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Aims: Our goal was to better understand tree growth and photosynthetic responses to variations in plant available water and elucidate the role of the inverse texture effect in snow dominated montane forests. Methods: We measured tree ring carbon isotope composition and annual growth over a 31-year record for lodgepole pine (Pinus contorta Douglas ex Loudon) growing on three different-aged glacial till surfaces in Wyoming, USA. Results: Soils of different ages developed on till surfaces from three separate glaciation events were significantly different in clay content and distribution with depth; maximum clay content at depth ranged from 19 to 20% on the two youngest till surfaces, but was as high as 36% on the oldest till surface. Ring growth was lowest at the youngest till sites, and only on these coarse-textured soils was growth positively correlated with annual maximum snow water equivalent (SWEmax). Δ13C was highest for trees at these young till sites, suggesting that hydraulic conductivity and stomatal conductance is comparatively high during growth periods on these coarse-textured soils. Conclusion: Taken together, we found that age of glacial till and related soil texture differences strongly influenced tree-ring growth and Δ13C response of lodgepole pine to interannual variation in precipitation and drought severity, but responses did not support an inverse texture effect in these semi-arid forest systems. [ABSTRACT FROM AUTHOR]

The degree to which recent bark beetle (Dendroctonus ponderosae) outbreaks may influence fire severity and postfire tree regeneration is of heightened interest to resource managers throughout western North America, but empirical data on actual fire effects are lacking. Outcomes may depend on burning conditions (i.e., weather during fire), outbreak severity, or intervals between outbreaks and subsequent fire. We studied recent fires that burned through green-attack/red-stage (outbreaks < 3 years before fire) and gray-stage (outbreaks 3-15 years before fire) subalpine forests dominated by lodgepole pine (Pinus contorta var. latifolia) in Greater Yellowstone, Wyoming, USA, to determine if fire severity was linked to prefire beetle outbreak severity and whether these two disturbances produced compound ecological effects on postfire tree regeneration. With field data from 143 postfire plots that burned under different conditions, we assessed canopy and surface fire severity, and postfire tree seedling density against prefire outbreak severity. In the green-attack/red stage, several canopy fire-severity measures increased with prefire outbreak severity under moderate burning conditions. Under extreme conditions, few fireseverity measures were related to prefire outbreak severity, and effect sizes were of marginal biological significance. The percentage of tree stems and basal area killed by fire increased with more green-attack vs. red-stage trees (i.e., the earliest stages of outbreak). In the gray stage, by contrast, most fire-severity measures declined with increasing outbreak severity under moderate conditions, and fire severity was unrelated to outbreak severity under extreme burning conditions. Postfire lodgepole pine seedling regeneration was unrelated to prefire outbreak severity in either post-outbreak stage, but increased with prefire serotiny. Results suggest bark beetle outbreaks can affect fire severity in subalpine forests under moderate burning conditions, but have little effect on fire severity under extreme burning conditions when most large wildfires occur in this system. Thus, beetle outbreak severity was moderately linked to fire severity, but the strength and direction of the linkage depended on both endogenous (outbreak stage) and exogenous (fire weather) factors. Closely timed beetle outbreak and fire did not impart compound effects on tree regeneration, suggesting the presence of a canopy seedbank may enhance resilience to their combined effects. [ABSTRACT FROM AUTHOR]

Abstract: Outbreaks of bark beetles and drought both lead to concerns about increased fire risk, but the relative importance of these two factors is the subject of much debate. We examined how mountain pine beetle (MPB) outbreaks and drought have contributed to the fire regime of lodgepole pine forests in northwestern Colorado and adjacent areas of southern Wyoming over the past century. We used dendroecological methods to reconstruct the pre-fire history of MPB outbreaks in twenty lodgepole pine stands that had burned between 1939 and 2006 and in 20 nearby lodgepole pine stands that were otherwise similar but that had not burned. Our data represent c. 80% of all large fires that had occurred in lodgepole pine forests in this study area over the past century. We also compared Palmer Drought Severity Index (PDSI) and actual evapotranspiration (AET) values between fire years and non-fire years. Burned stands were no more likely to have been affected by outbreak prior to fires than were nearby unburned stands. However, PDSI and AET values were both lower during fire years than during non-fire years. This work indicates that climate has been more important than outbreaks to the fire regime of lodgepole pine forests in this region over the past century. Indeed, we found no detectable increase in the occurrence of high-severity fires following MPB outbreaks. Dry conditions, rather than changes in fuels associated with outbreaks, appear to be most limiting to the occurrence of severe fires in these forests. [Copyright &y& Elsevier]

