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Understanding the factors influencing species range limits is increasingly crucial in anticipating migrations due to human‐caused climate change. In the boreal biome, ongoing climate change and the associated increases in the rate, size, and severity of disturbances may alter the distributions of boreal tree species. Notably, Interior Alaska lacks native pine, a biogeographical anomaly that carries implications for ecosystem structure and function. The current range of lodgepole pine (Pinus contorta var. latifolia) in the adjacent Yukon Territory may expand into Interior Alaska, particularly with human assistance. Evaluating the potential for pine expansion in Alaska requires testing constraints on range limits such as dispersal limitations, environmental tolerance limits, and positive or negative biotic interactions. In this study, we used field experiments with pine seeds and transplanted seedlings, complemented by model simulations, to assess the abiotic and biotic factors influencing lodgepole pine seedling establishment and growth after fire in Interior Alaska. We found that pine could successfully recruit, survive, grow, and reproduce across our broadly distributed network of experimental sites. Our results show that both mammalian herbivory and competition from native tree species are unlikely to constrain pine growth and that environmental conditions commonly found in Interior Alaska fall well within the tolerance limits for pine. If dispersal constraints are released, lodgepole pine could have a geographically expansive range in Alaska, and once established, its growth is sufficient to support pine‐dominated stands. Given the impacts of lodgepole pine on ecosystem processes such as increases in timber production, carbon sequestration, landscape flammability, and reduced forage quality, natural or human‐assisted migration of this species is likely to substantially alter responses of Alaskan forest ecosystems to climate change. [ABSTRACT FROM AUTHOR]

Anticipating consequences of disturbance interactions on ecosystem structure and function is a critical management priority as disturbance activity increases with warming climate. Across the Northern Hemisphere, extensive tree mortality from recent bark beetle outbreaks raises concerns about potential fire behavior and post-fire forest function. Silvicultural treatments (that is, partial or complete cutting of forest stands) may reduce outbreak severity and subsequent fuel loads, but longevity of pre-outbreak treatment effects on outbreak severity and post-outbreak fuel profiles remains underexplored. Further, treatments may present tradeoffs for other management objectives focused on ecosystem services (for example, carbon storage). We measured structure in old-growth subalpine forests following a recent (early 2000s) severe mountain pine beetle (MPB; Dendroctonus ponderosae) outbreak to examine effects of historical (1940s) cutting intensity on gray stage (~10 years after peak of outbreak) post-outbreak (1) fuel profiles and (2) aboveground biomass carbon. Compared to control (uncut) stands, historically cut stands subjected to the same MPB outbreak had approximately half the post-outbreak surface fuel loads, about 2–3x greater live canopy fuel loads, and greater within-stand spatial heterogeneity of dead canopy cover and available canopy fuel load. Post-outbreak total aboveground biomass carbon was similar across all stands, though historically cut stands had about 2x greater carbon in live biomass compared to uncut stands. These findings suggest tradeoffs with altered post-outbreak potential fire behavior and carbon storage in cut stands. Additional implications of historical silvicultural treatments for wildlife habitat, firefighting operations, and long-term carbon trajectories highlight temporal legacies of management on directing forest response to interacting disturbances. [ABSTRACT FROM AUTHOR]

Endophytes have been defined as microorganisms living inside plant tissues without causing negative effects on their hosts. Endophytic microbes have been extensively studied for their plant growth-promoting traits. However, analyses of endophytes require complete removal of epiphytic microorganisms. We found that the established tests to evaluate surface sterility, polymerase chain reaction, and leaf imprints, are unreliable. Therefore, we used scanning electron microscopy (SEM) as an additional assessment of epiphyte removal. We used a diverse suite of sterilization protocols to remove epiphytic microorganisms from the leaves of a gymnosperm and an angiosperm tree to test the influence of leaf morphology on the efficacy of these methods. Additionally, leaf tissue damage was also evaluated by SEM, as damaging the leaves might have an impact on endophytes and could lead to inaccurate assessment of endophytic communities. Our study indicates, that complete removal of the leaf cuticle by the sterilization technique assures loss of epiphytic microbes, and that leaves of different tree species may require different sterilization protocols. Furthermore, our study demonstrates the importance of choosing the appropriate sterilization protocol to prevent erroneous interpretation of host-endophyte interactions. Moreover, it shows the utility of SEM for evaluating the effectiveness of surface sterilization methods and their impact on leaf tissue integrity.

Endophytes have been defined as microorganisms living inside plant tissues without causing negative effects on their hosts. Endophytic microbes have been extensively studied for their plant growth-promoting traits. However, analyses of endophytes require complete removal of epiphytic microorganisms. We found that the established tests to evaluate surface sterility, polymerase chain reaction, and leaf imprints, are unreliable. Therefore, we used scanning electron microscopy (SEM) as an additional assessment of epiphyte removal. We used a diverse suite of sterilization protocols to remove epiphytic microorganisms from the leaves of a gymnosperm and an angiosperm tree to test the influence of leaf morphology on the efficacy of these methods. Additionally, leaf tissue damage was also evaluated by SEM, as damaging the leaves might have an impact on endophytes and could lead to inaccurate assessment of endophytic communities. Our study indicates, that complete removal of the leaf cuticle by the sterilization technique assures loss of epiphytic microbes, and that leaves of different tree species may require different sterilization protocols. Furthermore, our study demonstrates the importance of choosing the appropriate sterilization protocol to prevent erroneous interpretation of host-endophyte interactions. Moreover, it shows the utility of SEM for evaluating the effectiveness of surface sterilization methods and their impact on leaf tissue integrity.

Abstract Allometric equations predict organism attributes from simple measurements and underlie many global‐scale estimates, from plant productivity to ecosystem carbon stocks. In forests, destructive harvesting of trees in common groups (e.g. plant functional type) or at the species level is necessary to develop allometry but, since sampling is extremely difficult, predictions from these equations have high uncertainty due to low sample size, spatial bias and unrepresentative sampling of tree size. Terrestrial laser scanning (TLS) is a promising remote sensing technology that enables efficient and nondestructive estimates of tree‐level structure for developing allometric equations. Here, we nondestructively estimated component biomass of three coniferous tree species (Pinus ponderosa, Pinus contorta and Pseudotsuga menziesii) in Colorado, USA using TLS. We evaluated the suitability for this nondestructive data to be supplanted for destructive data in the development of species‐specific allometric equations and compared the prediction accuracy against a commonly used national‐scale allometry. We found TLS biomass estimates were consistently more precise across species (RMSE = ~19%) than nation‐scale allometry (RMSE = ~39%). Nondestructive biomass estimates from TLS are a suitable addition to or replacement for traditional sampling methods, with indistinguishable biomass predictions across most of the tested diameter range. We further show how TLS can be used to develop allometric equations compatible with airborne LiDAR and other remote sensing variables (e.g. height and crown area), developing generalized biomass predictions from crown area and tree height (R2 = 0.87). The ability for TLS to support the development of nondestructive allometry at a global scale will enable a more nuanced understanding of the drivers of individual tree architecture, while supporting the next generation of biomass remote sensing.

