Aims The response of understory plants is an important, yet understudied element of forest dynamics associated with mountain pine beetle (MPB) disturbance. Here, we provide a detailed assessment of both the changes in understory species and communities over a 5-year period following severe MPB-induced lodgepole pine mortality. Specifically, we examine species turnover, rarity and mobility over time, and evaluate shifts in community composition, including the potential for post-outbreak community homogenization or differentiation. Location Western Rocky Mountain National Park in the southern Rocky Mountains, Colorado. Methods We collected species occurrence data across lodgepole pine-dominated forest understories using repeated sampling of the same study sites (during the peak of MPB activity and 5 years after). Results More species appeared than disappeared 5 years after the peak of the outbreak. New species were comprised of both early- and late-successional species, suggesting the presence of rather high microhabitat heterogeneity. Although most species remained relatively rare, the number of exceedingly rare species decreased, and the number of highly common species increased. Several species were able to take advantage of the new stand conditions and expand throughout the study area. Some species appeared to spread locally over short distances mainly via vegetative reproduction, others expanded primarily upwards in elevation, probably through long-distance dispersal and some spread over both short and long distances. Non-native species doubled in occurrence and tended to spread upslope, towards the interior of the park, yet remained a small component of the understory overall. Shifts in community composition were minimal, but a slight convergence of plant community groups suggests a trend towards community homogenization. Conclusions Our results demonstrate that even relatively sparse and depauperate lodgepole pine forest understories may be altered in diverse ways following MPB-induced overstory mortality, depending on the unique ability of species to respond to changes in site conditions and resource availability.

Understory plants are an important element of forests, having a considerable influence on ecosystem functioning and canopy-tree development following disturbance. Recent bark beetle outbreaks across western North American forests have caused extensive canopy mortality, creating new growing conditions that provide the opportunity for changes within the intact understory. Over a five-year period following peak mountain pine beetle (MPB) activity across lodgepole pine-dominated forests in Rocky Mountain National Park, Colorado, we measured the changes in plant diversity, cover, and dominance by lifeform and quantified tree regeneration rates. Average species richness and diversity increased, but overall plant cover did not change. Graminoids appeared to benefit the most, increasing in average cover, richness, and relative dominance. The rise in graminoid dominance was largely at the expense of shrubs, which showed little ability to benefit from overstory mortality within the first years following attack. Most plant responses were positively related to the total tree basal area lost since the peak of the outbreak, suggesting that increased resource availability following tree death may benefit understory plants. However, a negative relationship between several understory variables and tree sapling density provides evidence that understory plants compete with saplings for the newly available resources. Tree seedling density nearly doubled over the duration of the study, indicating a strong regeneration pulse. Among species, lodgepole pine displayed the greatest tree seedling establishment. This is one of the first studies to use repeated measurements to describe this often-overlooked component of forest change associated with MPB disturbance.

The persistence and fall rate of snags (standing dead trees) generated during bark beetle outbreaks have consequences for the behavior, effects, and suppression of potential wildfires, hazard tree and timber salvage operations, wildlife habitat, and numerous ecosystem processes. However, post-beetle snagfall dynamics are poorly understood in most forest types. We tagged standing live and dead lodgepole pine (Pinus contorta), subalpine fir (Abies lasiocarpa), and Engelmann spruce (Picea engelmannii), including beetle-killed pine snags following the peak of a recent mountain pine bark beetle outbreak in watersheds at the Fraser Experimental Forest in northcentral Colorado and sampled snagfall 10 and 12 years later. Bark beetle attacks began in 2003, peaked by 2006, and killed 78% of overstory lodgepole pine in 133 plots distributed across a range of stand and site conditions. Of those snags, only 17% fell between 2007 and 2018. Most snags broke at ground level, due to butt rot, and were oriented downhill. In contrast, snags that tipped up or snapped off above the ground were oriented with the prevailing winds. Equal numbers of snags fell singly and in multiple-tree groups, and equal numbers remained elevated rather than in contact with the ground. Lodgepole pine snagfall was 1.6-times higher on steep slopes (>40%) where dead pine density was higher, compared to flatter sites. Based on our findings and previous research, we estimate that one-half the beetle-killed lodgepole pine in high-elevation forests such as those at Fraser may fall within 15-20 yr of beetle infestation, but that some pine snags are likely to persist for decades longer. Post-outbreak snagfall dynamics create a multiple-decade legacy of bark beetle outbreaks that will persist longer in high-elevation compared to lower-elevation forests.

