Search

Over 18 million hectares of lodgepole pine (Pinus contorta Dougl. var. latifolia Engelm.) forests have been affected by a massive mountain pine beetle outbreak in western Canada. The ensuing need for reforestation compels forest managers to deploy more drought-resistant stock in anticipation of a hotter and drier future climate. Recent studies suggest that naturally occurring lodgepole pine × jack pine (Pinus banksiana Lamb.) hybrids may be more drought-resistant than pure lodgepole pine. We used single-nucleotide polymorphism markers to determine the level of jack pine introgression present in the region J lodgepole pine seed orchard, in northwest Alberta, and we identified six hybrid genotypes with 10%–18% jack pine introgression for further study. We compared the performance of open-pollinated seedlings of lodgepole, jack, and hybrid pines under well-watered and drought conditions in a greenhouse experiment. Seedlings were subjected to repeg ated drought cycles, withholding water for either 2 or 3 weeks. We found that hybrid pines grew significantly faster than lodgepole pine under well-watered conditions while retaining the high tolerance to prolonged drought exhibited by lodgepole pine seedlings, a particularly beneficial combination of traits during the critical stage of establishment. We therefore suggest the deployment of hybrid lodgepole pine as a strategy to promote adaptation to hotter and drier future climates.

The article presents the results of the introduction of the North American flora woody species of the pine family (Pinaceae Lindl.) to the European North of Russia. The research purpose is to select the most valuable specimens for introduction into northern conditions from the tree species collection. The research has involved experimental, calculation, analytical, and comparative methods. The conclusion on the result of the introduction of a particular plant was based on the materials of long-term phenological observations, which were carried out according to the method of botanical gardens improved for the conditions of the European North of Russia. As a result of the introduction of North American species of the Pinaceae Lindl. family to the region, a large-scale test of foreign conifers was carried out for the first time; a collection of 2 species of fir, 4 species and 3 forms of spruce, 2 species of pine, and 1 species of Douglas fir was created. Most attention was paid to the introduction of species of the genus Pinus. Positive results of cultivation of Pinus contorta Loud. var. latifolia S. Wats. in the Dendrological garden allowed us to select it for further introduction testing in experimental plantations of the region. On plantations located in the Arkhangelsk and Vologda regions and the Komi Republic, 104.2 ths seedlings were grown from the seed samples, mostly from the northern natural range of this pine. Unfortunately, not all of the breeding material for the North American pine species tested was derived from natural growth sites, which puts it is advisable to use in testing additional samples from the most northern regions of their natural ranges in Canada. Many of these species are of both scientific and practical interest for forestry. To assess the economic value of those introduced species, adaptive capabilities of which in the conditions of the European North according to the results of studies are not in doubt, it is advisable to lay out pilot plantations in the region, similar to the already created plantings of North American lodgepole pine (Pinus contorta Loud. var. latifolia S. Wats.). The concentration of different species like balsam fir (Abies balsamea Mill.), Fraser fir (Abies fraseri (Pursch.) Poir.), American larch (Larix laricina (Du Roi) K.Koch.), Canadian spruce (Picea glauca Voss.), Engelmann’s spruce (Picea engelmannii Engelm.), black spruce (Picea mariana Britt.), Jack pine (Pinus banksiana Lamb.), lodgepole pine (Pinus contorta Loud. var. latifolia S. Wats.), and Douglas fir (Pseudotsuga menziesii Mirb.), involving several seed origins from the most northern regions of its natural range for each, in 2-3 forest areas will allow creating a valuable introduction object; scientific and practical importance of those mentioned cannot be overestimated. For citation: Demidova N.A., Durkina T.M., Gogoleva L.G., Vasiljeva N.N. Representatives of the Pine Family (Pinaceae Lindl.) of the North American Flora in the Collection of the Dendrological Garden named after V.N. Nilov (NRIF). Lesnoy Zhurnal [Russian Forestry Journal], 2021, no. 4, pp. 36–54. DOI: 10.37482/0536-1036-2021-4-36-54Funding: This work was carried out within the framework of the state assignment of the Federal Forestry Agency, registration No. НИОКТР 115042310018.

