Whitebark Pine Research Articles

Leaf Essential Oil Compositions and Enantiomeric Distributions of Monoterpenoids in Pinus Species: Pinus albicaulis, Pinus flexilis, Pinus lambertiana, Pinus monticola, and Pinus sabiniana.

Members of the Pinus genus are well known for their medicinal properties, which can be attributed to their essential oils. In this work, we have examined the leaf essential oils of five understudied Pinus species collected from various locations in western North America. The essential oils were obtained by hydrodistillation and analyzed by gas chromatographic methods, including enantioselective gas chromatography. Pinus albicaulis was dominated by (+)-δ-3-carene; Pinus flexilis was dominated by α-pinene (mostly (+)-α-pinene) and (-)-β-pinene; Pinus lambertiana was dominated by (-)-β-pinene; Pinus monticola was dominated by (-)-β-pinene, (+)-δ-3-carene, and (-)-α-pinene; and Pinus sabiniana was rich in (-)-α-pinene and limonene. While this work adds to our knowledge of Pinus essential oils, additional research is needed to more fully appreciate the geographic and altitudinal variations in the volatile compositions of these Pinus species.

Dynamic treeline and cryosphere response to pronounced mid-Holocene climatic variability in the US Rocky Mountains

Climate-driven changes in high-elevation forest distribution and reductions in snow and ice cover have major implications for ecosystems and global water security. In the Greater Yellowstone Ecosystem of the Rocky Mountains (United States), recent melting of a high-elevation (3,091 m asl) ice patch exposed a mature stand of whitebark pine (Pinus albicaulis) trees, located ~180 m in elevation above modern treeline, that date to the mid-Holocene (c. 5,950 to 5,440 cal y BP). Here, we used this subfossil wood record to develop tree-ring-based temperature estimates for the upper-elevation climate conditions that resulted in ancient forest establishment and growth and the subsequent regional ice-patch growth and downslope shift of treeline. Results suggest that mid-Holocene forest establishment and growth occurred under warm-season (May-Oct) mean temperatures of 6.2 °C (±0.2 °C), until a multicentury cooling anomaly suppressed temperatures below 5.8 °C, resulting in stand mortality by c. 5,440 y BP. Transient climate model simulations indicate that regional cooling was driven by changes in summer insolation and Northern Hemisphere volcanism. The initial cooling event was followed centuries later (c. 5,100 y BP) by sustained Icelandic volcanic eruptions that forced a centennial-scale 1.0 °C summer cooling anomaly and led to rapid ice-patch growth and preservation of the trees. With recent warming (c. 2000-2020 CE), warm-season temperatures now equal and will soon exceed those of the mid-Holocene period of high treeline. It is likely that perennial ice cover will again disappear from the region, and treeline may expand upslope so long as plant-available moisture and disturbance are not limiting.

Climate Dominates Geographic Influences on Whitebark Pine and Limber Pine Trends and Landscape Patterns in Canada

Climate Dominates Geographic Influences on Whitebark Pine and Limber Pine Trends and Landscape Patterns in Canada

Thinning enhances whitebark pine (Pinus albicaulis) growth and resin duct defenses