Recent mountain pine beetle (Dendroctonus ponderosae; MPB) outbreaks have caused one of the most widespread and dramatic changes in forest condition in North American forests in more than a century and highlighted challenges facing resource managers. To address uncertainty regarding the consequences of post-harvest woody residue management on soil productivity and tree regeneration following MPB outbreaks in lodgepole pine-dominated forests we compared three treatment prescriptions (bole-only harvest, whole-tree harvest, and whole-tree harvest with scarification) and uncut stands. The study was replicated at twelve sites across a range of operational project areas and stand conditions in northern Colorado. Salvage logging generated a new cohort of lodgepole pine at densities far above the threshold considered adequate to develop into well-stocked stands (1700-2300 t ha(-1) in logged compared to 537 t ha(-1) in uncut areas). Regeneration density was generally highest in whole-tree harvested areas. Growth of planted and naturally regenerating lodgepole pine recruits was best in the bole-only, residue-retention treatment, where soil moisture and inorganic nitrogen supply was also highest. However, we found no indication that whole-tree harvesting lowered soil moisture, soil nitrogen supply or pools relative to uncut stands. The density of trees regenerating beneath uncut stands indicates that post-outbreak forest structure should recover without management in these forests. The cohort of trees that regenerated following MPB-related overstory mortality, but prior to harvesting, comprise the fastest-growing component of the growing stock and 30% of its density. The broader watershed-scale outcomes of these treatments and their implications for wildfire behavior and other effects remain uncertain. However, the soil and tree patterns we report during the initial post-treatment period inform on-going decisions regarding harvest and residue retention and create a platform to guide future forest management research.

Abstract. Understanding the processes that underlie forest resilience is of increasing importance as climate change and shifting disturbance regimes continue to impact western forests. Forest research and management efforts within the low-diversity conifer forests of the U.S. Rocky Mountains have typically focused on relatively monotypic stands dominated by a single cohort, but mixed-conifer stands, such as those codominated by Abies lasiocarpa and Pinus contorta have been less widely studied. The presence of A. lasiocarpa may enhance resilience to fire- and mountain pine beetle–induced mortality and depends on successful A. lasiocarpa recruitment under a range of environmental conditions. The purpose of this study was to quantify the effects of key forest structural characteristics and environmental conditions on recruitment of A. lasiocarpa in a midelevation mixed-conifer forest in the central Rocky Mountains. To address this aim, A. lasiocarpa seedling density, light availability, neighborhood basal area, and soil fertility were measured across 24 plots, and the relative effects of each measured variable, temperature, and precipitation on seedling density were quantified within a Bayesian multilevel regression model. Model results showed nonsignificant effects of climate, light availability, and neighborhood index on seedling density; a significant positive association between seedling density and the interaction between soil fertility and neighborhood index; and a strong negative relationship between seedling density and soil fertility. We posit that the negative association with soil fertility in these nutrient-poor forests reflects an underlying gradient in soil moisture availability that corresponds with water flux pathways. Ultimately, much of the variance in seedling densities was explained by latent plot and year effects, indicating that A. lasiocarpa establishment in this mixed-conifer forest is likely governed by a complex suite of environmental factors that vary across fine spatiotemporal scales.

This chapter describes the snags and coarse woody debris (CWD) in the Greater Yellowstone Ecosystem. Severe forest fires, such as those that occurred in the Greater Yellowstone Ecosystem of 1988, create ephemeral forests of dead trees. For many people the trees are both an eyesore and a waste of salvageable wood. Harvesting the wood of burned trees is an option in many areas, but ecological processes in national parks are allowed to proceed whenever possible with minimal human intervention. The standing dead trees, commonly known as snags, have been falling to the ground and decomposing for millennia in most forest ecosystems. The approach to estimating the conversion of downed CWD to charcoal is based on the volume of a tapered cylinder.