En el año 2003 se estableció una investigación para evaluar la potencialidad de los sistemas silvopastorales con Pinus contorta (Dougl. ex Loud), en la zona intermedia de la provincia de Coyhaique, Región de Aysén, Patagonia chilena. Para su evaluación se comparan dos diseños silvopastorales con la especie, uno sin manejo y el otro con manejo, versus usos tradicionales de la tierra en Aysén, como es el manejo ganadero con bovinos sobre una pradera naturalizada. Los resultados del estudio arrojan que los sistemas silvopastorales mostraron ser una alternativa para los productores de la Patagonia, dado que les permite obtener ingresos intermedios por la venta de carne y madera proveniente de raleos, mientras que obtendrán un ingreso final por la venta de madera por el componente forestal. La producción de carne en peso vivo indicó que estos sistemas son apropiados para la zona, ya que la producción fue similar a la del sistema ganadero en los primeros dos años de evaluación, aunque el área efectiva del pastoreo se redujo, mostrando la influencia beneficiosa de los árboles en los factores climáticos como viento y temperatura.

1. 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. 2. 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. 3. 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. 4. 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.

The success of invasive plants is influenced by many interacting factors, but evaluating multiple possible mechanisms of invasion success and elucidating the relative importance of abiotic and biotic drivers is challenging, and therefore rarely achieved. We used live, sterile or inoculated soil from different soil origins (native range and introduced range plantation; and invaded plots spanning three different countries) in a fully factorial design to simultaneously examine the influence of soil origin and soil abiotic and biotic factors on the growth of invasive Pinus contorta. Our results displayed significant context dependency in that certain soil abiotic conditions in the introduced ranges (soil nitrogen, phosphorus or carbon content) influenced responses to inoculation treatments. Our findings do not support the enemy release hypothesis or the enhanced mutualism hypothesis, as biota from native and plantation ranges promoted growth similarly. Instead, our results support the missed mutualism hypothesis, as biota from invasive ranges were the least beneficial for seedling growth. Our study provides a novel perspective on how variation in soil abiotic factors can influence plant-soil feedbacks for an invasive tree across broad biogeographical contexts.

Suillus luteus is a common ectomycorrhizal (EM) fungus associated with several Pinus species. It is distributed throughout the Northern Hemisphere and has been introduced into South America and New Zealand. We examined the genetic structure and population biology of S. luteus, which was introduced into Patagonian plantations with Pinus species in Argentina. Overall, 106 samples were collected at 11 geographically separated sites (i.e., Pinus plantations) along a latitudinal gradient in Patagonia (ca. 38°–46° south latitude). Phylogenetic analyses confirmed placement in S. luteus. Genetic analysis demonstrated moderate within-site genetic diversity, but low differentiation between sites. No clear clusters were detected geographically or in relation to host species of Pinus. Our results suggest that the weak genetic structure of the species reflects the short time that has elapsed since the introduction of S. luteus into Patagonia, and its expansion with exotic afforestation there. Moreover, the lack of structure is consistent with a founder effect, suggesting the introduction of a small number of genets that spread throughout all the plantations. Therefore, the high level of gene flow and weak genetic structure observed are probably related to the anthropogenic movement of inoculum associated with forestry practices.