AbstractWhitebark pine (Pinus albicaulis) is an important component of high‐elevation ecosystems throughout the northern Rocky Mountains; however, the species was recently listed as “threatened” under the Endangered Species Act due to widespread mortality throughout its native range. Land managers tasked with whitebark pine conservation and restoration require information on strategies that can positively influence whitebark pine establishment and promote growth and defense of preexisting trees on the landscape. Our study investigated the response of whitebark pine tree growth and resin duct defenses to thinning and removal of competing shade‐tolerant conifers in the Caribou‐Targhee National Forest, Idaho, USA. In the nine years following thinning treatments, we found that whitebark pine trees (>12.7 cm diameter at breast height [dbh]) responded favorably to thinning and experienced a 60% increase in basal area increment (52% increase in ring width index) while also producing 34% more resin ducts. Additionally, resin ducts were 23% larger with 49% increased area in the nine years post‐thinning compared with trees in the control. Small diameter (<5 cm dbh) whitebark pine in the treatment experienced a dramatic growth release with over 90% increase in basal area increment (74% increase in ring width index). Thinning also created conditions favorable for whitebark pine seedling and sapling establishment. In the treatment, 75% of seedlings and 93% of saplings were whitebark pine, compared with 32% seedlings and 14% saplings in the control. In addition, 93% of overstory trees in the treatment were whitebark pine, compared with 42% in the control. Whitebark pine tree mortality was similar across both the control (26%) and the treatment (25%), indicating that the thinning treatments did not have any adverse impacts on tree mortality. The positive effects of thinning on whitebark pine establishment and tree growth, coupled with the lack of elevated mortality in the treatment, provide strong evidence in support of thinning as a viable silvicultural technique in whitebark pine conservation efforts.

Using Paleoecological Methods to Study Long-Term Disturbance Patterns in High-Elevation Whitebark Pine Ecosystems

Pinus albicaulis (whitebark pine) is a keystone species, providing food and habitat to wildlife, in high-elevation ecological communities. In recent years, this important species has been negatively impacted by changes in fire regimes, increased Dendroctonus ponderosae (mountain pine beetle) outbreaks associated with human landscape and climate modification, and the continued impact of the non-native Cronartium ribicola (white pine blister rust). This research investigates changes in fire occurrence, the establishment of Pinus albicaulis, and fuel availability at a high-elevation site in the Sawtooth National Recreation Area, Idaho, USA. Charcoal and pollen analyses were used to reconstruct fire and vegetation patterns for Phyllis Lake, Idaho, USA, over the past ~8200 cal y BP. We found that significant fire episodes occurred when the pollen accumulation rates (PARs) indicated more arboreal fuel availability, and we identified that Pinus albicaulis became well established at the site ~7200 cal y BP. The high-elevation nature of Phyllis Lake (2800 m) makes this record unique, as there are not many paleorecords at this high elevation from the Northern Rocky Mountains, USA. Additional high-elevation sites in Pinus albicaulis habitats will provide critical insight into the long-term dynamics of this threatened species.

Study of the Genetic Mechanisms of Siberian Stone Pine (Pinus sibirica Du Tour) Adaptation to the Climatic and Pest Outbreak Stresses Using Dendrogenomic Approach.

A joint analysis of dendrochronological and genomic data was performed to identify genetic mechanisms of adaptation and assess the adaptive genetic potential of Siberian stone pine (Pinus sibirica Du Tour) populations. The data obtained are necessary for predicting the effect of climate change and mitigating its negative consequences. Presented are the results of an association analysis of the variation of 84,853 genetic markers (single nucleotide polymorphisms-SNPs) obtained by double digest restriction-site associated DNA sequencing (ddRADseq) and 110 individual phenotypic traits, including dendrophenotypes based on the dynamics of tree-ring widths (TRWs) of 234 individual trees in six natural populations of Siberian stone pine, which have a history of extreme climatic stresses (e.g., droughts) and outbreaks of defoliators (e.g., pine sawfly [Neodiprion sertifer Geoff.]). The genetic structure of studied populations was relatively weak; samples are poorly differentiated and belong to genetically similar populations. Genotype-dendrophenotype associations were analyzed using three different approaches and corresponding models: General Linear Model (GLM), Bayesian Sparse Linear Mixed Model (BSLMM), and Bayesian-information and Linkage-disequilibrium Iteratively Nested Keyway (BLINK), respectively. Thirty SNPs were detected by at least two different approaches, and two SNPs by all three. In addition, three SNPs associated with mean values of recovery dendrophenotype (Rc) averaged across multiple years of climatic stresses were also found by all three methods. The sequences containing these SNPs were annotated using genome annotation of a very closely related species, whitebark pine (P. albicaulis Engelm.). We found that most of the SNPs with supposedly adaptive variation were located in intergenic regions. Three dendrophenotype-associated SNPs were located within the 10 Kbp regions and one in the intron of the genes encoding proteins that play a crucial role in ensuring the integrity of the plant's genetic information, particularly under environmental stress conditions that can induce DNA damage. In addition, we found a correlation of individual heterozygosity with some dendrophenotypes. Heterosis was observed in most of these statistically significant cases; signs of homeostasis were also detected. Although most of the identified SNPs were not assigned to a particular gene, their high polymorphism and association with adaptive traits likely indicate high adaptive potential that can facilitate adaptation of Siberian stone pine populations to the climatic stresses and climate change.