The environment could alter growth and resistance tradeoffs in plants by affecting the ratio of resource allocation to various competing traits. Yet, how and why functional tradeoffs change over time and space is poorly understood particularly in long-lived conifer species. By establishing four common-garden test sites for five lodgepole pine populations in western Canada, combined with genomic sequencing, we revealed the decoupling pattern and genetic underpinnings of tradeoffs between height growth, drought resistance based on δ13C and dendrochronology, and metrics of pest resistance based on pest suitability ratings. Height and δ13C correlation displayed a gradient change in magnitude and/or direction along warm-to-cold test sites. All cold test sites across populations showed a positive height and δ13C relationship. However, we did not observe such a clinal correlation pattern between height or δ13C and pest suitability. Further, we found that the study populations exhibiting functional tradeoffs or synergies to various degrees in test sites were driven by non-adaptive evolutionary processes rather than adaptive evolution or plasticity. Finally, we found positive genetic relationships between height and drought or pest resistance metrics and probed five loci showing potential genetic tradeoffs between northernmost and the other populations. Our findings have implications for deciphering the ecological, evolutionary, and genetic bases of the decoupling of functional tradeoffs due to environmental change. [ABSTRACT FROM AUTHOR]

Previous research on the effects of neighborhood crowding and soil moisture on tree height growth have been limited by time-consuming and sometimes inaccurate ground-based measurements of tree height. Recent developments in unmanned aerial vehicles (UAVs) allow detailed 3D point clouds of the canopy surface to be generated at relatively low cost. Using UAV-derived point clouds, we obtained height measurements of 4386 trees for the years 2019 and 2021. We also calculated four neighborhood crowding indices and a topography-based moisture index (depth-to-water) for these trees. Using initial tree height, neighborhood crowding indices and the depth-to-water index, we developed Bayesian hierarchical models to predict height growth for three tree species (Picea glauca (white spruce), Populus tremoluides (trembling aspen) and Pinus contorta (lodgepole pine)) across different stands. Bayes-R 2 values of the final models were highest for white spruce (35%) followed by trembling aspen (28%) and lodgepole pine (25%). Model outputs showed that the effect of crowding and depth-to-water on height growth are limited and species-dependent, adding a maximum of 7% to the Bayes-R 2 metric. Comparing different neighborhood crowding indices revealed that no index is clearly superior to others across all three species, as different neighborhood crowding indices resulted in only minor differences in model performance. While height growth can be partially explained by aerially derived neighborhood crowding indices and the depth-to-water index, future studies should focus on identifying relevant site characteristics to predict tree growth with greater accuracy. [ABSTRACT FROM AUTHOR]

Bark beetles maintain symbiotic associations with a diversity of microbial organisms, including ophiostomatoid fungi. Studies have frequently reported the role of ophiostomatoid fungi in bark beetle biology, but how fungal symbionts interact with host chemical defenses over time is needed. We first investigated how inoculations by three fungal symbionts of mountain pine beetle affect the terpene chemistry of live lodgepole pine trees. We then conducted a complimentary laboratory experiment specifically measuring the host metabolite degradation by fungi and collected the fungal organic volatiles following inoculations with the same fungal species on lodgepole pine logs. In both experiments, we analyzed the infected tissues for their terpene chemistry. Additionally, we conducted an olfactometer assay to determine whether adult beetles respond to the volatile organic chemicals emitted from each of the three fungal species. We found that all fungi upregulated terpenes as early as two weeks after inoculations. Similarly, oxygenated monoterpene concentrations also increased by several folds (only in logs). A large majority of beetles tested showed a strong attraction to two fungal species, whereas the other fungus repelled the beetles. Together this study shows that fungal symbionts can alter host defense chemistry, assist beetles in overcoming metabolite toxicity, and provide possible chemical cues for bark beetle attraction. [ABSTRACT FROM AUTHOR]

Allometric equations predict organism attributes from simple measurements and underlie many global‐scale estimates, from plant productivity to ecosystem carbon stocks. In forests, destructive harvesting of trees in common groups (e.g. plant functional type) or at the species level is necessary to develop allometry but, since sampling is extremely difficult, predictions from these equations have high uncertainty due to low sample size, spatial bias and unrepresentative sampling of tree size. Terrestrial laser scanning (TLS) is a promising remote sensing technology that enables efficient and nondestructive estimates of tree‐level structure for developing allometric equations.Here, we nondestructively estimated component biomass of three coniferous tree species (Pinus ponderosa, Pinus contorta and Pseudotsuga menziesii) in Colorado, USA using TLS. We evaluated the suitability for this nondestructive data to be supplanted for destructive data in the development of species‐specific allometric equations and compared the prediction accuracy against a commonly used national‐scale allometry.We found TLS biomass estimates were consistently more precise across species (RMSE = ~19%) than nation‐scale allometry (RMSE = ~39%). Nondestructive biomass estimates from TLS are a suitable addition to or replacement for traditional sampling methods, with indistinguishable biomass predictions across most of the tested diameter range. We further show how TLS can be used to develop allometric equations compatible with airborne LiDAR and other remote sensing variables (e.g. height and crown area), developing generalized biomass predictions from crown area and tree height (R2 = 0.87).The ability for TLS to support the development of nondestructive allometry at a global scale will enable a more nuanced understanding of the drivers of individual tree architecture, while supporting the next generation of biomass remote sensing. [ABSTRACT FROM AUTHOR]

Understanding how climate warming will affect the demographic rates of different ecotypes is critical to predicting shifts in species distributions. Here, we present results from a common garden, climate change experiment in which we measured seedling recruitment of lodgepole pine, a widespread North American conifer that is also planted globally. Seeds from a low-elevation provenance had more than three-fold greater recruitment to their third year than seeds from a high-elevation provenance across sites within and above its native elevation range and across climate manipulations. Heating halved recruitment to the third year of both low- and high-elevation seed sources across the elevation gradient, while watering more than doubled recruitment, alleviating some of the negative effects of heating. Demographic models based on recruitment data from the climate manipulations and long-term observations of adult populations revealed that heating could effectively halt modeled upslope range expansion except when combined with watering. Simulating fire and rapid postfire forest recovery at lower elevations accelerated lodgepole pine expansion into the alpine, but did not alter final abundance rankings among climate scenarios. Regardless of climate scenario, greater recruitment of low-elevation seeds compensated for longer dispersal distances to treeline, assuming colonization was allowed to proceed over multiple centuries. Our results show that ecotypes from lower elevations within a species' range could enhance recruitment and facilitate upslope range shifts with climate change.