Intra-specific variation in conifers has been extensively studied with respect to defense against herbivores and pathogens. While studies have shown the ability of individual or specific mixtures of compounds to influence insects and microbes, research testing biologically relevant mixtures of defense compounds reflecting intra-specific variation amongst tree populations to enemy complexes is needed. We characterized the variations in lodgepole pine monoterpenes from a progeny trial in western Canada and grouped trees in four clusters using their monoterpene profiles. We then selected 11 representative families across four clusters and amended their entire monoterpene profiles (with the exception of β-phellandrene) in media to determine how representative families affect the performance of the mountain pine beetle or its fungal symbiont. We placed adult beetles or inoculated fungus on the amended media and measured beetle performance and fungal growth as a proxy to host suitability. We found that different clusters or families differentially influenced beetle or fungal responses. However, monoterpene profiles of trees suitable to the beetle or the fungus were dissimilar. These outcomes reflect a co-evolutionary arms-race between the host and the bark beetle-fungus complex, which has resulted in the production of complementary defense metabolites among different pine populations to enhance tree survival.

The sister species, lodgepole pine (Pinus contorta var. latifolia Engelm.) and jack pine (Pinus banksiana Lamb), face pressures from a multitude of biotic agents, including mountain pine beetle (Dendroctonus ponderosae Hopkins, 1902) and their pathogenic fungal associates (e.g., Grosmannia clavigera (Rob.-Jeffr. & R.W. Davidson) Zipfel, Z.W. de Beer & M.J. Wingf.), mistletoe (Arceuthobium americanum Nutt. ex A.Gray), and the pathogen causing western gall rust (Cronartium harknessii E. Meinecke). Here, we report new stem tissue transcriptome resources developed for lodgepole and jack pines subjected to these biotic stresses. The annotated transcriptomes were compared to determine species-specific responses to the necrotrophic G. clavigera and the biotrophic Cronartium harknessii. We focused on chitinases, a family that includes members with well-documented roles in defense. Putative chitinase family members were identified via annotation, sequence similarity to reference chitinase genes, phylogenetic analyses, and in silico motif characterization. RNA-Seq revealed marked differences in the responses of lodgepole and jack pine chitinases to G. clavigera and Cronartium harknessii. The potential for adaptive variation in chitinases was investigated by assessing the level of genetic differentiation between and within lodgepole and jack pines using single nucleotide polymorphisms within chitinases. These analyses illustrate the potential of combining transcriptomic and genotyping resources to investigate genotype–phenotype correlations for non-model species.

Gravel mining is prevalent in forest landscapes across Canada, typically resulting in complete loss of vegetation and topsoil. Despite such extreme disturbance, lodgepole pine (Pinus contorta Dougl. var. latifolia Engelm.) trees are thriving at unreclaimed gravel pits located in central-interior British Columbia, possibly owing to, at least in part, the association of pine trees with their endophytic bacteria. To test this possibility, several bacterial strains were previously isolated from pine trees growing at these pits; of the strains isolated, 14 were identified as effective nitrogen fixers. In this study, we evaluated the inoculation effect of these 14 strains on lodgepole pine growth under nitrogen-poor conditions. Eighteen months after sowing and inoculation, each strain had colonized the rhizosphere and internal tissues of pine seedlings and had significantly enhanced their length (24%–65%) and biomass (100%–300%). Notably, three Pseudomonas strains increased pine seedling length by 1.6-fold and biomass by 4-fold. Most strains also demonstrated substantial potential to promote plant growth via phosphorus solubilization, siderophore production, 1-aminocyclopropane-1-carboxylic acid (ACC) deaminase activity, indole-3-acetic acid production, lytic enzyme activity, and catalase activity. Our results suggest that such nitrogen-fixing bacteria could be sustaining pine growth on bare gravel, indicating a possible ecological association that may explain natural tree regeneration in such a disturbed ecosystem.