Accuracy of Whitebark Pine and Limber Pine Identification by Forest Inventory and Analysis Field Crews

Abstract Accurate identification of whitebark and limber pine has become increasingly important following the 2022 listing of whitebark pine as a threatened species under the Endangered Species Act. However, morphological similarities make identification of the two species difficult where ranges overlap. Using a genetic test that differentiates whitebark and limber pine, we compared field identification by Forest Inventory and Analysis field crews with genetic identification for needle samples from 371 trees. Field identifications were 100% correct for the 76 samples collected from outside regions of species’ range overlap. A total of 83% of the field identifications were correct in regions of range overlap (89% for large trees, 88% for saplings, and 78% for seedlings). Field-identified samples were correct 60% of the time for limber pine and >99% for whitebark pine. Random forests analysis revealed that identification accuracy is influenced by crew experience, large (≥ 12.7cm diameter) limber or whitebark pines recorded by field crews on the plot, elevation, Julian day of sample collection, and habitat type. We found that whitebark pine has likely been underestimated, and limber pine overestimated, within their overlapping ranges. We provide insights on improving accuracy of future monitoring where these species overlap. Study Implications: Accurate identification of whitebark pine is critical for monitoring this threatened species, yet distinguishing whitebark from limber pine can be difficult. Genetic analysis determined accuracy of field identification by Forest Inventory and Analysis (FIA) crews was 83% where the species’ ranges overlap. Virtually all individuals identified as whitebark pine were genetically confirmed to be whitebark pine, although nearly 40% of individuals identified as limber pine were actually whitebark pine. Thus, previous data underestimated whitebark and overestimated limber pine abundance in the species’ range overlap. These results quantify reliability of FIA data for whitebark pine assessments and identify areas for improvement.

Contrasting demographic history and mutational load in three threatened whitebark pines (Pinus subsect. Gerardianae): implications for conservation.

Demographic history and mutational load are of paramount importance for the adaptation of the endangered species. However, the effects of population evolutionary history and genetic load on the adaptive potential in endangered conifers remain unclear. Here, using population transcriptome sequencing, whole chloroplast genomes and mitochondrial DNA markers, combined with niche analysis, we determined the demographic history and mutational load for three threatened whitebark pines having different endangered statuses, Pinus bungeana, P. gerardiana and P. squamata. Demographic inference indicated that severe bottlenecks occurred in all three pines at different times, coinciding with periods of major climate and geological changes; in contrast, while P. bungeana experienced a recent population expansion, P. gerardiana and P. squamata maintained small population sizes after bottlenecking. Abundant homozygous-derived variants accumulated in the three pines, particularly in P. squamata, while the species with most heterozygous variants was P. gerardiana. Abundant moderately and few highly deleterious variants accumulated in the pine species that have experienced the most severe demographic bottlenecks (P. gerardiana and P. squamata), most likely because of purging effects. Finally, niche modeling showed that the distribution of P. bungeana might experience a significant expansion in the future, and the species' identified genetic clusters are also supported by differences in the ecological niche. The integration of genomic, demographic and niche data has allowed us to prove that the three threatened pines have contrasting patterns of demographic history and mutational load, which may have important implications in their adaptive potential and thus are also key for informing conservation planning.