While droughts, intensified by climate change, have been affecting forests worldwide, pest epidemics are a major source of uncertainty for assessing drought impacts on forest trees. Thus far, little information has documented the adaptability and evolvability of traits related to drought and pests simultaneously. We conducted common-garden experiments to investigate how several phenotypic traits (i.e. height growth, drought avoidance based on water-use efficiency inferred from δ13C and pest resistance based on defence traits) interact in five mature lodgepole pine populations established in four progeny trials in western Canada. The relevance of interpopulation variation in climate sensitivity highlighted that seed-source warm populations had greater adaptive capability than cold populations. In test sites, warming generated taller trees with higher δ13C and increased the evolutionary potential of height growth and δ13C across populations. We found, however, no pronounced gradient in defences and their evolutionary potential along populations or test sites. Response to selection was weak in defences across test sites, but high for height growth particularly at warm test sites. Response to the selection of δ13C varied depending on its selective strength relative to height growth. We conclude that warming could promote the adaptability and evolvability of growth response and drought avoidance with a limited evolutionary influence from pest (biotic) pressures. [ABSTRACT FROM AUTHOR]

Mountain meadows have high biodiversity and help regulate stream water release following the snowmelt pulse. However, many meadows are experiencing woody plant encroachment, threatening these ecosystem services. While there have been field surveys of individual meadows and remote sensing-based landscape-scale studies of encroachment, what is missing is a broad-scale, ground-based study to understand common regional drivers, especially at high elevations, where land management has often played a less direct role. With this study, we ask: What are the climate and landscape conditions conducive to woody plant encroachment at the landscape scale, and how has historical climate variation affected tree recruitment in subalpine meadows over time? We measured density of encroaching trees across 340 subalpine meadows in the central Sierra Nevada, California, USA, and used generalized additive models (GAMs) to determine the relationship between landscape-scale patterns of encroachment and meadow environmental properties. We determined ages of trees in 30 survey meadows, used observed climate and GAMs to model the relationship between timing of recruitment and climate since the early 1900s, and extrapolated recruitment patterns into the future using downscaled climate scenarios. Encroachment was high among meadows with lodgepole pine (Pinus contorta Douglas ex Loudon var. murrayana (Balf.) Engelm.) in the immediate vicinity, at lower elevations, with physical conditions favoring strong soil drying, and with maximum temperatures above or below average. Climatic conditions during the year of germination were unimportant, with tree recruitment instead depending on a 3-yr seed production period prior to germination and a 6-yr seedling establishment period following germination. Recruitment was high when the seed production period had high snowpack, and when the seedling establishment period had warm summer maximum temperatures, high summer precipitation, and high snowpack. Applying our temporal model to downscaled output from four global climate models indicated that the average meadow will shift to forest by the end of the 21st century. Sierra Nevada meadow encroachment by conifers is ubiquitous and associated with climate conditions increasingly favorable for tree recruitment, which will lead to substantial changes in subalpine meadows and the ecosystem services they provide.

Bark beetles maintain symbiotic associations with a diversity of microbial organisms, including ophiostomatoid fungi. Studies have frequently reported the role of ophiostomatoid fungi in bark beetle biology, but how fungal symbionts interact with host chemical defenses over time is needed. We first investigated how inoculations by three fungal symbionts of mountain pine beetle affect the terpene chemistry of live lodgepole pine trees. We then conducted a complimentary laboratory experiment specifically measuring the host metabolite degradation by fungi and collected the fungal organic volatiles following inoculations with the same fungal species on lodgepole pine logs. In both experiments, we analyzed the infected tissues for their terpene chemistry. Additionally, we conducted an olfactometer assay to determine whether adult beetles respond to the volatile organic chemicals emitted from each of the three fungal species. We found that all fungi upregulated terpenes as early as two weeks after inoculations. Similarly, oxygenated monoterpene concentrations also increased by several folds (only in logs). A large majority of beetles tested showed a strong attraction to two fungal species, whereas the other fungus repelled the beetles. Together this study shows that fungal symbionts can alter host defense chemistry, assist beetles in overcoming metabolite toxicity, and provide possible chemical cues for bark beetle attraction.

Abstract Background Water-use efficiency (WUE) represents the coupling of forest carbon and water. Little is known about the responses of WUE to thinning at multiple spatial scales. The objective of this research was to use field measurements to understand short-term effects of two thinning treatments (T1: 4500 stems ha−1; and T2: 1100 stems ha−1) and the control (NT: 27,000 stems ha−1) on WUE at the three spatial scales (leaf level: the ratio of leaf photosynthesis to leaf transpiration; tree-level: tree growth to tree transpiration; and stand level: net primary production (NPP) to stand transpiration) and intrinsic WUEi (the ratio of leaf photosynthesis to stomatal conductance at leaf-level; and NPP to canopy conductance at stand-level) in a 16-year old natural lodgepole pine forest. Leaf-level measurements were conducted in 2017, while tree- and stand-level measurements were conducted in both 2016 (the normal precipitation year) and 2017 (the drought year). Results The thinning treatments did not significantly affect the tree- and stand-level WUE in the normal year of 2016. However, the thinning significantly affected WUE in the drought year of 2017: T2 exhibited significantly higher tree-level WUE (0.49 mm2 kg−1) than NT (0.08 mm2 kg−1), and compared to NT, the stand-level WUE values in the thinned stands (T1 and T2) were significantly higher, with means of 0.31, 0.56 and 0.70 kg m−3, respectively. However, the leaf-level and stand-level WUEi in the thinned stands in the drought year were significantly lower than those in the unthinned stands. No significant differences in the leaf-level WUE were found among the treatments in 2017. In addition, the thinning did not significantly change the WUE-VPD relationships at any studied spatial scale. Conclusions The thinning treatments did not cause significant changes in all studied WUE metrics in a normal year. However, their effects were significantly promoted under the drought conditions probably due to the decrease in soil water availability, demonstrating that thinning can improve WUE and consequently support forests to cope with the drought effects. The inconsistent results on the effects of the thinning on forest carbon and water coupling at the spatial scales and the lack of the consistent WUE metrics constraint across-scale comparison and transferring of WUE.

Promoting ecological resilience to increasing disturbance activity is a key management priority under warming climate. Across the Northern Hemisphere, tree mortality from widespread bark beetle outbreaks raises concerns for how forest management can foster resilience to future outbreaks. Density reduction (i.e., thinning) treatments can increase vigor of remaining trees, but the longevity of treatment efficacy for reducing susceptibility to future disturbance remains a key knowledge gap. Using one of the longest‐running replicated experiments in old‐growth subalpine forests, we measured stand structure following a recent (early 2000s) severe mountain pine beetle (MPB; Dendroctonus ponderosae) outbreak to examine the legacy of historical (1940s) thinning treatments on two components of resilience. We asked: 'How did historical thinning intensity affect (1) tree‐scale survival probability and stand‐scale survival proportion (collectively "resistance" to outbreak) for susceptible trees (lodgepole pine [Pinus contorta] ≥ 12 cm diameter) and (2) post‐outbreak stand successional trajectories?' Overall outbreak severity was high (MPB killed 59% of susceptible individuals and 78% of susceptible basal area), and historical thinning had little effect on tree‐scale and stand‐scale resistance. Tree‐scale survival probability decreased sharply with increasing tree diameter and did not differ from the control (uncut stands) in the historical thinning treatments. Stand‐scale proportion of surviving susceptible trees and basal area did not differ from the control in historically thinned stands, except for treatments that removed nearly all susceptible trees, in which survival proportion approximately doubled. Despite limited effects on resistance to MPB outbreak, the legacy of historical treatments shifted dominance from large‐diameter to small‐diameter lodgepole pine by the time of outbreak, resulting in historically thinned stands with ~2× greater post‐outbreak live basal area than control stands. MPB‐driven mortality of large‐diameter lodgepole pine in control stands and density‐dependent mortality of small‐diameter trees in historically thinned stands led to convergence in post‐outbreak live tree stand structure. One exception was the heaviest historical thinning treatments (59–77% basal area removed), for which sapling dominance of shade‐tolerant, unsusceptible conifers was lower than control stands. After six decades, thinning treatments have had minimal effect on resistance to bark beetle outbreaks, but leave persistent legacies in shaping post‐outbreak successional trajectories. [ABSTRACT FROM AUTHOR]