Many conifers have distributions that span wide ranges in both biotic and abiotic conditions, but the basis of response to biotic stress has received much less attention than response to abiotic stress. In this study, we investigated the gene expression response of lodgepole pine (Pinus contorta) to attack by the fungal pathogen Dothistroma septosporum, which causes Dothistroma needle blight, a disease that has caused severe climate-related outbreaks in northwestern British Columbia. We inoculated tolerant and susceptible pines with two D. septosporum isolates and analyzed the differentially expressed genes (DEGs), differential exon usage, and coexpressed gene modules using RNA-sequencing data. We found a rapid and strong transcriptomic response in tolerant lodgepole pine samples inoculated with one D. septosporum isolate, and a late and weak response in susceptible samples inoculated with another isolate. We mapped 43 of the DEG- or gene module–identified genes to the reference plant-pathogen interaction pathway deposited in the Kyoto Encyclopedia of Genes and Genomes database. These genes are present in PAMP-triggered and effector-triggered immunity pathways. Genes comprising pathways and gene modules had signatures of strong selective constraint, while the highly expressed genes in tolerant samples appear to have been favored by selection to counterattack the pathogen. We identified candidate resistance genes that may respond to D. septosporum effectors. Taken together, our results show that gene expression response to D. septosporum infection in lodgepole pine varies both among tree genotypes and pathogen strains and involves both known candidate genes and a number of genes with previously unknown functions. [Formula: see text] Copyright © 2021 The Author(s). This is an open access article distributed under the CC BY-NC-ND 4.0 International license .

Exotic conifers are rapidly spreading in many regions of New Zealand, as well as in many other countries, with detrimental impacts on both natural ecosystems and some productive sector environments. Herbicides, in particular the active ingredient triclopyr, are an important tool to manage invasive conifers, yet there is a paucity of information that quantifies the amount of herbicide required to kill trees of different sizes when applied as a basal bark treatment. Two sequential experiments were conducted to define the amount of triclopyr required to kill individual invasive lodgepole pine (Pinus contortaDouglas ex Loudon), trees of different sizes when applied in a methylated seed oil to bark (either the whole stem or base of the tree) and to determine which tree size variates (height, diameter at breast height [DBH], crown diameter [CD]) or derived attributes (crown area, crown volume index) best characterized this dose–response relationship. The outcomes of the dose–response research were compared with field operations where triclopyr was applied to the bark of trees from an aerial platform. Applying the herbicide to the whole stem, as opposed to the base of the tree only, significantly increased treatment efficacy. The tree size variates DBH, CD, crown area, and crown volume index all provided good fits to the tree mortality data, with >91% prediction accuracy. Of these variates, CD provided the most practical measure of tree size for ease of in-field calculation of dose by an operator. Herbicide rates used in field operations were seven to eight times higher than lethal doses calculated from experimental data. Our results highlight the potential for substantial reductions in herbicide rates for exotic conifer control, especially if dose–response data are combined with remotely sensed quantitative measurements of canopy area or volume using new precision technologies such as unmanned aerial vehicles.

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.

The mountain pine beetle, Dendroctonus ponderosae, has infested over ~16 Mha of pine forests in British Columbia killing >50% of mature lodgepole pine, Pinus contorta, trees in affected stands. At present, it is functionally an invasive species in Alberta, killing and reproducing in evolutionarily naive populations of lodgepole pine (P. contorta), novel jack pine (P. banksiana), and their hybrids. The entomopathogenic fungus Beauveria bassiana has shown some potential as a biocontrol agent of several bark beetle species. In this study, nine isolates of B. bassiana were examined for insect virulence characteristics, including conidiation rate, pigmentation, and infection rate in laboratory-reared D. ponderosae, to assess for their potential as biocontrol agents. The strains were categorized into three phenotypic groups based on pigmentation, conidial density, and myceliation rate. Virulence screening utilizing insect-based agar medium (D. ponderosae and European honeybee Apis mellifera carcasses) revealed no difference in selection of fungal growth. However, infection studies on D. ponderosae and A. mellifera showed contrasting results. In vivo A. mellifera infection model revealed ~5% mortality, representing the natural death rate of the hive population, whereas laboratory-reared D. ponderosae showed 100% mortality and mycosis. The LT50 (median lethal time 50) ranges from 2 to 5 ± 0.33 days, and LT100 ranges from 4 to 6 ± 0.5 days. We discuss the selective advantages of the three phenotypic groups in terms of virulence, pigmentation, conidial abundance, and tolerance to abiotic factors like UV and host tree monoterpenes. These results can further provide insights into the development of several phenotypically diverse B. bassiana strains in controlling the spread of the invasive D. ponderosae in Western Canada. • Three B. bassiana morphotype groups have been demonstrated to kill D. ponderosae. • A range of effective lethal times (LT50 and LT100) was established against D. ponderosae. • Variable tolerance to UV light and pine monoterpenes were observed in B. bassiana.