We characterised the remarkable heterogeneity following the large, severe fires of 1988 in Yellowstone National Park (YNP), in the northern Rocky Mountains, Wyoming, USA, by focussing on spatial variation in post-fire structure, composition and ecosystem function at broad, meso, and fine scales. Ecological heterogeneity at multiple scales may enhance resilience to large, severe disturbances by providing structural, biological and functional redundancy. Post-fire heterogeneity in stand age, coarse wood abundance, microbial and understorey communities reflected interactions between existing pre-fire patterns and fire severity at different scales, suggesting that environmental context plays an important role in successional responses to large fires. In response to these post-fire patterns, heterogeneity in carbon (C) and nitrogen (N) storage, N mineralisation, decomposition, and productivity was also evident at multiple scales and may confer resiliency to large fires. For example, at broad scales, C storage in YNP appears resistant to changes in age-class structure associated with large stand-replacing fires. In summary, the YNP landscape is recovering rapidly from the 1988 fires through natural mechanisms, owing to the abundance and spatial heterogeneity of post-fire residuals, but other systems with fewer biotic legacies may be less resilient to such large, severe fires. [ABSTRACT FROM AUTHOR]

Aim The historical variability of fire regimes must be understood in the context of drivers of the occurrence of fire operating at a range of spatial scales from local site conditions to broad-scale climatic variation. In the present study we examine fire history and variations in the fire regime at multiple spatial and temporal scales for subalpine forests of Engelmann spruce–subalpine fir ( Picea engelmannii, Abies lasiocarpa) and lodgepole pine ( Pinus contorta) of the southern Rocky Mountains. Location The study area is the subalpine zone of spruce–fir and lodgepole pine forests in the southern sector of Rocky Mountain National Park (ROMO), Colorado, USA, which straddles the continental divide of the northern Colorado Front Range (40°20′ N and 105°40′ W). Methods We used a combination of dendroecological and Geographic Information System methods to reconstruct fire history, including fire year, severity and extent at the forest patch level, for c. 30,000 ha of subalpine forest. We aggregated fire history information at appropriate spatial scales to test for drivers of the fire regime at local, meso, and regional scales. Results The fire histories covered c. 30,000 ha of forest and were based on a total of 676 partial cross-sections of fire-scarred trees and 6152 tree-core age samples. The subalpine forest fire regime of ROMO is dominated by infrequent, extensive, stand-replacing fire events, whereas surface fires affected only 1–3% of the forested area. Main conclusions Local-scale influences on fire regimes are reflected by differences in the relative proportions of stands of different ages between the lodgepole pine and spruce–fir forest types. Lodgepole pine stands all originated following fires in the last 400 years; in contrast, large areas of spruce–fir forests consisted of stands not affected by fire in the past 400 years. Meso-scale influences on fire regimes are reflected by fewer but larger fires on the west vs. east side of the continental divide. These differences appear to be explained by less frequent and severe drought on the west side, and by the spread of fires from lower-elevation mixed-conifer montane forests on the east side. Regional-scale climatic variation is the primary driver of infrequent, large fire events, but its effects are modulated by local- and meso-scale abiotic and biotic factors. The low incidence of fire during the period of fire-suppression policy in the twentieth century is not unique in comparison with the previous 300 years of fire history. There is no evidence that fire suppression has resulted in either the fire regime or current forest conditions being outside their historic ranges of variability during the past 400 years. Furthermore, in the context of fuel treatments to reduce fire hazard, regardless of restoration goals, the association of extremely large and severe fires with infrequent and exceptional drought calls into question the future effectiveness of tree thinning to mitigate fire hazard in the subalpine zone. [ABSTRACT FROM AUTHOR]

