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Increasing fire severity and warmer, drier postfire conditions are making forests in the western United States (West) vulnerable to ecological transformation. Yet, the relative importance of and interactions between these drivers of forest change remain unresolved, particularly over upcoming decades. Here, we assess how the interactive impacts of changing climate and wildfire activity influenced conifer regeneration after 334 wildfires, using a dataset of postfire conifer regeneration from 10,230 field plots. Our findings highlight declining regeneration capacity across the West over the past four decades for the eight dominant conifer species studied. Postfire regeneration is sensitive to high-severity fire, which limits seed availability, and postfire climate, which influences seedling establishment. In the near-term, projected differences in recruitment probability between low- and high-severity fire scenarios were larger than projected climate change impacts for most species, suggesting that reductions in fire severity, and resultant impacts on seed availability, could partially offset expected climate-driven declines in postfire regeneration. Across 40 to 42% of the study area, we project postfire conifer regeneration to be likely following low-severity but not high-severity fire under future climate scenarios (2031 to 2050). However, increasingly warm, dry climate conditions are projected to eventually outweigh the influence of fire severity and seed availability. The percent of the study area considered unlikely to experience conifer regeneration, regardless of fire severity, increased from 5% in 1981 to 2000 to 26 to 31% by mid-century, highlighting a limited time window over which management actions that reduce fire severity may effectively support postfire conifer regeneration.

Fire is an integral component of ecosystems globally and a tool that humans have harnessed for millennia. Altered fire regimes are a fundamental cause and consequence of global change, impacting people and the biophysical systems on which they depend. As part of the newly emerging Anthropocene, marked by human-caused climate change and radical changes to ecosystems, fire danger is increasing, and fires are having increasingly devastating impacts on human health, infrastructure, and ecosystem services. Increasing fire danger is a vexing problem that requires deep transdisciplinary, trans-sector, and inclusive partnerships to address. Here, we outline barriers and opportunities in the next generation of fire science and provide guidance for investment in future research. We synthesize insights needed to better address the long-standing challenges of innovation across disciplines to (i) promote coordinated research efforts; (ii) embrace different ways of knowing and knowledge generation; (iii) promote exploration of fundamental science; (iv) capitalize on the “firehose” of data for societal benefit; and (v) integrate human and natural systems into models across multiple scales. Fire science is thus at a critical transitional moment. We need to shift from observation and modeled representations of varying components of climate, people, vegetation, and fire to more integrative and predictive approaches that support pathways toward mitigating and adapting to our increasingly flammable world, including the utilization of fire for human safety and benefit. Only through overcoming institutional silos and accessing knowledge across diverse communities can we effectively undertake research that improves outcomes in our more fiery future.

Background Forest and nonforest ecosystems of the western United States are experiencing major transformations in response to land-use change, climate warming, and their interactive effects with wildland fire. Some ecosystems are transitioning to persistent alternative types, hereafter called “vegetation type conversion” (VTC). VTC is one of the most pressing management issues in the southwestern US, yet current strategies to intervene and address change often use trial-and-error approaches devised after the fact. To better understand how to manage VTC, we gathered managers, scientists, and practitioners from across the southwestern US to collect their experiences with VTC challenges, management responses, and outcomes. Results Participants in two workshops provided 11 descriptive case studies and 61 examples of VTC from their own field observations. These experiences demonstrate the extent and complexity of ecological reorganization across the region. High-severity fire was the predominant driver of VTC in semi-arid coniferous forests. By a large margin, these forests converted to shrubland, with fewer conversions to native or non-native herbaceous communities. Chaparral and sagebrush areas nearly always converted to non-native grasses through interactions among land use, climate, and fire. Management interventions in VTC areas most often attempted to reverse changes, although we found that these efforts cover only a small portion of high-severity burn areas undergoing VTC. Some areas incurred long (>10 years) observational periods prior to initiating interventions. Efforts to facilitate VTC were rare, but could cover large spatial areas. Conclusions Our findings underscore that type conversion is a common outcome of high-severity wildland fire in the southwestern US. Ecosystem managers are frontline observers of these far-reaching and potentially persistent changes, making their experiences valuable in further developing intervention strategies and research agendas. As its drivers increase with climate change, VTC appears increasingly likely in many ecological contexts and may require management paradigms to transition as well. Approaches to VTC potentially include developing new models of desired conditions, the use of experimentation by managers, and broader implementation of adaptive management strategies. Continuing to support and develop science-manager partnerships and peer learning groups will help to shape our response to ongoing rapid ecological transformations.