Upstream identification of wood properties using non-destructive testing methods such as acoustic velocity (AV) measurements is important for optimizing allocation of wood to mills or products. We evaluated the effectiveness of field AV measurement tools in predicting lodgepole pine wood stiffness (modulus of elasticity, MOE) as measured by Silviscan on wood samples. AV was measured on trees and logs from six sites in Alberta and British Columbia. We evaluated the effect on MOE estimation of calculating averages of the adjustment factor k and of green density (GD) at different spatial scales from individual tree to population. The effect of using forest inventory variables on MOE prediction were also examined. Prediction of tree-level MOE from tree-level measurements of AV, k and GD resulted in R2 values of 0.59. Using estimates of k and GD averaged at plot, site or population scales significantly diminished the R2 of the MOE predictions at tree level. Predicting MOE at plot or stand level from corresponding averages of AV, k and GD gave R2 values >0.8. Including inventory variables in tree-level MOE predictions increased the R2 to 0.62. AV measurements can give operationally useful estimates of MOE in lodgepole pine trees at the stand level.

We studied the recovery of tree- and stand-level carbon (C) storage in a lodgepole pine (Pinus contorta Dougl. ex Loud. var. latifolia Engelm.) forest in northern British Columbia that experienced substantial (∼83%) mortality in 2006–2007 (total loss by 2013 = 86%) during a severe mountain pine beetle (MPB; Dendroctonus ponderosae Hopkins, 1902) infestation. Earlier work suggested that this forest recovered positive annual C storage 3 years after attack based on eddy covariance measurements. We sought to confirm these results by examining C storage in surviving pine trees using tree core analysis. Average growth release of surviving lodgepole pine trees was 392% (range of –53% to 2326%) compared with mean decadal growth prior to MPB attack. Nearly 97% of trees underwent a growth release, considerably higher than the 15%–75% reported for lodgepole pine in previous studies. Mean annual stem C storage of the surviving trees in this study was highly correlated (r = 0.88) with 10 years of annual net ecosystem productivity estimates made using the eddy covariance technique, indicating that surviving lodgepole pine remain an important part of C recovery after MPB attack. Mean annual stem C storage was also highly correlated (r = 0.92) with the cumulative percentage of downed stems per hectare at the site, suggesting that increased availability of resources is likely assisting the growth release.

The ascomycete fungi Dothistroma septosporum (Dorogin) M. Morelet and D. pini Hulbary are both responsible for Dothistroma needle blight disease (DNB) on pines. Using a DNA-barcoding approach, we c...

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.

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.

Wildfire is a ubiquitous disturbance agent in subalpine forests in western North America. Lodgepole pine ( Pinus contorta var. latifolia), a dominant tree species in these forests, is largely resilient to high-severity fires, but this resilience may be compromised under future scenarios of altered climate and fire activity. We investigated fire occurrence and post-fire vegetation change in a lodgepole pine forest over the past 2500 years to understand ecosystem responses to variability in wildfire and climate. We reconstructed vegetation composition from pollen preserved in a sediment core from Chickaree Lake, Colorado, USA (1.5-ha lake), in Rocky Mountain National Park, and compared vegetation change to an existing fire history record. Pollen samples ( n = 52) were analyzed to characterize millennial-scale and short-term (decadal-scale) changes in vegetation associated with multiple high-severity fire events. Pollen assemblages were dominated by Pinus throughout the record, reflecting the persistence of lodgepole pine. Wildfires resulted in significant declines in Pinus pollen percentages, but pollen assemblages returned to pre-fire conditions after 18 fire events, within c.75 years. The primary broad-scale change was an increase in Picea, Artemisia, Rosaceae, and Arceuthobium pollen types, around 1155 calibrated years before present. The timing of this change is coincident with changes in regional pollen records, and a shift toward wetter winter conditions identified from regional paleoclimate records. Our results indicate the overall stability of vegetation in Rocky Mountain lodgepole pine forests during climate changes and repeated high-severity fires. Contemporary deviations from this pattern of resilience could indicate future recovery challenges in these ecosystems.