Can climate variability and landscape position predict white pine blister rust incidence, mortality due to the disease, and regeneration in whitebark pine?

Invasive forest pathogens, like Cronartium ribicola (Fisch), the fungus that causes white pine blister rust, threaten native tree species. Federally listed whitebark pine ( Pinus albicaulis Engelmann) is highly susceptible and faces extensive mortality due to this disease. Understanding infection conditions and disease incidence variability is crucial for management and restoration efforts. We surveyed whitebark pine stands in Glacier National Park, Montana, USA from 2020 to 2022, examining the impact of landform and climate on tree mortality, rust incidence, cone production, and regeneration. Our analysis revealed that tree diameter, elevation, and aspect significantly influenced mortality, rust incidence, and cone production, while climatic factors such as spring solar radiation, humidity, and late summer snowpack also played key roles. Regeneration was primarily affected by elevation, geographic location, and humidity. Although landform variables similarly predicted disease incidence in this and other studies, climatic drivers varied by region, emphasizing the need to consider region-specific landform and climate for effective management. Our study highlights the importance of protecting large trees, which harbor genetic diversity crucial for recovery and adaptation to disturbance and climate change.

Transcriptional Profiling of Early Defense Response to White Pine Blister Rust Infection in Pinus albicaulis (Whitebark Pine).

Pathogen perception generates the activation of signal transduction cascades to host defense. White pine blister rust (WPBR) is caused by Cronartium ribicola J.C. Fisch and affects a number of species of Pinus. One of the most severely affected species is Pinus albicaulis Engelm (whitebark pine). WPBR resistance in the species is a polygenic and complex trait that requires an optimized immune response. We identified early responses in 2-year-old seedlings after four days of fungal inoculation and compared the underlying transcriptomic response with that of healthy non-inoculated individuals. A de novo transcriptome assembly was constructed with 56,796 high quality-annotations derived from the needles of susceptible and resistant individuals in a resistant half-sib family. Differential expression analysis identified 599 differentially expressed transcripts, from which 375 were upregulated and 224 were downregulated in the inoculated seedlings. These included components of the initial phase of active responses to abiotic factors and stress regulators, such as those involved in the first steps of flavonoid biosynthesis. Four days after the inoculation, infected individuals showed an overexpression of chitinases, reactive oxygen species (ROS) regulation signaling, and flavonoid intermediates. Our research sheds light on the first stage of infection and emergence of disease symptoms among whitebark pine seedlings. RNA sequencing (RNA-seq) data encoding hypersensitive response, cell wall modification, oxidative regulation signaling, programmed cell death, and plant innate immunity were differentially expressed during the defense response against C. ribicola.

Lethal combination for seedlings: extreme heat drives mortality of drought-exposed high-elevation pine seedlings.

Hotter drought- and biotically-driven tree mortality are expected to increase with climate change in much of the western United States, and species persistence will depend upon ongoing establishment under novel conditions or migration to track ecological niche requirements. High-elevation tree species may be particularly vulnerable to increasing water stress as snowpack declines, increasing the potential for adult mortality and simultaneous regeneration failures. Seedling survival will be determined by ecophysiological limitations in response to changing water availability and temperature. We exposed seedlings from populations of Pinus longaeva, Pinus flexilis, and Pinus albicaulis to severe drought and concurrent temperature stress in common gardens testing timing of drought onset under two different temperature regimes. We monitored seedling functional traits, physiological function, and survival. The combined stressors of water limitation and extreme heat led to conservative water use strategies and declines in physiological function, with these joint stressors ultimately exceeding species' tolerances and leading to complete episodic mortality across all species. Growing conditions were the primary determinant of seedling trait expression, with seedlings exhibiting more drought-resistant traits such as lower specific leaf area in the hottest, driest treatment conditions. Water stress-induced stomatal closure was also widely apparent. Under adequate soil moisture, seedlings endured prolonged exposure to high air and surface temperatures, suggesting broad margins for survival. The critical interaction between soil moisture and temperature suggests that rising temperatures will exacerbate growing season moisture stress. Our results highlight the importance of local conditions over population- and species-level influences in shaping strategies for stress tolerance and resistance to desiccation at this early life stage. By quantifying some of the physiological consequences of drought and heat that lead to seedling mortality, we can better understand the future effects of global change on the composition and distribution of high-elevation conifer forests.