Significance Forests are experiencing growing risks of drought-induced mortality in a warming world. Yet, ecosystem dynamics following drought mortality remain unknown, representing a major limitation to our understanding of the ecological consequences of climate change. We provide an emerging picture of postdrought ecological trajectories based on field indicators of forest dynamics. Replacement patterns following mortality indicate limited short-term persistence of predrought dominant tree species, highlighting the potential for major ecosystem reorganization in the coming decades. The great variability of the observed dynamics within and among species reinforces the primary influence of drought characteristics and ecosystem legacies, modulated by land use, management, and past disturbances, on ongoing drought-related species turnover and their potential implications for future forest biodiversity and ecosystem services., Forest vulnerability to drought is expected to increase under anthropogenic climate change, and drought-induced mortality and community dynamics following drought have major ecological and societal impacts. Here, we show that tree mortality concomitant with drought has led to short-term (mean 5 y, range 1 to 23 y after mortality) vegetation-type conversion in multiple biomes across the world (131 sites). Self-replacement of the dominant tree species was only prevalent in 21% of the examined cases and forests and woodlands shifted to nonwoody vegetation in 10% of them. The ultimate temporal persistence of such changes remains unknown but, given the key role of biological legacies in long-term ecological succession, this emerging picture of postdrought ecological trajectories highlights the potential for major ecosystem reorganization in the coming decades. Community changes were less pronounced under wetter postmortality conditions. Replacement was also influenced by management intensity, and postdrought shrub dominance was higher when pathogens acted as codrivers of tree mortality. Early change in community composition indicates that forests dominated by mesic species generally shifted toward more xeric communities, with replacing tree and shrub species exhibiting drier bioclimatic optima and distribution ranges. However, shifts toward more mesic communities also occurred and multiple pathways of forest replacement were observed for some species. Drought characteristics, species-specific environmental preferences, plant traits, and ecosystem legacies govern postdrought species turnover and subsequent ecological trajectories, with potential far-reaching implications for forest biodiversity and ecosystem services.

Changing disturbance regimes and climate can overcome forest ecosystem resilience. Following high-severity fire, forest recovery may be compromised by lack of tree seed sources, warmer and drier postfire climate, or short-interval reburning. A potential outcome of the loss of resilience is the conversion of the prefire forest to a different forest type or nonforest vegetation. Conversion implies major, extensive, and enduring changes in dominant species, life forms, or functions, with impacts on ecosystem services. In the present article, we synthesize a growing body of evidence of fire-driven conversion and our understanding of its causes across western North America. We assess our capacity to predict conversion and highlight important uncertainties. Increasing forest vulnerability to changing fire activity and climate compels shifts in management approaches, and we propose key themes for applied research coproduced by scientists and managers to support decision-making in an era when the prefire forest may not return.

Recent, widespread spruce beetle (Dendroctonus rufipennis) outbreaks have driven extensive tree mortality across western North America. Post-disturbance forest management often includes salvage logging to capture economic value of dead timber, reduce fire hazard, and meet other social or ecological objectives. Little is known about effects of salvage logging on surface fuel loads or plant understory communities in Engelmann spruce (Picea engelmannii)-dominated forests. We sampled fine and coarse woody debris, ground cover, and plant species composition along transects in spruce-beetle impacted stands in southwestern Colorado, USA. Twenty stands had been subject to clearcut 1–2 years prior to sampling; 32 stands were unlogged controls. Salvage logged stands exhibited altered surface fuels, ground cover, plant species cover, and community composition. Salvage increased 1-, 10-, and 100-hr fuels, and cover by bare ground and woody debris; cover by litter and cryptogams was reduced. Understory plant cover was reduced in salvaged stands, primarily due to losses of shrub cover. We found no difference in species diversity or richness between treated stands and controls. Salvage logging also drove shifts in plant community composition. Mean cover by non-native species was low, and not different, between control and salvage stands. Our study characterizes short-term effects that will undoubtedly change substantially over longer periods, in particular due to anticipated tree seedling growth and movement of standing dead wood from the canopy to the surface. In the near term, abundant fine woody surface fuels at salvage sites could influence the likelihood and rate of spread of surface fires, though over time we expect surface fuels in untreated stands to increase to comparable or greater levels than in salvaged stands. Differences in vascular plant cover and composition imparted by salvage harvests is also expected to change over time, though whether treated and untreated communities diverge or converge is not known. As such, we recommend that salvage harvest effects be monitored over extended time frames in order to detect longer-term trends.