To evaluate the long-term effects of clearcutting and slash treatment on understory plant species composition in Rocky Mountain coniferous forests, we collected comparable data from 30- to 50-year-old post-harvest stands and adjacent mature (>100-year-old) stands that originated after wildfire. In general, plant species richness was higher in the post-harvest stands than in adjacent mature stands, due in large part to the presence of species that benefit from disturbance. Species composition in post-harvest and mature stands was more similar in montane forests (2710–2805 m elevation) than in subalpine forests (2800–3164 m elevation), suggesting greater resilience of montane understories. Composition in post-harvest and mature stands was least similar when slash was piled and burned, and most similar when slash was lopped and scattered. Eighty-seven percent of the post-harvest stands had at least one exotic species, but total cover of exotic plant species in all post-harvest stands was ≤1%. Exotic species were largely absent from mature forests. Our results suggest that understory species of low-elevation montane forests recover more quickly from disturbance than do those of subalpine forests, with slash treatment and species life history traits strongly influencing this recovery. [Copyright &y& Elsevier]

• Snags are important components of forest ecosystems. • Snag fall occurred in every snag age class with the greatest proportion occurring 4–8 years since death (YSD). • Snags 13 and 14 years YSD had the lowest fall rates despite being the oldest. • Slope aspect had the strongest influence on fall rates. • The predicted half-life of snags was ~16 YSD. Snags (standing dead trees) are important components of forest ecosystems that, among other benefits, provide critical habitat for many species of wildlife, but also represent important safety concerns to firefighters, forest workers, and the public. We identified factors that influence the fall rates of lodgepole pines, Pinus contorta Dougl. ex Loud., killed by bark beetles during a severe regional-scale outbreak of mountain pine beetle, Dendroctonus ponderosae Hopkins, that occurred in 2004–2012. Data were obtained during annual assessments (2010–2019) of a network of 107 0.081-ha circular plots installed in P. contorta forests in Colorado, Idaho, Montana, Utah and Wyoming, U.S. A total of 3789 P. contorta snags were monitored, and in each case we recorded, among other variables, the year the snag was created and the year the snag fell to the forest floor. Among snag age classes (i.e., number of years since tree death), the highest number of snags (1046) were 12 years since death (YSD), and those 13 and 14 YSD exhibited the lowest fall rates to date (<10%) despite being the oldest in our study. Snags 13 and 14 YSD were among the largest in diameter. Snag fall occurred in every snag age class from 1 to 14 YSD, with the greatest proportion of snag fall events occurring 4–8 YSD. By 2019, 24.7% (937) of snags fell to the forest floor. We modeled snag fall using a Cox's proportional-hazards model that included six covariates of interest: elevation (m), slope aspect (categorical, cardinal direction), slope (%), canopy cover (%), snag height (m), and snag height:dbh (m, diameter at 1.37 m in height). Slope aspect had the strongest influence on fall rates. Northern aspects, increased canopy cover, and taller snag heights decreased the probability of snag fall. Conversely, southern aspects and increased height:dbh ratios (i.e., taller, skinner snags) increased the probability of snag fall. The predicted half-life (i.e., the amount of time since death required for 50% of the snag population to fall to the forest floor) was ~16 YSD after which the function predicted a linear, ~0.04/year decline in snag survival probability 15–30 YSD. The observed longevity of P. contorta snags confers important ecological benefits for some wildlife, and may offer opportunities for extended periods of salvage. However, checking (cracks) rapidly occurred, and many snags 6 YSD and older had at least one check at 1.37 m and 5.5 m in height, which can negatively impact lumber recovery. The relatively slow snag fall rates observed in our study also lengthen concerns regarding hazard trees, human safety, and protection of critical infrastructure. The implications of these and other results to management of P. contorta forests are discussed. [ABSTRACT FROM AUTHOR]