Abstract Background Biomass maps are valuable tools for estimating forest carbon and forest planning. Individual-tree biomass estimates made using allometric equations are the foundation for these maps, yet the potentially-high uncertainty and bias associated with individual-tree estimates is commonly ignored in biomass map error. We developed allometric equations for lodgepole pine (Pinus contorta), ponderosa pine (P. ponderosa), and Douglas-fir (Pseudotsuga menziesii) in northern Colorado. Plot-level biomass estimates were combined with Landsat imagery and geomorphometric and climate layers to map aboveground tree biomass. We compared biomass estimates for individual trees, plots, and at the landscape-scale using our locally-developed allometric equations, nationwide equations applied across the U.S., and the Forest Inventory and Analysis Component Ratio Method (FIA-CRM). Total biomass map uncertainty was calculated by propagating errors from allometric equations and remote sensing model predictions. Two evaluation methods for the allometric equations were compared in the error propagation—errors calculated from the equation fit (equation-derived) and errors from an independent dataset of destructively-sampled trees (n = 285). Results Tree-scale error and bias of allometric equations varied dramatically between species, but local equations were generally most accurate. Depending on allometric equation and evaluation method, allometric uncertainty contributed 30–75% of total uncertainty, while remote sensing model prediction uncertainty contributed 25–70%. When using equation-derived allometric error, local equations had the lowest total uncertainty (root mean square error percent of the mean [% RMSE] = 50%). This is likely due to low-sample size (10–20 trees sampled per species) allometric equations and evaluation not representing true variability in tree growth forms. When independently evaluated, allometric uncertainty outsized remote sensing model prediction uncertainty. Biomass across the 1.56 million ha study area and uncertainties were similar for local (2.1 billion Mg; % RMSE = 97%) and nationwide (2.2 billion Mg; % RMSE = 94%) equations, while FIA-CRM estimates were lower and more uncertain (1.5 billion Mg; % RMSE = 165%). Conclusions Allometric equations should be selected carefully since they drive substantial differences in bias and uncertainty. Biomass quantification efforts should consider contributions of allometric uncertainty to total uncertainty, at a minimum, and independently evaluate allometric equations when suitable data are available.

Mountain pine beetle (MPB; Dendroctonus ponderosae Hopkins) is a devastating forest insect pest that has killed millions of hectares of pines in western North America over the past two decades. Like other bark beetles, MPB vectors ophiostomatoid fungal species, some of which are pathogenic to host pine species. The phytopathogenicity of these fungal symbionts has sparked considerable debate regarding their role in facilitating MPB attack success. We tested the hypothesis that MPB ophiostomatoid fungal associates like Grosmannia clavigera (Robinson-Jeffrey and Davidson) Zipfel, de Beer and Wingfield contribute to overwhelming host defenses during MPB mass attack. We compared responses of mature lodgepole pine ( Pinus contorta Dougl. ex Loud. var. latifolia Engelm.) trees growing in natural stands that were mass attacked by MPB with those inoculated with G. clavigera by examining host defense hormones, secondary metabolites, and gene expression profiles. The jasmonate and ethylene signatures of necrotrophic pathogen-triggered response were identified in G. clavigera -inoculated trees, but only the jasmonate signature of a herbivore-triggered response was measured in MPB-attacked trees. Several G. clavigera -induced changes in pine phenolic metabolite profiles and phenolic biosynthesis gene expression patterns were absent in MPB-attacked pines. These findings indicate that ophiostomatoid fungi like G. clavigera are not a major factor in overwhelming host defenses during MPB mass attack. Instead, fungal pathogenicity likely is more important in aiding MPB colonization and development within the host tree. Phenolics appear to play a larger role in the host response to G. clavigera than to MPB, although phenolics may also influence MPB feeding and behavior. [Formula: see text] Copyright © 2024 The Author(s). This is an open access article distributed under the CC BY 4.0 International license., Competing Interests: The author(s) declare no conflict of interest.

Tree‐species mixture effects (e.g. complementarity and facilitation) have been found to increase individual‐tree productivity, lessen mortality and improve recruitment in forests worldwide. By promoting more efficient and complete resource use, mixture effects may also lessen individual‐tree‐level water stress, thus improving drought resistance. We investigated the influence of mixture effects on tree productivity, mortality and recruitment across broad compositional and moisture gradients in high‐elevation Interior West US mixed‐conifer communities, where Populus tremuloides (trembling aspen) is the major contributor to functional diversity. Our goal was to provide a more complete scientific foundation for managing these drought‐prone, fire‐excluded systems under an uncertain climate.We used landscape‐scale national forest inventory data to examine mixture effects on P. tremuloides and the major associated conifer species, Pseudotsuga menziesii, Pinus contorta, Abies lasiocarpa and Picea engelmannii. Using generalized linear mixed modelling, we isolated the influences of P. tremuloides relative density and climate on tree‐level (stems ≥ 12.7 cm DBH) growth, mortality and stand‐level recruitment (presence/absence of new trees). Cold‐season precipitation (PPT) and warm‐season vapour pressure deficit (VPD) served to represent soil moisture supply and demand, respectively.Populus tremuloides growth declined as interspecific density increased. In contrast, Pinus contorta and A. lasiocarpa growth increased with P. tremuloides density. For all species except A. lasiocarpa and P. menziesii, growth increased under higher PPT and VPD. Populus tremuloides mortality increased under high VPD but not with interspecific relative density. We found limited evidence that A. lasiocarpa mortality decreased as P. tremuloides density increased. Populus tremuloides recruitment declined steeply above 25% interspecific relative density. We found a decline in conifer recruitment odds as P. tremuloides density increased, ranging from strong in P. contorta to insubstantial in P. engelmannii.Synthesis. Our findings have implications for sustaining mixed‐conifer communities impacted by climate change and historical fire exclusion. Mixtures of P. tremuloides and conifers may improve conifer growth while adversely impacting P. tremuloides growth relative to pure stands. Higher conifer productivity combined with lower P. tremuloides recruitment at high conifer relative density may accelerate succession. [ABSTRACT FROM AUTHOR]