Widespread tree mortality from mountain pine beetle (MPB; Dendroctonus ponderosae Hopkins) outbreaks has prompted forest management activities to reduce crown fire hazard in the Rocky Mountain region. However, little is known about how beetle-related salvage logging and biomass utilization options affect woody surface fuel loads and fuel moisture dynamics. We compared these attributes in salvage-logged lodgepole pine (Pinus contorta var. lab:fact Engelm. Ex S. Wats.) stands harvested using either biomass removal (whole-tree harvest) or biomass retention (bole only harvest) prescriptions with untreated MPB-infested stands. Both prescriptions roughly doubled 1-h and 10-h fuel loads compared to untreated forest. Biomass retention left ten times more 1000-h fuels compared to biomass removal prescription (28 vs 3 Mg ha(-1)). Overall, the woody fuel load was more than twice as high with biomass retention compared to biomass removal (60 vs 25 Mg ha(-1)). Fuel moisture content was lower in salvage logged units compared to untreated forest plots, but it did not differ among the biomass prescriptions. Fine (10-h) and heavy (1000-h) fuels dried to a critical ignition threshold 3-8 weeks earlier in the two prescriptions, respectively, compared to the untreated forests. Salvage logging removes canopy fuels and crown fire hazard, but we found that depending on the amount of biomass retained it can both increase surface fuel load and decrease fuel moisture compared to untreated stands. In the coming years, snag fall will transfer crown to surface fuels in untreated beetle-killed stands adding coarse surface fuel loads surpassing those in treated stands. (C) 2017 Elsevier B.V. All rights reserved.

Species distributions have often been assumed to represent climatic limitations, yet recent evidence has challenged these assumptions and emphasized the potential importance of biotic interactions, dispersal limitation, and disturbance. Despite significant investigation into these factors, an integrated understanding of where and when they may be important is lacking. Here, we review evidence for the factors underlying the historical and contemporary distributions of North American tree species and argue that a cohesive conceptual framework must be informed by an understanding of species ecological and evolutionary history. We further demonstrate that available evidence offers little indication of a significant, independent influence of biotic interactions or dispersal limitation on species distributions. Disturbance may provide important constraints on distributions in limited contexts. Overall, historic and contemporary evidence suggests that species distributions are strongly influenced by climate, yet examples of disequilibrium with climate abound. We propose that differences among life stages and the impacts of human land use may contribute to explain these inconsistencies and are deserving of greater research attention.

Early-seral forests are expanding throughout western North America as fire frequency and annual area burned increase, yet fire behaviour in young postfire forests is poorly understood. We simulated fire behaviour in 24-year-old lodgepole pine (Pinus contorta var. latifolia) stands in Yellowstone National Park, Wyoming, United States using operational models parameterised with empirical fuel characteristics, 50–99% fuel moisture conditions, and 1–60kmhr−1 open winds to address two questions: [1] How does fireline intensity, and crown fire initiation and spread vary among young, lodgepole pine stands? [2] What are the contributions of fuels, moisture and wind on fire behaviour? Sensitivity analysis indicated the greatest contributors to output variance were stand structure mediated wind attenuation, shrub fuel loads and 1000-h fuel moisture for fireline intensity; crown base height for crown fire initiation; and crown bulk density and 1-h fuel moisture for crown fire spread. Simulation results predicted crown fire (e.g. passive, conditional or active types) in over 90% of stands at 50th percentile moisture conditions and wind speeds greater than 3kmhr−1. We conclude that dense canopy characteristics heighten crown fire potential in young, postfire lodgepole pine forests even under less than extreme wind and fuel moisture conditions.