Fire regimes in many dry forests of western North America are substantially different from historical conditions, and there is concern about the ability of these forests to recover following severe wildfire. Fire refugia, unburned or low-severity burned patches where trees survived fire, may serve as essential propagule sources that enable forest regeneration. To quantify the influence of fire refugia spatial pattern and other biophysical factors on the process of post-fire tree regeneration; in particular examining both the proximity and density of surrounding refugia to characterize the landscape of refugial seed sources. We surveyed regeneration at 135 sites in stand-replacement patches across a gradient of fire refugia density in eastern Oregon, USA. We characterized the influence of refugial seed source pattern and other environmental factors on the abundance of regenerating seedlings, and examined the relationship between post-fire climate and the temporal pattern of ponderosa pine seedling establishment. Tree seedlings were present in 83% of plots 12–17 years post-fire, and densities varied substantially (0–67800 stems ha−1, median = 1100). Variation in seedling abundance was driven by the spatial patterns of refugial seed sources. Despite widespread post-fire shrub cover, high-severity burned forests have not undergone a persistent type conversion to shrublands. Ponderosa pine seedling establishment peaked 5–11 years after fire and was not closely associated with post-fire climate. Fire refugia and the seed sources they contain fostered tree regeneration in severely burned patches. Management practices that reduce refugia within post-fire landscapes may negatively influence essential forest recovery processes.

Author(s): Krawchuk, Meg A; Meigs, Garrett W; Cartwright, Jennifer M; Coop, Jonathan D; Davis, Raymond; Holz, Andres; Kolden, Crystal; Meddens, Arjan JH

The increasing incidence of wildfires across the southwestern United States (US) is altering the contemporary forest management template within historically frequent-fire conifer forests. An increasing fraction of southwestern conifer forests have recently burned, and many of these burned landscapes contain complex mosaics of surviving forest and severely burned patches without surviving conifer trees. These heterogeneous burned landscapes present unique social and ecological challenges. Severely burned patches can present numerous barriers to successful conifer regeneration, and often contain heavy downed fuels which have cascading effects on future fire behavior and conifer regeneration. Conversely, surviving forest patches are increasingly recognized for their value in postfire reforestation but often are overlooked from a management perspective. Here we present a decision-making framework for landscape-scale management of complex postfire landscapes that allows for adaptation to a warming climate and future fire. We focus specifically on historically frequent-fire forests of the southwestern US but make connections to other forest types and other regions. Our framework depends on a spatially-explicit assessment of the mosaic of conifer forest and severely burned patches in the postfire landscape, evaluates likely vegetation trajectories, and identifies critical decision points to direct vegetation change via manipulations of fuels and live vegetation. This framework includes detailed considerations for postfire fuels management (e.g., edge hardening within live forest patches and repeat burning) and for reforestation (e.g., balancing tradeoffs between intensive and extensive planting strategies, establishing patches of seed trees, spatial planning to optimize reforestation success, and improving nursery capacity). In a future of increasing fire activity in forests where repeated low- to moderate-severity fire is essential to ecosystem resilience, the decision-making framework developed here can easily be integrated with existing postfire management strategies to optimize allocation of limited resources and more actively manage burned landscapes.