Legacy effects from one disturbance may influence successional pathways by amplifying or buffering forest regeneration after the next disturbance. We assessed vegetation and tree regeneration in non-serotinous Sierra lodgepole pine (Pinus contorta var. murrayana) stands after a 1984 wildfire which burned with variable severity and again after a high-severity subsequent fire in 2012. The legacy effects of the 1984 fire were amplified; seedlings and saplings were abundant in areas initially burned at low severity (1267 stems ha−1) despite high reburn severity, but regeneration was low in areas twice burned at high severity (31 stems ha−1). Our results suggest that the severity of the 1984 fire may have influenced post-2012 tree regeneration by creating variable fuel loading, which may have affected soils, litter cover and shade after the 2012 fire and therefore affected seedling establishment and survival. A canopy seed bank of unburnt cones from trees killed by the 2012 fire potentially contributed to a strong effect of prior burn severity on regeneration after the 2012 fire despite a lack of serotinous or resprouting tree species, although the influence of this canopy seedbank was likely limited to the year following the fire. Our results suggest that a low- to moderate-severity fire increases forest resilience relative to a high-severity fire even when the next fire burns at high severity. [ABSTRACT FROM AUTHOR]

Increased climatic variability can impact tree physiological processes beyond what is predicted from changes in mean conditions. We assessed the sensitivity of conifer saplings to spatial and temporal variability in meteorological conditions, taking advantage of the end of California's historic drought and the exceedingly wet winter of 2017. We sought to understand how very dry and very wet conditions constrain photosynthesis and growth in four regionally dominant conifers and whether sensitivity in these processes changes across a 500 m gradient in elevation. All species demonstrated phenotypic plasticity in response to temporal differences in precipitation on both inter-annual and seasonal timescales. Net photosynthesis in Pinus contorta decreased from an early season 2016 average of 12.4 to 6.89 μmol CO2 m−2 s−1 later in the summer, but increased 14.1% between seasons in the wet year. By contrast, elevation had almost no effect on instantaneous photosynthetic gas exchange, CO2 response curve parameters, or stem water potential in any of the years for any of the species. Effects of the heavy snow year (2017) on needle growth differed between elevations. Pinus contorta showed a 38.9% increase in average needle length at the lower two elevations but a 31.6% decrease at the highest site compared to the height of the drought. Despite these differences, biological variation was dampened compared to the physical variation between years, suggesting these trees can effectively withstand substantial meteorological variability. Our results show that these species demonstrated considerable ability to tolerate and recover from an extreme drought event. [ABSTRACT FROM AUTHOR]

The effects of changing climate and disturbance on mountain forest carbon (C) stocks vary with tree species distributions and over elevational gradients. Warming can not only increase C uptake by stimulating productivity at high elevations but also enhance C release by increasing respiration and the frequency, intensity and size of wildfires.To understand the consequences of climate change for temperate mountain forests, we simulated interactions among climate, wildfire, tree species and their combined effects on regional C stocks in forests of the Greater Yellowstone Ecosystem, USA (GYE) with the LANDIS‐II landscape change model. Simulations used historical climate and future potential climate represented by downscaled projections from five general circulation models (GCMs) that bracket the range of variability under the representative concentration pathway (RCP) 8.5 emissions scenario.Total ecosystem C increased by 67% through 2100 in simulations with historical climate, and by 38%–69% with GCM climate. Differences in C uptake among GCMs resulted primarily from variation in area burned, not productivity. Warming increased productivity by extending the growing season, especially near upper tree line, but did not offset biomass losses to fire. By 2100, simulated area burned increased by 27%–215% under GCM climate, with the largest increases after 2050. With warming >3°C in mean annual temperature, the increased frequency of large fires reduced live C stocks by 4%–36% relative to the control, historical climate scenario. However, relative losses in total C were delayed under GCMs with large increases in summer precipitation and buffered by C retained in soils and the wood of fire‐killed trees. Increasing fire size limited seed dispersal, and reductions in soil moisture limited seedling establishment; both effects will likely constrain long‐term forest regeneration and C uptake.Synthesis. Forests in the GYE can maintain a C sink through the mid‐century in a warming climate but continued warming may cause the loss of forest area, live above‐ground biomass and, ultimately, ecosystem C. Future changes in C stocks in similar forests throughout western North America will depend on regional thresholds for extensive wildfire and forest regeneration and therefore, changes may occur earlier in drier regions. [ABSTRACT FROM AUTHOR]

Aim: Climate change poses significant challenges for tree species, which are slow to adapt and migrate. Insight into genetic and phenotypic variation under current landscape conditions can be used to gauge persistence potential to future conditions and determine conservation priorities, but landscape effects have been minimally tested in trees. Here, we use Pinus contorta, one of the most widely distributed conifers in North America, to evaluate the influence of landscape heterogeneity on genetic structure as well as the magnitude of local adaptation versus phenotypic plasticity in a widespread tree species. Location: Western North America. Methods: We paired landscape genetics with fully reciprocal in situ common gardens to evaluate landscape influence on neutral and adaptive variation across all subspecies of P. contorta. Results: Landscape barriers alone play a minor role in limiting gene flow, creating marginal geographically‐based structure. Local climate determines population performance, with survival highest at home but growth greatest in mild climates (e.g., warm, wet). Survival of two of the three populations tested was consistent with patterns of local adaptation documented for P. contorta, while growth was indicative of plasticity for populations grown under novel conditions and suggesting that some populations are not currently occupying their climatic optimum. Main Conclusions: Our findings provide insight into the role of the landscape in shaping population genetic structure in a widespread tree species as well as the potential response of local populations to novel conditions, knowledge critical to understanding how widely distributed species may respond to climate change. Geographically based genetic structure and reduced survival under water‐limited conditions may make some populations of widespread tree species more vulnerable to local maladaptation and extirpation. However, genetically diverse and phenotypically plastic populations of widespread trees, such as many of the P. contorta populations sampled and tested here, likely possess high persistence potential. [ABSTRACT FROM AUTHOR]

Bark beetle outbreaks alter forests in many ways including stand structure, fuels and fire behavior, wildlife habitat, and aesthetic value. Less understood are the effects outbreaks have on understory vegetation, despite the importance for overstory succession, nutrient cycling, water quality, soil erosion, and wildlife. Beetle outbreaks also change forests in ways that could promote invasion by nonnative weeds, but this is rarely studied. We assessed changes in cover of understory vegetation and presence/abundance of weeds in lodgepole pine forests in Colorado, Idaho, Montana, Utah and Wyoming, USA following recent mountain pine beetle outbreaks. Incipient beetle populations began in 2004, peaked around 2008, and returned to endemic levels by 2012. Understory vegetation was sampled in 2010, 2012, 2014 and 2018. Total understory cover and cover of shrubs and graminoids remained unchanged, while cover of forbs generally increased. Forb cover was negatively correlated with shrub and canopy cover. Approximately 20% of plots contained weeds and weed abundance increased over time. Presence of weeds was negatively correlated with graminoid cover and positively correlated with tree mortality and snag fall. By 2018, weed abundance was only positively correlated with snag fall. Localized soil disturbance created when snags uproot and fall appears to act as sites of weed invasion and proliferation. Lastly, we provide a global review of the effects of bark beetle outbreaks on understory vegetation which reveals a prevailing short-term increase following outbreaks, but the magnitude of response and cover type affected varies with beetle species/forest type and time since outbreak.