Disturbance and succession have long been of interest in ecology, but how landscape patterns of ecosystem structure and function evolve following large disturbances is poorly understood. After nearly 25 years, lodgepole pine (Pinus contorta var. latifolia) forests that regenerated after the 1988 Yellowstone Fires (Wyoming, USA) offer a prime opportunity to track the fate of disturbance-created heterogeneity in stand structure and function in a wilderness setting. In 2012, we resampled 72 permanent plots to ask (1) How have postfire stand structure and function changed between 11 and 24 yr postfire, and what variables explain these patterns and changes? (2) How has landscape-level (among-stand) variability in postfire stand structure and function changed between 11 and 24 yr postfire? We expected to see evidence of convergence beginning to emerge, but also that initial postfire stem density would still determine trajectories of biomass accumulation. After 24 yr, postfire lodgepole pine density remained very high (mean = 21,738 stems/ha, range = 0-344,067 stems/ha). Stem density increased in most plots between 11 and 24 yr postfire, but declined sharply where 11-yr-postfire stem density was > 72,000 stems/ha. Stems were small in high-density stands, but stand-level lodgepole pine leaf area, foliage biomass, and live aboveground biomass increased over time and with increasing stem density. After 24 yr, mean annual lodgepole pine aboveground net primary production (ANPP) was high (mean = 5 Mg · ha⁻¹ · yr⁻¹, range = 0-16.5 Mg · ha⁻¹ · yr⁻¹). Among stands, lodgepole pine ANPP increased with stem density, which explained 69% of the variation; another 8% of the variation was explained by environmental covariates. Early patterns of postfire lodgepole pine regeneration, which were contingent on prefire serotiny and fire severity, remained the dominant driver of stand structure and function. We observed mechanisms that would lead to convergence in stem density (structure) over time, but it was landscape variation in functional variables that declined substantially. Stand structure and function have not converged across the burned landscape, but our evidence suggests function will converge sooner than structure.

Community-level models (CLMs) aim to improve species distribution modeling (SDM) methods by attempting to explicitly incorporate the influences of interacting species. However, the ability of CLMs to appropriately account for biotic interactions is unclear. We applied CLM and SDM methods to predict the distributions of three dominant conifer tree species in the U.S. Rocky Mountains and compared CLM and SDM predictive accuracy as well as the ability of each approach to accurately reproduce species co-occurrence patterns. We specifically evaluated the performance of two statistical algorithms, MARS and CForest, within both CLM and SDM frameworks. Across all species, differences in SDM and CLM predictive accuracy were slight and can be attributed to differences in model structure rather than accounting for the effects of biotic interactions. In addition, CLMs generally over-predicted species co-occurrence, while SDMs under-predicted co-occurrence. Our results demonstrate no real improvement in the ability of CLMs to account for biotic interactions relative to SDMs. We conclude that alternative modeling approaches are needed in order to accurately account for the effects of biotic interactions on species distributions.

Young, recently burned forests are increasingly widespread throughout western North America, but forest development after large wildfires is not fully understood, especially regarding effects of variable burn severity, environmental heterogeneity, and changes in drivers over time. We followed development of subalpine forests after the 1988 Yellowstone fires by periodically resampling permanent plots established soon after the fires. We asked two questions about patterns and processes over the past 25 years: (1) Are plant species richness and community composition converging or diverging across variation in elevation, soils, burn -severity, and post-fire lodgepole pine (Pinus contorta var. latifolia) density? (2) What are the major controls on post-fire species composition, and has the relative importance of controls changed over time? For question 1, we sampled 10-m(2) plots (n = 552) distributed among three geographic areas that differ in elevation and substrate; plots spanned the spectrum of fire severities and were resampled periodically from 1991 to 2013. For question 2, we sampled 0.25--ha plots (n = 72), broadly distributed across areas that burned as stand-replacing fire, in 1999 and 2012. Richness and species composition diverged early on between infertile low-elevation areas (lower richness) and more fertile high-elevation areas (greater richness). Richness increased rapidly for the first 5 yr post-fire, then leveled off or increased only slowly thereafter. Only 6% of 227 recorded species were nonnative. Some annuals and species with heat-stimulated soil seed banks were associated with severely burned sites. However, most post-fire species had been present before the fire; many survived as roots or rhizomes and regenerated rapidly by sprouting. Among the 72 plots, substrate, temperature, and precipitation (the abiotic template) were consistently important drivers of community composition in 1999 and 2012. Post-fire lodgepole pine abundance was not significant in 1999 but was the most important driving variable by 2012, with a negative effect on presence of most understory species, especially annuals and shade-intolerant herbs. Burn severity was significant in 1999 but not in 2012, and distance to unburned forest had no influence in either year. The 1988 fires did not fundamentally alter subalpine forest community assemblages in Yellowstone, and ecological memory conferred resilience to high-severity fire.