Forest mastication treatments are increasingly utilized by land managers as a means of reducing tree cover for fire hazard mitigation and other habitat objectives in pinon-juniper (P-J) woodlands. Mastication converts trees into small pieces (e.g., wood chips), in the process creating canopy openings, redistributing fuel from the canopy to the surface, converting large diameter to small diameter fuels, and covering the ground with piles of woody debris. We measured vegetation and fuels at 192 sites in 24 pairs of 1–11-year-old mastication treatments and untreated adjacent controls in P-J woodlands of the Arkansas River valley, Colorado, and used paired t -tests, mixed-effects models, and gradient analysis (non-metric multidimensional scaling) to assess mastication effects. Treatments were associated with major, persistent ecological shifts relative to controls. Tree cover and canopy fuels were reduced in treatments; concomitantly, dead and down woody surface fuels, forb, and graminoid cover were elevated. Treatments exhibited much higher frequency, richness, and cover by a suite of non-native plant species including cheatgrass ( Bromus tectorum ). Non-native plant expansion appears linked to the disturbance associated with treatment activities, reductions in tree canopy, and alterations to ground cover, and effective mitigation of increases by these species may necessitate both pre- and post-treatment control measures. Shifts from native-dominated woodlands to open, weedy, herb- and shrub-dominated communities are likely to change patterns of abundance and habitat use by woodland- and forest-dependent wildlife. Decreased canopy fuels and increased herbaceous surface fuels including exotic annuals are expected to alter potential fire behavior. We encourage managers carrying out P-J mastication projects to explicitly address potential trade-offs between desired treatment outcomes and potentially unwelcome impacts, and how these might be mitigated. It may also be worth considering whether or not tree removal treatments will be warranted given anticipated climate change impacts to these woodlands.

Tree species are highly vulnerable to anthropogenic environmental change, and are increasingly being challenged by non-native pests and climate change. Rocky Mountain bristlecone pines are long-lived, exhibit delayed maturation, have low genetic diversity, and inhabit cold, high-elevation environments. They are threatened by the non-native disease white pine blister rust, warming temperatures, changing precipitation patterns, and altered disturbance regimes. The goal of this work was to (1) sample the range-wide genetic diversity of bristlecone pine for ex situ seed and tissue collections and research before the populations are impacted, (2) assess the health and ecological conditions of the species across its geographic range (61 stands) to provide a baseline by which to evaluate future changes, and (3) assess relationships between stand and regeneration characteristics and topographic, geographic, and climatic factors to identify vulnerabilities and proactive management interventions to increase population resilience. Local variation in topoclimate was strongly related to stand structure, composition, health, and regeneration. Bristlecone pine currently showed only minor impacts from insects and pathogens, but relationships between cone production and regeneration with climate variables suggest vulnerabilities to projected climate change. Both cone production and seedling regeneration were also linked to stand structure and ground cover that provide opportunities for proactive in situ management to increase population size to mitigate potential future climate- and pathogen-driven declines. These efforts represent the first proactive coordinated range-wide genetic collection design and forest health assessment for a threatened, but not yet impacted, tree species and offers a model for other species at risk.

Wildfires in forest ecosystems produce landscape mosaics that include relatively unaffected areas, termed fire refugia. These patches of persistent forest cover can support fire-sensitive species and the biotic legacies important for post-fire forest recovery, yet little is known about their abundance and distribution within fire perimeters. Readily accessible 30-m resolution satellite imagery and derived burn severity products are commonly employed to characterize post-fire landscapes, however, coarse image resolution, generalized burn severity thresholds, and other limitations can constrain accurate representation of fire refugia. This study quantifies the abundance and pattern of fire refugia within 10 fires occurring in ponderosa pine and dry mixed-conifer forests between 2000 and 2003. We developed high-resolution maps of post-fire landscapes using semi-automated, object-based classification of 1-m aerial imagery, conducted imagery- and field-based accuracy assessments, and contrasted these with Landsat-derived burn severity metrics. Fire refugia area within burn perimeters ranged from 20% to 57%. Refugia proportion generally decreased with increasing Landsat-derived burn severity, but still accounted for 3&ndash, 12% of areas classified as high severity. Patch size ranged from 1-m2 isolated trees to nearly 8000 ha, and median patch size was 0.01 ha&mdash, substantially smaller than a 30-m Landsat pixel. Patch size was negatively related to burn severity, distance to fire refugia from open areas was positively related to burn severity. Finally, optimized thresholds of 30-m post-fire normalized burn ratio (NBR) and relative differenced normalized burn ratio (RdNBR) delineated fire refugia with an accuracy of 77% when validated against the 1-m resolution maps. Estimations of fire refugia abundance based on Landsat-derived burn severity metrics are unlikely to detect small, isolated fire refugia patches. Finer-resolution maps can improve understanding of the distribution of forest legacies and inform post-fire management activities including reforestation and treatments.