• Significant reductions in tree size and density occurred. • Tree mortality was positively correlated with tree density. • Significant increases in snag density occurred. • Lodgepole pine remained the dominant tree species despite increases in subalpine fir. • A long-term shift in tree composition is unlikely. Mountain pine beetle, Dendroctonus ponderosae Hopkins (Coleoptera: Curculionidae), is the most important forest insect in western North America. We determined causes and rates of tree mortality and changes in forest structure and composition associated with D. ponderosae outbreaks in the Intermountain West, U.S. during 2004–2019 based on a network of 125 0.081-ha circular plots installed in Colorado, Idaho, Montana, Utah and Wyoming. Incipient populations of D. ponderosae began in 2004; peaked in 2007; and returned to endemic levels in 2011 in Idaho, Montana, Utah and Wyoming. In Colorado, incipient populations began in 2004; peaked in 2009; and returned to endemic levels in 2012. A total of 5107 trees died, 98.6% were lodgepole pine, Pinus contorta Dougl. ex Loud. Fifteen contributing factors were identified, including (in order of importance, highest to lowest) D. ponderosae , unknown causes, pine engraver, Ips pini (Say), wind, breakage and/or adjacent tree fall, Pityogenes knechteli Swaine/ Pityopthorus confertus Swaine , suppression, spruce beetle, Dendroctonus rufipennis (Kirby), root disease, western balsam bark beetle, Dryocoetes confusus (Swain), lodgepole pine dwarf mistletoe, Arceuthobium americanum Nutt. ex. Engelm., stem diseases, woodborers, North American porcupine, Erethizon dorsatum (L.), mule deer, Odocoileus hemionus (Rafinesque), and lodgepole pine beetle, Dendroctonus murrayanae Hopkins. Most tree mortality (68.8%) was attributed solely to D. ponderosae , although D. ponderosae also occurred in association with other contributing factors. Overall, significant reductions in mean dbh (by 5.3%), mean quadratic mean diameter (by 8.6%), mean tree height (by 15.9%), mean number of trees (by 40.8%), mean basal area (by 52.9%), and mean stand density index (SDI) (by 51.8%) were observed. Significant reductions in tree density were observed in all diameter classes, except the smallest (midpoint = 10 cm, 5-cm classes). Significant increases in the mean number of snags (by 1324.7%) were observed, and most snags remain standing (71.3%). Pinus contorta remains the dominant tree species, and while significant increases in the number of subalpine fir, Abies lasiocarpa (Hooker) Nuttall, seedlings and saplings were observed, a long-term shift in tree composition is unlikely. Tree mortality (number of trees killed) was positively correlated with the initial number of live trees, basal area of live trees, SDI, and aspect, but not slope or elevation. The implications of these and other results to recovery and management of P. contorta forests in the Intermountain West are discussed. [ABSTRACT FROM AUTHOR]

Forests store significant quantities of carbon, and accurate quantification of the fate of this carbon after fire is necessary for global carbon accounting. Pyrogenic carbon (PyC) encompasses various carbonaceous products of incomplete combustion formed during fires and has potential to act as a carbon sink for up to millennia, but current estimates of PyC production in wildfires vary widely. Northern hardwood forests have changed dramatically in recent decades due to insect epidemics, such as the bark beetle epidemic in the Rocky Mountain Region which has caused widespread mortality. This study assessed impacts of bark beetle-induced mortality on fuel pyrolysis kinetics, carbon partitioning of combustion products, and net heat output to aid in forest fire modeling and carbon accounting by comparing healthy and beetle-killed lodgepole pine tree boles burned in a 2018 forest fire in southeast Wyoming, USA with unburned boles. Results showed charring predominantly restricted to the bark and cambium. Significant differences between burned and unburned healthy and beetle-impacted bark/cambium compositions were identified, and PyC production and energy output were quantified. Charring extent and PyC content were found to be greater in beetle-impacted boles due to a reduction in bark/cambium resistance to heating and charring, with 80 times more PyC produced in a beetle-killed bark/cambium than in a healthy bark/cambium. Upon scale-up, total PyC production in the fire-affected area was estimated to be 0.71 GgPyC (82.5 kgPyC/ha). This was found to be significantly enhanced compared to an estimated PyC production of 0.036 GgPyC (4.12 kgPyC/ha) in a hypothetical healthy lodgepole pine ecosystem of equal area. The results of this investigation concluded that the 58% beetle-induced mortality in the Badger Creek Fire area resulted in 3 times more carbon released to the global atmosphere, 20 times more PyC retained onsite and 32% greater heat output during wildfire., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2020 Elsevier B.V. All rights reserved.)