Escalating wildfire in subalpine forests with stand-replacing fire regimes is increasing the extent of early-seral forests throughout the western USA. Post-fire succession generates the fuel for future fires, but little is known about fuel loads and their variability in young post-fire stands. We sampled fuel profiles in 24-year-old post-fire lodgepole pine (Pinus contorta var. latifolia) stands (n = 82) that regenerated from the 1988 Yellowstone Fires to answer three questions. (1) How do canopy and surface fuel loads vary within and among young lodgepole pine stands? (2) How do canopy and surface fuels vary with pre-and post-fire lodgepole pine stand structure and environmental conditions? (3) How have surface fuels changed between eight and 24 years post-fire? Fuel complexes varied tremendously across the landscape despite having regenerated from the same fires. Available canopy fuel loads and canopy bulk density averaged 8.5 Mg/ha (range 0.0-46.6) and 0.24 kg/m(3) (range: 0.0-2.3), respectively, meeting or exceeding levels in mature lodgepole pine forests. Total surface-fuel loads averaged 123 Mg/ha (range: 43-207), and 88% was in the 1,000-h fuel class. Litter, 1-h, and 10-h surface fuel loads were lower than reported for mature lodgepole pine forests, and 1,000-h fuel loads were similar or greater. Among-plot variation was greater in canopy fuels than surface fuels, and within-plot variation was greater than among-plot variation for nearly all fuels. Post-fire lodgepole pine density was the strongest positive predictor of canopy and fine surface fuel loads. Pre-fire successional stage was the best predictor of 100-h and 1,000-h fuel loads in the post-fire stands and strongly influenced the size and proportion of sound logs (greater when late successional stands had burned) and rotten logs (greater when early successional stands had burned). Our data suggest that 76% of the young post-fire lodgepole pine forests have 1,000-h fuel loads that exceed levels associated with high-severity surface fire potential, and 63% exceed levels associated with active crown fire potential. Fire rotations in Yellowstone National Park are predicted to shorten to a few decades and this prediction cannot be ruled out by a lack of fuels to carry repeated fires.

Understanding the rates, trajectories, and spatial variability in succession following severe wildfire is increasingly important for forest managers in western North America and critical for anticipating the resilience or vulnerability of forested landscapes to changing environmental conditions. However, few long-term studies have considered succession following severe, stand-replacing wildfires over large areas. This research focused on succession after the 1988 Yellowstone fires and tested hypotheses within three overarching questions:(1) Are stand structure and function beginning to converge twenty-five years after the Yellowstone Fires, and what mechanisms may contribute to convergence or divergence?(2) Are plant community composition and species richness converging or diverging across gradients in local fire severity, postfire lodgepole pine density, elevation and soil type a quarter-century after the 1988 fires?(3) How do canopy and surface fuels vary across the postfire landscape, and how will the variation in fuels influence potential fire behavior a quarter century postfire? Twenty-five years after the 1988 fires, we resampled permanent plots in lodgepole-pine (Pinus contorta var. latifolia) forests in Yellowstone National Park (Wyoming, USA) that burned in the 1988 fires and were distributed widely across the landscape.Stand structure and function. A quarter century after the 1988 fires, postfire lodgepole pine density remained very high (mean= 21,738 stems ha-1) but varied substantially across the landscape, ranging from zero to 344,000 stems ha-1. Stem density increased in most study plots between postfire years 11 and 24, declining …

More frequent fire activity associated with climate warming is expected to increase the extent of young forest stands in fire-prone landscapes, yet growth rates and biomass allocation patterns in young forests that regenerated naturally following stand-replacing fire have not been well studied. We assessed the structural and functional characteristics of young, postfire lodgepole pine (Pinus contorta var. latifolia) trees across the Yellowstone subalpine plateaus to understand the influence of postflre stand density and age on tree-level aboveground biomass (AB), component biomass (bole, branch, foliage), partitioning to components, tree-level aboveground net primary productivity (ANPP) and leaf area (LA). Sixty 24-year-old lodgepole pine trees were harvested from 21 sites ranging from 500 to 74,667 stems.ha(-1) for development of allometric equations to predict biomass, ANPP and LA. All traits increased nonlinearly with increasing tree basal diameter. Tree-level total AB and component biomass decreased with increasing stand density and increased with age when compared with measurements from 11-year-old trees. Bole partitioning increased with stand density, while foliage and branch wood partitioning declined. Tree-level ANPP and LA decreased significantly with stand density and age. Overall, our results indicate that stand density and age explain much of the variation in tree characteristics and that 24 years after fire, the initial postfire regeneration density is still exerting significant influence on the structure and function of individual trees. (C) 2014 Elsevier B.V. All rights reserved.