Altered fire regimes can drive major and enduring compositional shifts or losses of forest ecosystems. In western North America, ponderosa pine and dry mixed‐conifer forest types appear increasingly vulnerable to uncharacteristically extensive, high‐severity wildfire. However, unburned or only lightly impacted forest stands that persist within burn mosaics—termed fire refugia—may serve as tree seed sources and promote landscape recovery. We sampled tree regeneration along gradients of fire refugia proximity and density at 686 sites within the perimeters of 12 large wildfires that occurred between 2000 and 2005 in the interior western United States. We used generalized linear mixed‐effects models to elucidate statistical relationships between tree regeneration and refugia pattern, including a new metric that incorporates patch proximity and proportional abundance. These relationships were then used to develop a spatially explicit landscape simulation model. We found that regeneration by ponderosa pine and obligate‐seeding mixed‐conifer tree species assemblages was strongly and positively predicted by refugia proximity and density. Simulation models revealed that for any given proportion of the landscape occupied by refugia, small patches produced greater landscape recovery than large patches. These results highlight the disproportionate importance of small, isolated islands of surviving trees, which may not be detectable with coarse‐scale satellite imagery. Findings also illustrate the interplay between patch‐scale resistance and landscape‐scale resilience: Disturbance‐resistant settings (fire refugia) can entrain resilience (forest regeneration) across the burn matrix. Implications and applications for land managers and conservation practitioners include strategies for the promotion and maintenance of fire refugia as components of resilient forest landscapes.

Natural resource managers are increasingly applying tree reduction treatments to piñon–juniper woodlands to meet a range of ecological, social, and economic goals. However, treatment effects on woodland-obligate bird species are not well understood. We measured multiscale avian occupancy on 29 paired (control/treatment) sites in piñon–juniper woodlands in central Colorado, USA. We conducted point counts at 232 stations, 3 times each season in 2014 and 2015. We used hierarchical multiscale modeling to obtain unbiased estimates of landscape and local occupancy (i.e. probability of use) in treated and untreated sites for 31 species. Treatments reduced the occupancy of conifer obligates, including Mountain Chickadee (Poecile gambeli), Clark’s Nutcracker (Nucifraga columbiana), and White-breasted Nuthatch (Sitta carolinensis), and increased occupancy of Lark Sparrow (Chondestes grammacus) and Mountain Bluebird (Sialia currucoides). Occupancy of Virginia’s Warbler (Oreothylpis virginiae) and Gray Flycatcher (Empidonax wrightii), two piñon–juniper specialists, decreased at the landscape scale in treated sites, and Pinyon Jay (Gymnorhinus cyanocephalus) occupancy decreased at the local scale. Tree reduction treatments in piñon–juniper woodlands have the potential to reduce habitat quality for a suite of bird species of conservation concern. We suggest that treatments designed to retain higher tree density and basal area will benefit conifer-obligate and piñon–juniper specialist bird species.

Aim Studies of fire activity along environmental gradients have been undertaken, but the results of such studies have yet to be integrated with fire-regime analysis. We characterize fire-regime components along climate gradients and a gradient of human influence. Location We focus on a climatically diverse region of north-western North America extending from northern British Columbia, Canada, to northern Utah and Colorado, USA. Methods We used a multivariate framework to collapse 12 climatic variables into two major climate gradients and binned them into 73 discrete climate domains. We examined variation in fire-regime components (frequency, size, severity, seasonality and cause) across climate domains. Fire-regime attributes were compiled from existing databases and Landsat imagery for 1897 large fires. Relationships among the fire-regime components, climate gradients and human influence were examined through bivariate regressions. The unique contribution of human influence was also assessed. Results A primary climate gradient of temperature and summer precipitation and a secondary gradient of continentality and winter precipitation in the study area were identified. Fire occupied a distinct central region of such climate space, within which fire-regime components varied considerably. We identified significant interrelations between fire-regime components of fire size, frequency, burn severity and cause. The influence of humans was apparent in patterns of burn severity and ignition cause. Main conclusions Wildfire activity is highest where thermal and moisture gradients converge to promote fuel production, flammability and ignitions. Having linked fire-regime components to large-scale climate gradients, we show that fire regimes – like the climate that controls them – are a part of a continuum, expanding on models of varying constraints on fire activity. The observed relationships between fire-regime components, together with the distinct role of climatic and human influences, generate variation in biotic communities. Thus, future changes to climate may lead to ecological changes through altered fire regimes.