1. Significant research has been devoted to the development of semiochemical repellents for mountain pine beetle Dendroctonus ponderosae Hopkins. Several formulations of the beetle's antiaggregation pheromone verbenone (4,6,6‐trimethylbicyclo[3.1.1]hept‐3‐en‐2‐one) are registered to protect pine trees and stands. 2. We compared the efficacy of verbenone and Verbenone Plus for protecting individual lodgepole pine Pinus contorta Dougl. ex Loud. and small stands of P. contorta from mortality attributed to D. ponderosae . Verbenone Plus is a blend of semiochemicals [(−)‐verbenone, acetophenone, and (E )‐2‐hexen‐1‐ol + (Z )‐2‐hexen‐1‐ol] originally developed to inhibit aggregation of western pine beetle D. brevicomis LeConte. It is the only effective semiochemical repellent that protects ponderosa pine P. ponderosa Dougl. ex Laws. from mortality attributed to D. brevicomis , but has yet to be registered or commercialized. 3. Both verbenone and Verbenone Plus were effective for protecting individual P. contorta and small stands of P. contorta from mortality attributed to D. ponderosae . Verbenone alone was as effective as Verbenone Plus. 4. An obstacle to the broader use of semiochemical repellents for the management of bark beetles is the relatively high cost and specificity of the semiochemicals involved. A semiochemical repellent with efficacy across multiple bark beetle species and hosts, such as Verbenone Plus, would be desirable, assuming the additional components are not cost‐prohibitive or limit registration.

Herbivores modulate the structure and composition of plant communities, including plant invasions. This is conditioned by plant palatability which can be reduced by its chemical or physical traits. The effects that ungulates browsing has on pine invasions are variable and the empirical evidence on the causes of this variability is scarce. We experimentally explored how sheep browsing preference varies between seedlings of pine species with different invasiveness; Pinus contorta (high invasiveness), P. ponderosa (medium invasiveness), P. radiata (medium invasiveness) and P. jeffreyi (low invasiveness). Secondly, we quantified anti-herbivory chemical compounds and physical traits of these species and related them with sheep preference observed. The browsing incidence of P. contorta was 68%, P. ponderosa 58%, P. radiata 29%, and P. jeffreyi 84%. Among anti-herbivory traits analyzed, α-pinene concentration had a negative effect on the probability of a terminal bud being browsed and on browsing intensity. Meanwhile, foliar toughness was negatively related to browsing intensity and water concentration was positively related to browsing intensity. Also, the most invasive species, P. contorta, was highly damaged. Thus, sheep herbivory could be slowing pine invasion rate; suggesting that could be considered a tool to control early invasions, especially for this particular species.

Three kinds of tree bark extractives from trembling aspen (Populus tremuloides) (AE), lodgepole pine (Pinus contorta) (LE), and western red cedar (Thuja plicata) (RE) were produced and used as antioxidants. The antioxidation activities of those extractives were evaluated and compared with commercial antioxidant butylated hydroxytoluene (BHT). The obtained extractives were applied as natural photostabilizers to produce wood flour/polypropylene (WF/PP) composites at an addition level of 2 wt%. The composites were subjected to QUV accelerated weathering tester for a total of 1200 h. The changes in surface color, surface morphology, and surface chemistry of composites were investigated during the exposure. The results showed that all bark extractives alleviated the photodegradation of composites successfully. Among them, RE was proved to be the most effective. It exhibited excellent UV absorption and similar free-radical inhibition ability compared with BHT. Consequently, composites containing RE exhibited less discoloration, less severe surface cracking, and less changes in surface chemistry during the whole weathering process. Overall, these results revealed that RE had great potential as a natural photostabilizer in polymer systems.