A genome sequence for the threatened whitebark pine.

Whitebark pine (WBP, Pinus albicaulis) is a white pine of subalpine regions in the Western contiguous United States and Canada. WBP has become critically threatened throughout a significant part of its natural range due to mortality from the introduced fungal pathogen white pine blister rust (WPBR, Cronartium ribicola) and additional threats from mountain pine beetle (Dendroctonus ponderosae), wildfire, and maladaptation due to changing climate. Vast acreages of WBP have suffered nearly complete mortality. Genomic technologies can contribute to a faster, more cost-effective approach to the traditional practices of identifying disease-resistant, climate-adapted seed sources for restoration. With deep-coverage Illumina short reads of haploid megagametophyte tissue and Oxford Nanopore long reads of diploid needle tissue, followed by a hybrid, multistep assembly approach, we produced a final assembly containing 27.6 Gb of sequence in 92,740 contigs (N50 537,007 bp) and 34,716 scaffolds (N50 2.0 Gb). Approximately 87.2% (24.0 Gb) of total sequence was placed on the 12 WBP chromosomes. Annotation yielded 25,362 protein-coding genes, and over 77% of the genome was characterized as repeats. WBP has demonstrated the greatest variation in resistance to WPBR among the North American white pines. Candidate genes for quantitative resistance include disease resistance genes known as nucleotide-binding leucine-rich repeat receptors (NLRs). A combination of protein domain alignments and direct genome scanning was employed to fully describe the 3 subclasses of NLRs. Our high-quality reference sequence and annotation provide a marked improvement in NLR identification compared to previous assessments that leveraged de novo-assembled transcriptomes.

Increasing radial growth in old-growth high-elevation conifers in Southern California, USA, during the exceptional "hot drought" of 2000-2020.

Hot droughts, droughts attributed to below-average precipitation and exceptional warmth, are increasingly common in the twenty-first century, yet little is known about their effect on coniferous tree growth because of their historical rarity. In much of the American West, including California, radial tree growth is principally driven by precipitation, and narrow ring widths are typically associated with either drier or drought conditions. However, for species growing at high elevations (e.g., Larix lyalli, Pinus albicaulis), growth can be closely aligned with above-average temperatures with maximum growth coinciding with meteorological drought, suggesting that the growth effects of drought span from adverse to beneficial depending on location. Here, we compare radial growth responses of three high-elevation old-growth pines (Pinus jeffreyi, P. lambertiana, and P. contorta) growing in the San Jacinto Mountains, California, during a twenty-first-century hot drought (2000-2020) largely caused by exceptional warmth and a twentieth-century drought (1959-1966) principally driven by precipitation deficits. Mean radial growth during the hot drought was 12% above average while 18% below average during the mid-century drought illustrating that the consequences of environmental stress exhibit spatiotemporal variability. We conclude that the effects of hot droughts on tree growth in high-elevation forests may produce responses different than what is commonly associated with extended dry periods for much of western North America's forested lands at lower elevational ranges and likely applies to other mountainous regions (e.g., Mediterranean Europe) defined by summer-dry conditions. Thus, the climatological/biological interactions discovered in Southern California may offer clues to the unique nature of high-elevation forested ecosystems globally.

Paired acoustic recordings and point count surveys reveal Clark's nutcracker and whitebark pine associations across Glacier National Park.