Aspen stands in the Rocky Mountains form especially productive and diverse plant understory communities. However, little is known about canopy–understory relationships or understory dynamics, especially in light of widespread aspen decline. The purpose of our research was to assess recent aspen plant community dynamics by resampling 19 sites sampled in 1964 and 1994 near Crested Butte, Colorado. In 2010, we replicated previous sampling methods to measure canopy structure and understory composition at each site. In contrast with the conclusions of previous work in the study area, sampled aspen stands did not exhibit long-term stability, nor were they succeeding to conifers. Over the 46-year sample period, live aspen density and basal area both decreased significantly (3151–1605 stems ha−1; 39.2–33.7 m2 ha−1), with more pronounced decreases over the most recent sampling interval, 1994–2010. These changes appear linked to natural patterns of stand maturation, but may also have been enhanced by recent, drought-triggered mortality, as well as constraints on aspen regeneration imposed by elk and livestock. A striking decrease in the cover and frequency of fireweed (Chamerion angustifolium) was consistent with expectations of aging stands initiated by historic fire, and the expansion of Kentucky bluegrass (Poa pratensis) indicated strong effects of ungulate herbivory. Recent overstory losses had little effect on understory communities, which were dominated by a suite of large perennial herbs that showed little change over the sampling interval, including Fendler’s meadowrue (Thalictrum fendleri), osha (Ligusticum porteri), elk and dryspike sedge (Carex geyeri and C. siccata), and blue wildrye (Elymus glaucus). Understory community composition was not closely related to current stand conditions or light levels, but was linked with regional elevational and geographic gradients and historic (1994) stand structure.

The increasing frequency and severity of wildfires in semi-arid conifer forests as a result of global change pressures has raised concern over potential impacts on biodiversity. Ground-dwelling arthropod communities represent a substantial portion of diversity in conifer forests, and could be particularly impacted by wildfires. In addition to direct mortality, wildfires can affect ground-dwelling arthropods by altering understory characteristics and associated deterministic community assembly processes (e.g., environmental sorting). Alternatively, disturbances have been reported to increase the importance of stochastic community assembly processes (e.g., probabilistic dispersal and colonization rates). Utilizing pitfall traps to capture ground-dwelling arthropods within forest stands that were burned by one or two wildfires since 1996 in the Jemez Mountains of northern New Mexico, United States (USA), we examined the potential influences of deterministic versus stochastic processes on the assembly of these diverse understory communities. Based on family-level and genera-level arthropod identifications, we found that the multivariate community structures differed among the four fire groups surveyed, and were significantly influenced by the quantities of duff, litter, and coarse woody debris, in addition to tree basal area and graminoid cover. Taxon diversity was positively related to duff quantities, while taxon turnover was positively linked to exposed-rock cover and the number of logs on the ground. Despite the significant effects of these understory properties on the arthropod community structure, a combination of null modeling and metacommunity analysis revealed that both deterministic and stochastic processes shape the ground-dwelling arthropod communities in this system. However, the relative influence of these processes as a function of time since the wildfires or the number of recent wildfires was not generalizable across the fire groups. Given that different assembly processes shaped arthropod communities among locations that had experienced similar disturbances over time, increased efforts to understand the processes governing arthropod community assembly following disturbance is required in this wildfire-prone landscape.