Summary: Disturbances have altered community dynamics in boreal forests with unknown consequences for belowground ecological processes. Soil fungi are particularly sensitive to such disturbances; however, the individual response of fungal guilds to different disturbance types is poorly understood.Here, we profiled soil fungal communities in lodgepole pine forests following a bark beetle outbreak, wildfire, clear‐cut logging, and salvage‐logging. Using Illumina MiSeq to sequence ITS1 and SSU rDNA, we characterized communities of ectomycorrhizal, arbuscular mycorrhizal, saprotrophic, and pathogenic fungi in sites representing each disturbance type paired with intact forests. We also quantified soil fungal biomass by measuring ergosterol.Abiotic disturbances changed the community composition of ectomycorrhizal fungi and shifted the dominance from ectomycorrhizal to saprotrophic fungi compared to intact forests. The disruption of the soil organic layer with disturbances correlated with the decline of ectomycorrhizal and the increase of arbuscular mycorrhizal fungi. Wildfire changed the community composition of pathogenic fungi but did not affect their proportion and diversity. Fungal biomass declined with disturbances that disrupted the forest floor.Our results suggest that the disruption of the forest floor with disturbances, and the changes in C and nutrient dynamics it may promote, structure the fungal community with implications for fungal biomass–C. See also the Commentary on this article by Marín & Kohout, 229: 656–658. [ABSTRACT FROM AUTHOR]

Silicification is the most important process of fossilization resulting in the preservation of internal tissues in plants, thereby providing essential information on the anatomy, life history, and evolution of land plants. However, fundamental knowledge of silica uptake, precipitation, and contribution to in situ plant fossilization is limited. To identify the cellular pathway of aqueous silica and subsequent fluid–wood interaction processes, we investigated upright standing young trees of lodgepole pine (Pinus contorta) in the hot‐spring environment of Cistern Spring, Yellowstone National Park, USA. Our multi‐method analytical approach using x‐ray diffraction, Raman spectroscopy, scanning electron microscopy, and electron probe microanalyses shows that the surficial and internally co‐precipitated silica–halite–gypsum assemblage traces the flow of silica‐rich fluid and documents fluid retention after short‐term hot‐spring water immersion and evaporation. Element distribution maps reveal systematic 2D and 3D differences between silica quantities deposited in earlywood and latewood. The distribution of inorganic impurities in cell walls traces the anatomical structure of the wood and indicates rapid migration of homogeneous fluid into the waterlogged organic substrate. Our results show that the preferential pathway of silica‐rich fluid into the above‐ground wood was through the decorticated periphery which took place during a short‐term flooding, ranging from days to weeks, of hot‐spring fluid. In conclusion, the fluid retention capability in cellular pore space controls the in situ silicification process. The silicification of trees in growth position is a rapid process in which the in vivo transport of silica‐rich fluid upward through the secondary xylem plays an insignificant role. [ABSTRACT FROM AUTHOR]

Background: Biomass maps are valuable tools for estimating forest carbon and forest planning. Individual-tree biomass estimates made using allometric equations are the foundation for these maps, yet the potentially-high uncertainty and bias associated with individual-tree estimates is commonly ignored in biomass map error. We developed allometric equations for lodgepole pine (Pinus contorta), ponderosa pine (P. ponderosa), and Douglas-fir (Pseudotsuga menziesii) in northern Colorado. Plot-level biomass estimates were combined with Landsat imagery and geomorphometric and climate layers to map aboveground tree biomass. We compared biomass estimates for individual trees, plots, and at the landscape-scale using our locally-developed allometric equations, nationwide equations applied across the U.S., and the Forest Inventory and Analysis Component Ratio Method (FIA-CRM). Total biomass map uncertainty was calculated by propagating errors from allometric equations and remote sensing model predictions. Two evaluation methods for the allometric equations were compared in the error propagation—errors calculated from the equation fit (equation-derived) and errors from an independent dataset of destructively-sampled trees (n = 285). Results: Tree-scale error and bias of allometric equations varied dramatically between species, but local equations were generally most accurate. Depending on allometric equation and evaluation method, allometric uncertainty contributed 30–75% of total uncertainty, while remote sensing model prediction uncertainty contributed 25–70%. When using equation-derived allometric error, local equations had the lowest total uncertainty (root mean square error percent of the mean [% RMSE] = 50%). This is likely due to low-sample size (10–20 trees sampled per species) allometric equations and evaluation not representing true variability in tree growth forms. When independently evaluated, allometric uncertainty outsized remote sensing model prediction uncertainty. Biomass across the 1.56 million ha study area and uncertainties were similar for local (2.1 billion Mg; % RMSE = 97%) and nationwide (2.2 billion Mg; % RMSE = 94%) equations, while FIA-CRM estimates were lower and more uncertain (1.5 billion Mg; % RMSE = 165%). Conclusions: Allometric equations should be selected carefully since they drive substantial differences in bias and uncertainty. Biomass quantification efforts should consider contributions of allometric uncertainty to total uncertainty, at a minimum, and independently evaluate allometric equations when suitable data are available. [ABSTRACT FROM AUTHOR]

The Scots pine (Pinus sylvestris L.) is the second-most widely used tree species in forestry in Latvia and is the only species used for afforestation on nutrient poor soils that cover considerable forest land in Latvia. Several studies have shown that, in such conditions, the lodgepole pine (Pinus contorta var. latifolia) may be more productive in terms of biomass and yield. It is important to consider climate change studies to assess the potential for a larger-scale use of the lodgepole pine in forestry. The aim was to assess the intra-annual height growth patterns of both species, the differences between them, and the influence of meteorological parameters on their height growth. Their height growth was monitored on a weekly basis in two sampling sites in central Latvia, and the height increment curves were described by Gompertz's model. The height growth dynamics of individual trees and species differed notably, indicating the potential for the selection of the best-adapted genotypes. Our results indicate that the early onset of the active growth phase might be the most important factor determining the total height increment for both species. Temperature-related meteorological parameters were the only ones with a statistically significant influence on pines height growth and only when at least one of the variables were standardised prior to the analysis. A temperature increase had a slightly stronger positive effect on the growth of the lodgepole pine, indicating that it might be suitable for more intensive use in forestry under the climate change scenarios for Latvia. [ABSTRACT FROM AUTHOR]

Ectomycorrhizal fungi (plant symbionts) are diverse and exist within spatially variable communities that play fundamental roles in the functioning of terrestrial ecosystems. However, the underlying ecological mechanisms that maintain and regulate the spatial structuring of ectomycorrhizal fungal communities are both complex and remain poorly understood. Here, we use a gradient of mountain pine beetle (Dendroctonus ponderosae) induced tree mortality across eleven stands in lodgepole pine (Pinus contorta) forests of western Canada to investigate: (i) the degree to which spatial structure varies within this fungal group, and (ii) how these patterns may be driven by the relative importance of tree mortality from changes in understory plant diversity, productivity and fine root biomass following tree death. We found that the homogeneity of the ectomycorrhizal fungal community increased with increasing tree death, aboveground understory productivity and diversity. Whereas, the independent effect of fine root biomass, which declined along the same gradient of tree mortality, increased the heterogeneity of the ectomycorrhizal fungal community. Together, our results demonstrate that large-scale biotic disturbance homogenizes the spatial patterns of ectomycorrhizal fungal communities.

