Effects of drought with beetles mask generalizable risk factors of mortality.
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As the climate warms, drought will increasingly occur under elevated temperatures, placing forest ecosystems at growing risk of extensive dieback and mortality. In some cases, increases in tree ...density following early 20th-century fire suppression may exacerbate this risk. Treatments designed to restore historical stand structure and enhance resistance to high-severity fire might also alleviate drought stress by reducing competition, but the duration of these effects and the underlying mechanisms remain poorly understood. To elucidate these mechanisms, we evaluate tree growth, mortality, and tree-ring stable-carbon isotope responses to stand-density reduction treatments with and without prescribed fire in a ponderosa pine forest of western Montana. Moderate and heavier cutting experiments (basal area reductions of 35% and 56%, respectively) were initiated in 1992, followed by prescribed burning in a subset of the thinned units. All treatments led to a growth release that persisted to the time of resampling. The treatments had little effect on climate–growth relationships, but they markedly altered seasonal carbon isotope signals and their relationship to climate. In burned and unburned treatments, carbon isotope discrimination (Δ13C) increased in the earlywood (EW) and decreased in the latewood (LW) relative to the control. The sensitivity of LW Δ13C to late-summer climate also increased in all treatments, but not in the control. Such increased sensitivity indicates that the reduction in competition enabled trees to continue to fix carbon for new stem growth, even when the climate became sufficiently stressful to stop new assimilation in slower-growing trees in untreated units. These findings would have been masked had we not separated EW and LW. The importance of faster growth and enhanced carbon assimilation under late-summer climatic stress became evident in the second decade post-treatment, when mountain pine beetle activity increased locally, and tree mortality rates in the controls of both experiments increased to more than twice those in their respective treatments. These findings highlight that, when thinning is used to restore historical forest structure or increase resistance to high-severity fire, there will likely be additional benefits of enhanced growth and physiological activity under climatic stress, and the effects may persist for more than two decades.
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Trees use nonstructural carbohydrates (NSCs) to support many functions, including recovery from disturbances. However, NSC's importance for recovery following fire and whether NSC depletion ...contributes to post-fire delayed mortality are largely unknown. We investigated how fire affects NSCs based on fire-caused injury from a prescribed fire in a young Pinus ponderosa (Lawson & C. Lawson) stand. We assessed crown injury (needle scorch and bud kill) and measured NSCs of needles and inner bark (i.e., secondary phloem) of branches and main stems of trees subject to fire and at an adjacent unburned site. We measured NSCs pre-fire and at six timesteps post-fire (4 days-16 months). While all trees initially survived the fire, NSC concentrations declined quickly in burned trees relative to unburned controls over the same post-fire period. This decline was strongest for trees that eventually died, but those that survived recovered to unburned levels within 14 months post-fire. Two months post-fire, the relationship between crown scorch and NSCs of the main stem inner bark was strongly negative (Adj-R2 = 0.83). Our results support the importance of NSCs for tree survival and recovery post-fire and suggest that post-fire NSC depletion is in part related to reduced photosynthetic leaf area that subsequently limits carbohydrate availability for maintaining tree function. Crown scorch is a commonly measured metric of tree-level fire severity and is often linked to post-fire tree outcome (i.e., recovery or mortality). Thus, our finding that NSC depletion may be the mechanistic link between the fire-caused injury and tree outcome will help improve models of post-fire tree mortality and forest recovery.
Fuel and restoration treatments seeking to mitigate the likelihood of uncharacteristic high‐severity wildfires in forests with historically frequent, low‐severity fire regimes are increasingly ...common, but long‐term treatment effects on fuels, aboveground carbon, plant community structure, ecosystem resilience, and other ecosystem attributes are understudied. We present 20‐year responses to thinning and prescribed burning treatments commonly used in dry, low‐elevation forests of the western United States from a long‐term study site in the Northern Rockies that is part of the National Fire and Fire Surrogate Study. We provide a comprehensive synthesis of short‐term (<4 years) and mid‐term (<14 years) results from previous findings. We then place these results in the context of a mountain pine beetle (MPB; Dendroctonus ponderosae) outbreak that impacted the site 5–10 years post‐treatment and describe 20‐year responses to assess the longevity of restoration and fuel reduction treatments in light of the MPB outbreak. Thinning treatments had persistently lower forest density and higher tree growth, but effects were more pronounced when thinning was combined with prescribed fire. The thinning+prescribed fire treatment had the additional benefit of maintaining the highest proportion of ponderosa pine (Pinus ponderosa) for overstory and regeneration. No differences in understory native plant cover and richness or exotic species cover remained after 20 years, but exotic species richness, while low relative to native species, was still higher in the thinning+prescribed fire treatment than the control. Aboveground live carbon stocks in thinning treatments recovered to near control and prescribed fire treatment levels by 20 years. The prescribed fire treatment and control had higher fuel loads than thinning treatments due to interactions with the MPB outbreak. The MPB‐induced changes to forest structure and fuels increased the fire hazard 20 years post‐treatment in the control and prescribed fire treatment. Should a wildfire occur now, the thinning+prescribed fire treatment would likely have the lowest intensity fire and highest tree survival and stable carbon stocks. Our findings show broad support that thinning and prescribed fire increase ponderosa pine forest resilience to both wildfire and bark beetles for up to 20 years, but efficacy is waning and additional fuel treatments are needed to maintain resilience.
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Mountain pine beetle (MPB, Dendroctonus ponderosae) is a significant mortality agent of Pinus, and climate-driven range expansion is occurring. Pinus defenses in recently invaded areas, including ...high elevations, are predicted to be lower than in areas with longer term MPB presence. MPB was recently observed in high-elevation forests of the Great Basin (GB) region, North America. Defense and susceptibility in two long-lived species, GB bristlecone pine (Pinus longaeva) and foxtail pine (P. balfouriana), are unclear, although they are sympatric with a common MPB host, limber pine (P. flexilis).
We surveyed stands with sympatric GB bristlecone–limber pine and foxtail–limber pine to determine relative MPB attack susceptibility and constitutive defenses.
MPB-caused mortality was extensive in limber, low in foxtail and absent in GB bristlecone pine. Defense traits, including constitutive monoterpenes, resin ducts and wood density, were higher in GB bristlecone and foxtail than in limber pine.
GB bristlecone and foxtail pines have relatively high levels of constitutive defenses which make them less vulnerable to climate-driven MPB range expansion relative to other highelevation pines. Long-term selective herbivore pressure and exaptation of traits for tree longevity are potential explanations, highlighting the complexity of predicting plant–insect interactions under climate change.
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Summary
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.
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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.
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10.
Tree physiology and bark beetles Ryan, Michael G.; Sapes, Gerard; Sala, Anna ...
The New phytologist,
02/2015, Volume:
205, Issue:
3
Journal Article
Peer reviewed
Open access
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