Global declines in tree populations have led to dramatic shifts in forest ecosystem composition, biodiversity, and functioning. These changes have consequences for both forest plant and wildlife communities, particularly when declining species are involved in coevolved mutualisms. Whitebark pine (Pinus albicaulis) is a declining keystone species in western North American high-elevation ecosystems and an obligate mutualist of Clark's nutcracker (Nucifraga columbiana), an avian seed predator and disperser. By leveraging traditional point count surveys and passive acoustic monitoring, we investigated how stand characteristics of whitebark pine in a protected area (Glacier National Park, Montana, USA) influenced occupancy and vocal activity patterns in Clark's nutcracker. Using Bayesian spatial occupancy models and generalized linear mixed models, we found that habitat use of Clark's nutcracker was primarily supported by greater cone density and increasing diameter of live whitebark pine. Additionally, we demonstrated the value of performing parallel analyses with traditional point count surveys and passive acoustic monitoring to provide multiple lines of evidence for relationships between Clark's nutcracker and whitebark pine forest characteristics. Our findings allow managers to gauge the whitebark pine conditions important for retaining high nutcracker visitation and prioritize management efforts in whitebark pine ecosystems with low nutcracker visitation.

Field-testing whitebark pine resistance to white pine blister rust: A simple, effective approach to progeny testing for restoration

The endangered conifer whitebark pine (Pinus albicaulis), native to high elevation forests of western North America, is declining mainly due to the introduced pathogen Cronartium ribicola, causing the disease white pine blister rust. Low levels of genetic resistance to blister rust are present in whitebark pine populations. Traditional methods of inoculating seedlings to determine resistance to the rust are costly, labour intensive, and have limited capacity due to facilities. This presents a bottleneck in the development of resistant material for restoring stands that have been heavily infected by the rust. Here we test an alternative approach to controlled inoculations that could screen large numbers of families with lower costs and requiring fewer specialized resources. We screened 214 open-pollinated families from 44 provenances from British Columbia (BC), Washington (WA), Oregon (OR), Idaho (ID), Montana (MT), and Nevada (NV) to determine: (1) the effectiveness of natural rust inoculation from Ribes nigrum in a common garden; (2) family- and provenance-level resistance to blister rust; and (3) climate variables related to population-level rust resistance. Eighty-one of these families were previously screened by the US Forest Service in Cottage Grove, Oregon using standard artificial inoculation methods. The natural inoculation was effective, with 73 % of seedlings displaying stem symptoms of the disease, and 95 % showing rust infection. A clear relationship was found between distance from the nearest Ribes nigrum plant and severity of blister rust. Linear mixed models with spatial correlations were fitted to rust infection data using ASReml-R to estimate breeding values, heritability, and among-population differentiation (QST) for rust resistance. Population differentiation for rust resistance was moderate, with the highest resistance in provenances from the Cascade Mountains of Washington, and the most susceptible families from the BC Chilcotin and Okanagan regions. Heritability of rust resistance was moderate (0.23). The field testing was broadly consistent with classifications of families into resistant, moderate, and susceptible groups based on artificial inoculations. This method of screening could be used at a broader scale to determine families resistant to white pine blister rust without specialized testing facilities and increase the availability of resistant seedlings for restoration.

Clark's nutcracker forest community visitation: Whitebark pine maintains a keystone seed disperser.

Clark's nutcrackers (Nucifraga columbiana) are obligate seed dispersers for whitebark pine (Pinus albicaulis), but they frequently use other conifer seed resources because of annual variability in cone production or geographic variation in whitebark pine availability. Whitebark pine is declining from several threats including white pine blister rust, leading to potential population declines in the nutcracker and the pine. We hypothesize that where there are few additional seed resources, whitebark pine becomes the key and limiting resource supporting nutcracker populations. We investigated how nutcrackers use coniferous forest community types within Yellowstone National Park to determine potential seed resources and the importance of whitebark pine. We established sites representing five forest community types, including whitebark pine, lodgepole pine (P. contorta), Engelmann spruce (Picea engelmannii), limber pine (P. flexilis), and Douglas-fir (Pseudotsuga menziesii). Each transect annually generated nutcracker point counts, conifer cone production indices, community composition data, and seed resource use observations. We compared hierarchical distance sampling models, estimating nutcracker density and its relationship to forest community type, seed harvesting time-period, year, study site, and cone seed energy. We found cone production varied across years indicating annual variability in energy availability. Nutcracker density was best predicted by forest community type and survey time-period and was highest in whitebark pine stands during the mid-harvesting season. Nutcracker density was comparatively low for all other forest community types. This finding underscores the importance of whitebark pine as a key seed resource for Clark's nutcracker in Yellowstone National Park. The decline of whitebark pine potentially leads to a downward spiral in nutcrackers and whitebark pine, arguing for continued monitoring of nutcrackers and implementation of restoration treatments for whitebark pine.