Subalpine forests in the northern Rocky Mountains have been resilient to stand-replacing fires that historically burned at 100- to 300-year intervals. Fire intervals are projected to decline drastically as climate warms, and forests that reburn before recovering from previous fire may lose their ability to rebound. We studied recent fires in Greater Yellowstone (Wyoming, United States) and asked whether short-interval (<30 years) stand-replacing fires can erode lodgepole pine ( Pinus contorta var. latifolia ) forest resilience via increased burn severity, reduced early postfire tree regeneration, reduced carbon stocks, and slower carbon recovery. During 2016, fires reburned young lodgepole pine forests that regenerated after wildfires in 1988 and 2000. During 2017, we sampled 0.25-ha plots in stand-replacing reburns ( n = 18) and nearby young forests that did not reburn ( n = 9). We also simulated stand development with and without reburns to assess carbon recovery trajectories. Nearly all prefire biomass was combusted ("crown fire plus") in some reburns in which prefire trees were dense and small (≤4-cm basal diameter). Postfire tree seedling density was reduced sixfold relative to the previous (long-interval) fire, and high-density stands (>40,000 stems ha -1 ) were converted to sparse stands (<1,000 stems ha -1 ). In reburns, coarse wood biomass and aboveground carbon stocks were reduced by 65 and 62%, respectively, relative to areas that did not reburn. Increased carbon loss plus sparse tree regeneration delayed simulated carbon recovery by >150 years. Forests did not transition to nonforest, but extreme burn severity and reduced tree recovery foreshadow an erosion of forest resilience., Competing Interests: The authors declare no conflict of interest.

The extent of young postfire conifer forests is growing throughout western North America as the frequency and size of high-severity fires increase, making it important to understand ecosystem structure and function in early seral forests. Understanding nitrogen (N) dynamics during postfire stand development is especially important because northern conifers are often N limited. We resampled lodgepole pine (Pinus contorta var. latifolia) stands that regenerated naturally after the 1988 fires in Yellowstone National Park (Wyoming, USA) to ask (1) How have N pools and fluxes changed over a decade (15 to 25 yr postfire) of very rapid forest growth? (2) At postfire year 25, how do N pools and fluxes vary with lodgepole pine density and productivity? Lodgepole pine foliage, litter (annual litterfall, forest-floor litter), and mineral soils were sampled in 14 plots (0.25 ha) that varied in postfire lodgepole pine density (1,500 to 344,000 stems/ha) and aboveground net primary production (ANPP; 1.4 to 16.1 Mg·ha -1 ·yr -1 ). Counter to expectation, foliar N concentrations in lodgepole pine current-year and composite needles (1.33 and 1.11% N, respectively) had not changed over time. Further, all measured ecosystem N pools increased substantially: foliar N increased to 89 kg N/ha (+93%), O-horizon N increased to 39 kg N/ha (+38%), and mineral soil percent total N (0-15 cm) increased to 0.08% (+33%). Inorganic N availability also increased to 0.69 μg N·[g resin] -1 ·d -1 (+165%). Thus, soil N did not decline as live biomass N pools increased. Among stands, biomass N pools at postfire year 25 remained strongly influenced by early postfire tree density: foliar and litterfall N concentrations declined with lodgepole pine density and ANPP, but the foliar N pool increased. Lodgepole pine ANPP correlated negatively with annual resin-sorbed N, and we found no indication of widespread N limitation. The large increases in N pools cannot be explained by atmospheric N deposition or presence of known N fixers. These results suggest an unmeasured N source and are consistent with recent reports of N fixation in young lodgepole pine., (© 2019 by the Ecological Society of America.)