Large and severe wildfires have raised concerns about the future of forested landscapes in the southwestern United States, especially under repeated burning. In 2011, under extreme weather and drought conditions, the Las Conchas fire burned over several previous burns as well as forests not recently exposed to fire. Our purpose was to examine the influences of prior wildfires on plant community composition and structure, subsequent burn severity, and vegetation response. To assess these relationships, we used satellite-derived measures of burn severity and a nonmetric multidimensional scaling of pre- and post- Las Conchas field samples. Earlier burns were associated with shifts from forested sites to open savannas and meadows, oak scrub, and ruderal communities. These non-forested vegetation types exhibited both resistance to subsequent fire, measured by reduced burn severity, and resilience to reburning, measured by vegetation recovery relative to forests not exposed to recent prior fire. Previous shifts toward non-forested states were strongly reinforced by reburning. Ongoing losses of forests and their ecological values confirm the need for restoration interventions. However, given future wildfire and climate projections, there may also be opportunities presented by transformations toward fire-resistant and resilient vegetation types within portions of the landscape.

Objective: Our purpose was to characterize vegetation compositional patterns, tree regeneration, and plant diversity, and their relationships to landscape context, topography, and light availability across the margins of four stand-replacing subalpine burns. Location: Four 1977 to 1978 burns east of the Continental Divide in Colorado: the Ouzel burn, a burn near Kenosha Pass, the Badger Mountain burn, and the Maes Creek burn. Methods: Vegetation and environmental factors were sampled in 200 0.01-ha plots on transects crossing burn edges, and stratified by elevation. We utilized dissimilarity indices, mixed-effects models, and randomization tests to assess relationships between vegetation and environment. Results: Three decades after wildfire, plant communities exhibited pronounced compositional shifts across burn edges. Tree regeneration decreased with increasing elevation and distance into burn interiors; concomitant increases in forbs and graminoids were linked to greater light availability. Richness was roughly doubled in high-severity burn interiors due to the persistence of a suite of native species occurring primarily in this habitat. Richness rose with distance into burns, but declined with increasing elevation. Only three of 188 plant species were non-native; these were widespread, naturalized species that comprised o1% total cover. Conclusions: These subalpine wildfires generated considerable, persistent increases in plant species richness at local and landscape scales, and a diversity of plant communities. The findings suggest that fire suppression in such systems must lead to reduced diversity. Concerns about post-fire invasion by exotic plants appear unwarranted in high-elevation wilderness settings.

Rocky Mountain bristlecone pine ( Pinus aristata ) and limber pine ( Pinus flexilis ) are important high-elevation pines of the southern Rockies that are forecast to decline due to the recent spread of white pine blister rust ( Cronartium ribicola ) into this region. Proactive management strategies to promote the evolution of rust resistance and maintain ecosystem function require an improved understanding of the role of disturbance on the population dynamics of both species and environmental conditions that favor seedling establishment. We examined patterns of bristlecone and limber pine regeneration across the perimeters of three, 29-year-old, high-severity burns in northern, central, and southern Colorado: Ouzel, Badger Mountain, and Maes Creek, respectively. Both species exhibited a very protracted regeneration response to these fires. Bristlecone pine regeneration was concentrated near burn edges and beneath surviving seed sources. This spatial pattern is consistent with limitations incurred by wind-dispersal, also borne out by the low occurrence of seedling clusters. Relative to unburned stands, the absolute abundance of bristlecone pine generally increased only on plots retaining some surviving trees. Limber pine regeneration pattern varied between sites: high in the burn interior at Ouzel, concentrated at burn edges at Badger, and mostly in unburned stands at Maes. Clark’s Nutcracker dispersal of limber pine in each study area was indicated by high seedling distance from possible seed sources and high frequencies of clustered stems. Except at Ouzel, the absolute abundance of limber pine decreased in burns. Across sites, establishment by both species was boosted by nearby nurse objects (rocks, fallen logs, and standing tree trunks), a relationship that extended out at least as far as the closest three such objects, usually found within 50 cm. Fire decreased the frequency of Pedicularis but increased Castilleja and Ribes species (alternate hosts of white pine blister rust), though only one species, R. cereum , was positively associated with either pine species. We conclude that regeneration of bristlecone and limber pine may benefit from natural disturbance or proactive management creating appropriately sized openings and microtopographic structure (e.g., abundant fallen logs); however, beneficial responses may require many decades to be achieved.