Climate Change-Driven Cumulative Mountain Pine Beetle-Caused Whitebark Pine Mortality in the Greater Yellowstone Ecosystem

An aerial survey method called the Landscape Assessment System (LAS) was used to assess mountain pine beetle (Dendroctonus ponderosae)-caused mortality of whitebark pine (Pinus albicaulis) across the Greater Yellowstone Ecosystem (59,000 km2; GYE). This consisted of 11,942 km of flightlines, along which 4434 geo-tagged, oblique aerial photos were captured and processed. A mortality rating of none to severe (0–4.0 recent attack or 5.0–5.4 old attack) was assigned to each photo based on the amount of red (recent attack) and gray (old attack) trees visible. The method produced a photo inventory of 74 percent of the GYE whitebark pine distribution. For the remaining 26 percent of the distribution, mortality levels were estimated based on an interpolated mortality surface. Catchment-level results combining the photo-inventoried and interpolated mortality indicated that 44 percent of the GYE whitebark pine distribution showed severe old attack mortality (5.3–5.4 rating), 37 percent showed moderate old attack mortality (5.2–5.29 rating), 19 percent showed low old attack mortality (5.1–5.19 rating) and less than 1 percent showed trace levels of old attack mortality (5.0–5.09). No catchments were classified as recent attacks indicating that the outbreak of the early 2000’s has ended. However, mortality continues to occur as chronic sub-outbreak-level mortality. Ground verification using field plots indicates that higher LAS mortality values are moderately correlated with a higher percentage of mortality on the ground.

Disease resistance in whitebark pine and potential for restoration of a threatened species

Societal impact statementForests world‐wide are being negatively affected by non‐native, invasive pathogens and pests, and some tree species face uncertain futures. To retain these species as components of future forests, the rare genetic resistance that exists needs to be identified and harnessed. The applied tree improvement program for whitebark pine (Pinus albicaulis), a threatened (in the United States) and endangered (in Canada) keystone species in many forests in western North America, provides an example of what can be accomplished in a relatively short timeframe. The level and frequency of resistance vary by location, and this information will be used to implement the national restoration plan.Summary Forest trees face serious threats from non‐native diseases and pests, often causing high mortality of both the existing trees and regeneration. Developing populations with genetic resistance can help restore forests and retain affected species. Resistance programs have historically focused on species of high economic importance; however, the threats to species of little direct economic value that provide other important ecosystem services are also great. We examined the frequency, level, and geographic variation in genetic resistance to white pine blister rust in the threatened Pinus albicaulis (whitebark pine), a keystone species in high‐elevation ecosystems in western North America. In the two trials reported here, 2‐year‐old seedling progeny of 225 whitebark pine parent trees were inoculated with two geographic sources of the fungal pathogen Cronartium ribicola and evaluated over 5 years for an array of resistance traits. The trials focused primarily on parent trees from the Oregon and Washington populations. We found unexpectedly high levels of quantitative resistance in some seedling families and populations, in stark contrast to levels observed in similar resistance programs with other North American white pine species such as Pinus monticola and Pinus lambertiana. The level of resistance found in some whitebark pine populations provides optimism about potential recovery efforts for this species. Restoration efforts are underway by government agencies, tribes, and non‐government organizations in both the United States and Canada. These efforts may help boost support for applied genetic resistance programs in other forest tree species severely affected by non‐native pathogens or pests.