The effects of herbivory on plant invasions are broadly discussed, and many studies have led to widely debated theories. In particular, the effects of herbivores on pine invasion found in different studies vary; in some cases, they controlled their expansion, and in others, they promoted it. On the other hand, vulnerability to invasion by pines differs between community types. Sites with dunes and bare ground are the most heavily invaded, followed by grasslands, while shrublands and forests are least invaded. Because current evidence is mostly observational, some of the varying responses of pine invasions to herbivory should be examined further through replicated experiments. Here, we address experimentally the extent to which preference for the non‐native invasive Pinus contorta by domestic sheep (Ovis aries) depends on the vegetation type. We installed experimental enclosures within two adjacent communities, grassland and shrubland, and in each one, we planted seedlings of P. contorta Douglas and established a sheep density typically recommended for the study area. The number of browsed seedlings, the number and type of branches browsed per seedling, the reduction in height and probability of survival immediately after browsing period were recorded. The number of browsed seedlings and damage to the terminal bud were higher in grassland than in shrubland, while the number of browsed branches per seedling was higher in shrubland than grassland. The reductions in height and probability of survival immediately after browsing were similar in both communities. These results show that moderate levels of sheep herbivory could reduce 20% seedling survival in both communities; nevertheless, the damage patterns differ between them. The sheep browsed more substantial number of seedlings in grasslands than in shrublands. However, if sheep find the seedlings, they damage it more in shrublands. These results suggest that experimental studies comparing communities are important for pine invasion management. [ABSTRACT FROM AUTHOR]

Phytopathogenic ophiostomatoid fungi are common associates of bark beetles and contribute to beetle-associated mortality of trees. Mountain pine beetle outbreaks in Canada are facilitating novel associations between its vectored fungi (Grosmannia clavigera , Leptographium longiclavatum and Ophiostoma montium) and jack pine. How the induced defense-related metabolite responses of jack and lodgepole pines vary in response to the fungi is unknown. Understanding this variation is important to clarifying pine susceptibility to and the physiological impacts of infection. We used a comparative metabolite profiling approach to investigate the defense-related signaling, carbon utilization/mobilization, and synthesis responses of both pines to the fungi. Both pine species largely exhibited similar metabolite responses to the fungi. The magnitude of pine metabolite responses positively reflected pathogen virulence. Our findings indicate that pines can recognize and metabolomically respond to novel pathogens, likely due to signals common between the novel fungi and fungi coevolved with the pine. Thus, jack pine is likely as susceptible as lodgepole pine to infections by each of the MPB-vectored fungi. Furthermore, the magnitude of the metabolite responses of both pines varied by the eliciting fungal species, with the most virulent pathogen causing the greatest reduction in carbohydrates and the highest accumulation of defensive terpenes. [ABSTRACT FROM AUTHOR]

Subalpine forests in the northern Rocky Mountains have been resilient to stand-replacing fires that historically burned at 100-to 300-year intervals. Fire intervals are projected to decline drastically as climate warms, and forests that reburn before recovering from previous fire may lose their ability to rebound. We studied recent fires in Greater Yellowstone (Wyoming, United States) and asked whether short-interval (<30 years) stand-replacing fires can erode lodgepole pine (Pinus contorta var. latifolia) forest resilience via increased burn severity, reduced early postfire tree regeneration, reduced carbon stocks, and slower carbon recovery. During 2016, fires reburned young lodgepole pine forests that regenerated after wildfires in 1988 and 2000. During 2017, we sampled 0.25-ha plots in stand-replacing reburns (n = 18) and nearby young forests that did not reburn (n = 9). We also simulated stand development with and without reburns to assess carbon recovery trajectories. Nearly all prefire biomass was combusted ("crown fire plus") in some reburns in which prefire trees were dense and small (=4-cm basal diameter). Postfire tree seedling density was reduced sixfold relative to the previous (long-interval) fire, and high-density stands (>40,000 stems ha-1) were converted to sparse stands (<1,000 stems ha-1). In reburns, coarse wood biomass and aboveground carbon stocks were reduced by 65 and 62%, respectively, relative to areas that did not reburn. Increased carbon loss plus sparse tree regeneration delayed simulated carbon recovery by >150 years. Forests did not transition to nonforest, but extreme burn severity and reduced tree recovery foreshadow an erosion of forest resilience. [ABSTRACT FROM AUTHOR]

Ectomycorrhizal fungi (plant symbionts) are diverse and exist within spatially variable communities that play fundamental roles in the functioning of terrestrial ecosystems. However, the underlying ecological mechanisms that maintain and regulate the spatial structuring of ectomycorrhizal fungal communities are both complex and remain poorly understood. Here, we use a gradient of mountain pine beetle (Dendroctonus ponderosae) induced tree mortality across eleven stands in lodgepole pine (Pinus contorta) forests of western Canada to investigate: (i) the degree to which spatial structure varies within this fungal group, and (ii) how these patterns may be driven by the relative importance of tree mortality from changes in understory plant diversity, productivity and fine root biomass following tree death. We found that the homogeneity of the ectomycorrhizal fungal community increased with increasing tree death, aboveground understory productivity and diversity. Whereas, the independent effect of fine root biomass, which declined along the same gradient of tree mortality, increased the heterogeneity of the ectomycorrhizal fungal community. Together, our results demonstrate that large-scale biotic disturbance homogenizes the spatial patterns of ectomycorrhizal fungal communities. [ABSTRACT FROM AUTHOR]

Non-indigenous conifers are considered invasive to the coastal dune heathland in Denmark, and massive clearing is carried out in an attempt to recreate and keep the original heathland. Burning is a common method for managing, but its feasibility to control the seed bank of conifers has not been investigated. This project shows that the burning of logged conifer trees will often eliminate seeds of lodgepole pine, mugo pine and Sitka spruce, even when the seeds were placed into a depth of five centimeters in the soil. The effect on seeds depends on the fuel load and the fire conditions (e.g., dryness, wind, and temperature). If the seeds were exposed to a high temperature, the seeds were not able to germinate afterward. The temperature was about 80 °C for all species. If the sum of temperatures based on temperature records every 30 s exceeded between 12,000 and 14,000 °C no seeds were able to germinate. The relationship between the mean temperature of the burns and the germination rate at seeds placed in various soil depths was modelled. Findings should be interpreted cautiously as each depth-species combinations were not replicated in space or time due to practical constraints.

Abstract Spatial variation in cone serotiny in Rocky Mountain lodgepole pine (Pinus contorta ssp. latifolia) across Yellowstone National Park influences initial pine recruitment after stand‐replacing fire with tremendous population, community, and ecosystem consequences. A previous study showed that much of the spatial variation in serotiny results from the balance of selection arising from high frequencies of fire favoring serotiny countered by opposing selection exerted by American red squirrels (Tamiasciurus hudsonicus) as seed predators. This earlier study, however, assumed stable local red squirrel densities over multiple generations of pines. Here, we examine environmental properties that might contribute to long‐term stability in the densities of red squirrels among sites. We found that the amount of clay in the soil, an indicator of plant and fungal growth—the latter an important food resource for red squirrels—and the coefficient of variation (CV) in diameter at breast height (DBH) of forest trees together account for a substantial amount of variation in red squirrel density. Soil development occurs over very long time scales, and thus, intersite variation in the amount of clay is unlikely to shift across pine generations. However, CV of DBH and squirrel density increase with stand age, which acts to amplify selection against serotiny with increasing interfire interval. Regardless, much of the variation in the CV of DBH is accounted for by soil bulk density, mean annual temperature, and surface curvature, which are unlikely to vary in their relative differences among sites over time. Consequently, these soil and abiotic attributes could contribute to consistent spatial patterns of red squirrel densities from one pine generation to the next, resulting in consistent local and spatial variation in selection exerted by red squirrels against serotiny.