Understanding how live and dead forest fuel moisture content (FMC) varies with seasonal weather and stand structure will improve researchers’ and forest managers’ ability to predict the cumulative effects of weather on fuel drying during the fire season and help identify acute conditions that foster wildfire ignition and high rates of fire spread. No studies have investigated the efficacy of predicting FMC using mechanistic water budget models at daily time scales through the fire season nor have they investigated how FMC may vary across space. This study addresses these gaps by (1) validating a novel mechanistic live FMC model and (2) applying this model with an existing dead FMC model at three forest sites using five climate change scenarios to characterize how FMC changes through time and across space. Sites include post-fire 24-year old forest, mature forest with high canopy cover, and mature forest affected by the mountain pine beetle with moderate canopy cover. Climate scenarios include central tendency, warm/dry, warm/wet, hot/dry, and hot/wet.

A warming climate may increase the frequency and severity of stand‐replacing wildfires, reducing carbon (C) storage in forest ecosystems. Understanding the variability of postfire C cycling on heterogeneous landscapes is critical for predicting changes in C storage with more frequent disturbance. We measured C pools and fluxes for 77 lodgepole pine (Pinus contortaDougl. ex Loud var.latifoliaEngelm.) stands in and around Yellowstone National Park (YNP) along a 300‐year chronosequence to examine how quickly forest C pools recover after a stand‐replacing fire, their variability through time across a complex landscape, and the role of stand structure in this variability.Carbon accumulation after fire was rapid relative to the historical mean fire interval of 150–300 years, recovering nearly 80% of prefire C in 50 years and 90% within 100 years. Net ecosystem carbon balance (NECB) declined monotonically, from 160 g C·m−2·yr−1at age 12 to 5 g C·m−2·yr−1at age 250, but was never negative after disturbance. Decomposition and accumulation of dead wood contributed little to NECB relative to live biomass in this system. Aboveground net primary productivity was correlated with leaf area for all stands, and the decline in aboveground net primary productivity with forest age was related to a decline in both leaf area and growth efficiency. Forest structure was an important driver of ecosystem C, with ecosystem C, live biomass C, and organic soil C varying with basal area or tree density in addition to forest age. Rather than identifying a single chronosequence, we found high variability in many components of ecosystem C stocks through time; a >50% random subsample of the sampled stands was necessary to reliably estimate the nonlinear equation coefficients for ecosystem C. At the spatial scale of YNP, this variability suggests that landscape C develops via many pathways over decades and centuries, with prior stand structure, regeneration, and within‐stand disturbance all important. With fire rotation projected to be <30 years by mid century in response to a changing climate, forests in YNP will store substantially less C (at least 4.8 kg C/m2or 30% less).

Range-margin populations may have different life history characteristics than other populations, and the montane forests of Glacier National Park represent the easternmost limit of several western conifer species including Thuja plicata Donn ex D. Don (western redcedar) and Tsuga heterophylla (Raf.) Sarg. (western hemlock). Understanding regeneration strategies of range-margin populations is important for predicting species distribution shifts. We identify successional status, seedling substrate preferences, and the degree to which different species establish within canopy gaps and the forest matrix. Thuja plicata and Tsuga heterophylla were each found to comprise at least 35% of all tree size classes, with six other species each contributing less than 6%. Similarly, other species each comprised less than 5% of canopy gap and understory seedlings. Thuja plicata is typically thought of as a late-successional species but it, along with Tsuga heterophylla, dominated stands during all stages of successional development. Similarly, they dominated both closed forest and gap establishment sites on all rooting substrates but preferred wood for establishment. Even with the likelihood of increased fire frequency, these range-margin populations likely have the capacity to maintain their current distributions. As disturbance regimes are modified across landscapes, it may be possible to observe the potential to adapt to local conditions in other range-margin populations.