Snag-fall patterns following stand-replacing fire vary with stem characteristics and topography in subalpine forests of Greater Yellowstone

Standing dead tree stems (snags) become abundant following disturbances like bark beetle outbreaks and stand-replacing fire. Snags are an important element of wildlife habitat, and when they eventually fall can injure or damage people and infrastructure and contribute to coarse wood and fuels accumulation. While species-specific and general trends in snag persistence following disturbance have been well-studied, less attention has been paid to how these patterns vary across broad topographical gradients. We studied the ca. 250,000 ha of fire-killed snags created by the extensive and severe 1988 Yellowstone fires in the Greater Yellowstone Ecosystem (Wyoming, USA) and asked: (1) What characteristics of individual tree stems contribute to the likelihood and mode of snag-fall? (2) How do snag-fall patterns vary across broad topographical gradients? In 2002 and 2003 (14–15 years postfire) we determined the abundance of standing snags vs. fallen stems to identify patterns of snag persistence in relation to stem characteristics and topography. In 2022 (34 years postfire), we sampled a separate set of plots to determine which, if any, trends identified 14–15 years postfire persisted for another two decades. Trends in snag persistence were species specific, with lodgepole pine (Pinus contorta var. latifolia) stems less likely to remain standing than either Engelmann spruce (Picea engelmannii), Douglas-fir (Pseudotsuga menziesii var. glauca), subalpine fir (Abies lasiocarpa), or whitebark pine (Pinus albicaulis). Across all species, larger snags and those dead at the time of fire remained standing longer than smaller snags and those alive at the time of fire. The mode of snag-fall (uprooted vs. snapped) also varied with stem characteristics and species identity. Greater proportions of snags (all species and sizes combined) remained standing at higher elevations and on steeper slopes, likely driven by soils, stem allometry, decay rates, wind patterns, and pre-fire stand composition. As the extent and frequency of large, severe fires increases in western subalpine forests, understanding snag dynamics and the ecology and management of both standing and fallen snags is increasingly important.

Evidence for density-dependent effects on body composition of a large omnivore in a changing Greater Yellowstone Ecosystem.

Understanding the density-dependent processes that drive population demography in a changing world is critical in ecology, yet measuring performance-density relationships in long-lived mammalian species demands long-term data, limiting scientists' ability to observe such mechanisms. We tested performance-density relationships for an opportunistic omnivore, grizzly bears (Ursus arctos, Linnaeus, 1758) in the Greater Yellowstone Ecosystem, with estimates of body composition (lean body mass and percent body fat) serving as indicators of individual performance over two decades (2000-2020) during which time pronounced environmental changes have occurred. Several high-calorie foods for grizzly bears have mostly declined in recent decades (e.g., whitebark pine [Pinus albicaulis, Engelm, 1863]), while increasing human impacts from recreation, development, and long-term shifts in temperatures and precipitation are altering the ecosystem. We hypothesized that individual lean body mass declines as population density increases (H1), and that this effect would be more pronounced among growing individuals (H2). We also hypothesized that omnivory helps grizzly bears buffer energy intake from changing foods, with body fat levels being independent from population density and environmental changes (H3). Our analyses showed that individual lean body mass was negatively related to population density, particularly among growing-age females, supporting H1 and partially H2. In contrast, population density or sex had little effect on body fat levels and rate of accumulation, indicating that sufficient food resources were available on the landscape to accommodate successful use of shifting food sources, supporting H3. Our results offer important insights into ecological feedback mechanisms driving individual performances within a population undergoing demographic and ecosystem-level changes. However, synergistic effects of continued climate change and increased human impacts could lead to more extreme changes in food availability and affect observed population resilience mechanisms. Our findings underscore the importance of long-term studies in protected areas when investigating complex ecological relationships in an increasingly anthropogenic world.