Montane and subalpine grasslands are prominent, but poorly understood, features of the Rocky Mountains. These communities frequently occur below reversed tree lines on valley floors, where nightly cold air accumulation is spatially coupled with fine soil texture. We used field experiments to assess the roles of minimum temperature, soil texture, grass competition, and ungulate browsing on the growth, photosynthetic performance, and survival of transplanted ponderosa pine (Pinus ponderosa) seedlings at 32 sites straddling such reversed tree lines in the Valles Caldera National Preserve (VCNP) of the Jemez Mountains, New Mexico (USA). Seedling growth increased most strongly with increasing nighttime minimum temperatures away from the valley bottoms; seedlings experiencing the coldest temperatures on the caldera floor exhibited stunted needles and often no measurable height growth. Based on the chlorophyll fluorescence ratios PhiPSII and Fv/Fm, we found that low minimum temperatures, low soil moisture, and fine soil texture all contributed to photoinhibition. Neighboring herbs had only minor negative effects on seedlings. We found no effect of ungulates, but golden-mantled ground squirrels (Spermophilus lateralis) caused substantial seedling mortality. Second-year seedling survival was highest on sandy soils, and third-year survival was highest at sites with higher minimum temperatures. We conclude that differential tree seedling establishment driven by low minimum temperatures in the valley bottoms is the primary factor maintaining montane grasslands of the VCNP, although this process probably operated historically in combination with frequent surface fire to set the position of the tree line ecotone. As at alpine tree lines, reversed tree lines bordering montane and subalpine grasslands can represent temperature-sensitive boundaries of the tree life form.

Aim Recent forest encroachment into montane and subalpine grasslands has occurred in the Rocky Mountains and many other mountain ranges globally. The timing, rate, and extent of tree invasion can depend on interactions among topography, positive spatial feedbacks, and temporally variable factors (especially climate, grazing, and fire). Here we examine spatial and temporal patterns of tree invasion in the Valles Caldera of the Jemez Mountains. Location This study was conducted in the Valles Caldera (35°50′–36°00′ N; 106°24′–106°37′ W), a 24-km-wide volcanic basin in northern New Mexico, USA. Grasslands in this otherwise forested region occur in broad valley bottoms of the caldera floor between 2575 and 2700 m, and on south-facing slopes and mountain tops up to 3300 m. Methods We used a GIS analysis of orthorectified aerial photos taken in 1935 and 1996, covering a 40,000-ha study area, to quantify the extent of tree invasion and to assess its relationship to spatial factors. We obtained dates of establishment from 299 increment cores and basal disks from 50 sites in the Valles Caldera National Preserve (VCNP) to reconstruct temporal patterns of tree invasion. Results The area of grasslands in our study area declined from 11,747 to 9336 ha (nearly 18%) between 1935 and 1996. Tree invasion increased with slope, elevation, and proximity to the previous tree line, but showed no relationship to aspect. Tree invasion was more rapid and continuous on upper mountain slopes, while the invasion of valley-bottom grasslands below reversed tree lines was more episodic, and appeared to track mean summer minimum temperatures. Main conclusions The rapid and continuous invasion of steep, high-elevation slopes suggests that frequent fire was the single most important factor in maintaining grassy communities in these sites. The slower, episodic invasion of valley-bottom grasslands, and the apparent relationship between increased invasion and years of higher summer minimum temperatures are consistent with the hypothesis that these grasslands have been maintained by low temperatures or frosts damaging to tree seedlings. We encourage prescribed fire to restore and maintain grasslands in the VCNP, especially small patches on steep, high-elevation slopes.

We observed black bears (Ursus americanus) foraging on aggregations of army cutworm moths (Euxoa auxiliaris) (Lepidoptera: Noctuidae) in subalpine felsenmeers (block fields) in the Jemez Mountains of New Mexico. Moth aggregations serve as food for grizzly bears (Ursus arctos horribilis) in the northern Rocky Mountains of Wyoming and Montana. However, black bears have not been reported to use these aggregations, nor have such aggregations been documented to occur this far south in the Rocky Mountains.