In many regions, alien conifers have spread widely at lower elevations and are increasingly found colonizing alpine areas. Although studies have addressed conifer invasions at low elevations, little is known about the rates and constraints on spread into higher elevations. Here, we assess the relative importance of climate and the availability of regeneration microsites on the establishment of the alien species Pinus contorta into a high elevation site in New Zealand. Spread has occurred from two stands planted at the elevation of the native treeline (1347–1388 masl) in the 1960s. Most stems established between 1350 and 1450 masl and P. contorta individuals were found up to 270 m above the original plantings. Although the population has increased by 180% in the last 20 years, population growth rate has been declining. Furthermore, comparisons with studies from other mountain ranges around the world and at low elevations in New Zealand suggest this is a relatively limited spread. Our results suggest that climate variation did not have a significant effect on establishment patterns, as opposed to availability of regeneration microsites. Soil and alpine mat microsites favoured establishment of P. contorta and, although these microsites did not appear to be saturated, microsite availability may be an important limiting factor for the spread of P. contorta. Thus management strategies should focus on preventing spread in addition to removing already established stems.

Increased mortality of forest trees, driven directly or indirectly by climate change, is occurring around the world. In western North America, whitebark pine, a high elevation keystone species, and lodgepole pine, a widespread ecologically and economically important tree, have experienced extensive mortality in recent climate-driven outbreaks of the mountain pine beetle. However, even in stands experiencing high levels of mortality, some mature trees have survived. We hypothesized that the outbreak acted as a natural selection event, removing trees most susceptible to the beetle and least adapted to warmer drier conditions. If this was the case, genetic change would be expected at loci underlying beetle resistance. Given we did not know the basis for resistance, we used inter-simple sequence repeats to compare the genetic profiles of two sets of trees, survivors (mature, living trees) and general population (trees just under the diameter preferred by the beetles and expected to approximate the genetic structure of each tree species at the site without beetle selection). This method detects high levels of polymorphism and has often been able to detect patterns associated with phenotypic traits. For both whitebark and lodgepole pine, survivors and general population trees mostly segregated independently indicating a genetic basis for survivorship. Exceptions were a few general population trees that segregated with survivors in proportions roughly reflecting the proportion of survivors versus beetle-killed trees. Our results indicate that during outbreaks, beetle choice may result in strong selection for trees with greater resistance to attack. Our findings suggest that survivorship is genetically based and, thus, heritable. Therefore, retaining survivors after outbreaks to act as primary seed sources could act to promote adaptation. Further research will be needed to characterize the actual mechanism(s) of resistance.

Mountain pine beetle (Dendroctonus ponderosae Hopkins; MPB) is an aggressive bark beetle that attacks numerous Pinus spp. and causes extensive mortality in lodgepole pine (Pinus contorta Douglas ex Loudon; LPP) forests in the western United States and Canada. We used pre-outbreak LPP attributes, cumulative MPB attack severity, and areal extent of mortality data to identify subwatershed-scale forest attributes associated with severe MPB-caused tree mortality that occurred across the Northern Rockies, USA from 1999-2014. We upscaled stand-level data to the subwatershed scale to allow identification of large LPP areas vulnerable to MPB. The highest mortality occurred in subwatersheds where LPP mean basal area was greater than 11.5 m² ha-1 and LPP quadratic mean diameter was greater than or equal to 18 cm. A coarse assessment of federally-owned LPP-dominated forestland in the analysis area indicated about 42% could potentially be silviculturally treated. Silvicultural management may be a suitable option for many LPP forests, and our hazard model can be used to identify subwatersheds with LPP attributes associated with high susceptibility to MPB across landscape spatial scales. Identifying highly susceptible subwatersheds can help prioritize general areas for potential treatments, especially where spatially extensive areas of contiguous, highly susceptible LPP occur. [ABSTRACT FROM AUTHOR]

Increased mortality of forest trees, driven directly or indirectly by climate change, is occurring around the world. In western North America, whitebark pine, a high elevation keystone species, and lodgepole pine, a widespread ecologically and economically important tree, have experienced extensive mortality in recent climate-driven outbreaks of the mountain pine beetle. However, even in stands experiencing high levels of mortality, some mature trees have survived. We hypothesized that the outbreak acted as a natural selection event, removing trees most susceptible to the beetle and least adapted to warmer drier conditions. If this was the case, genetic change would be expected at loci underlying beetle resistance. Given we did not know the basis for resistance, we used inter-simple sequence repeats to compare the genetic profiles of two sets of trees, survivors (mature, living trees) and general population (trees just under the diameter preferred by the beetles and expected to approximate the genetic structure of each tree species at the site without beetle selection). This method detects high levels of polymorphism and has often been able to detect patterns associated with phenotypic traits. For both whitebark and lodgepole pine, survivors and general population trees mostly segregated independently indicating a genetic basis for survivorship. Exceptions were a few general population trees that segregated with survivors in proportions roughly reflecting the proportion of survivors versus beetle-killed trees. Our results indicate that during outbreaks, beetle choice may result in strong selection for trees with greater resistance to attack. Our findings suggest that survivorship is genetically based and, thus, heritable. Therefore, retaining survivors after outbreaks to act as primary seed sources could act to promote adaptation. Further research will be needed to characterize the actual mechanism(s) of resistance. [ABSTRACT FROM AUTHOR]

This paper analyzes sapwood variability and allometry within individuals of Populus tremuloides, Pinus contorta, Pinus banksiana, Picea mariana, and Picea glauca. Outside bark diameter at breast height (DBH) and sapwood depth (sd) in four cardinal directions were measured in individuals in stands in Alberta and Saskatchewan, Canada. The microscopical analysis of wood anatomy was used to measure sd, and the error associated with the measures was observed. Sapwood allometry analyses examined the influence of DBH on sd and on sapwood area (SA). All species were observed to have varying sapwood depths around the trunk with statistical analyses showing that Pinus banksiana has a well defined preference to grow thicker in the North-East side. The largest sd values were observed for the Populus tremuloides set. Unlike Populus tremuloides and Picea glauca, for the species Pinus contorta, Pinus banksiana, and Picea mariana, incremental growth in DBH does not directly drive sapwood growth in any direction. For these three species, SA increases only because of increases in DBH as sd remains nearly constant. These results show that sapwood depth and sapwood area seem to behave differently in each studied species and are not always proportional to the tree size as is normally assumed. [ABSTRACT FROM AUTHOR]