There is an expected decrease in the short to mid-term timber supply of lodgepole pine (Pinus contorta var. latifolia) in Alberta, Canada due to the impacts of past fires and the mountain pine beetle outbreak. Commercial thinning is a potential option for increasing mid-term sawlog timber supply by decreasing the time needed for individual trees to reach merchantable size, through providing access to fiber earlier in the rotation, and from the ability to keep mature stands on the stump longer. In the current study, we evaluated the effects of commercial thinning in stands of different ages. Stands less than 70 years old at thinning were classified as commercial thinning (CT), whereas stands greater than 70 years at thinning were classified as salvage thinning (ST). Tree growth and survival were measured every five years from 1996 to 2016 in both thinned and unthinned stands. Fifteen years post-thinning, stand volume was less in the thinned stands relative to the unthinned controls. However, when volume removed at thinning was considered, volume gain from thinned stands was greater than that from the controls in both thinning treatments. Individual tree DBH and live crown ratio were also greater in thinned stands relative to controls. Thinning favored diameter gain mostly in the medium-sized trees in CT but both small to medium-sized trees in ST. Thinning reduced mortality in all stands relative to the controls and a maximum of 50% reduction in mortality was observed in CT. Overall, the findings suggest that natural lodgepole pine stands may respond to commercial thinning with a net gain in cumulative volume at final harvest.

Understanding tree vulnerability to freezing temperatures will help resource managers to mitigate the effects of climatic variability. To test the effectiveness of tissue dehardening curves to represent whole seedling responses, we tested the cold tolerance of 4 conifer species to a range of freezing temperatures, − 15, − 25, and − 35 °C in late March, and − 5, − 10, and − 15 °C in early May. Results show that, after a 30-min exposure to freezing temperatures, needle mortality started at − 25 °C in late March (early spring dehardening) before budbreak activation and at − 10 °C in early May (late spring dehardening) when pine buds had flushed and white spruce (Picea glauca) buds had started swelling. Freezing temperatures delayed the timing of budbreak; increased needle, terminal bud, and seedling mortality; and reduced shoot growth and number of top laterals. White spruce and lodgepole pine (Pinus contorta) were more sensitive to freezing in terms of needle, terminal bud, and seedling mortality and growth reduction. The effects of March freezing were generally detected in the first year, whereas those of the May freezing were still evident in the second year on the shoot growth of black spruce (Picea mariana) and jack pine (Pinus banksiana). Although they seem relatively hardier based on first year post-freezing assessments (i.e., less immediate physical damage from freezing), black spruce and jack pine may have sustained more internal or physiological shock/damage. Our study results support the use of tissue cold hardiness–forcing temperature relationships to assess the risk of freezing damage for conifer seedlings.

Ectomycorrhizal fungi supply their plant partners with nitrogen but can also retain substantial amounts. The concentration of nitrogen in the soil and the amount of carbon supplied from the host seem to influence the proportion of N retained by the fungus. In an experiment designed to determine whether differential supply of nitrogen to two plants influenced nitrogen transfer from fungus to plant within a mycorrhizal network, we observed rapid, substantial loss of nitrogen from pine seedlings. The loss occurred when the mycorrhizal fungus experienced a sudden increase in nitrogen supply. We grew Pinus contorta seedlings in association with Suillus tomentosus in low-nitrogen microcosms where some nitrogen was accessible only by hyphae. After 70 days, foliage of some seedlings was treated with nitrogen. Three days later, hyphal nutrient media were replaced with water or a solution containing nitrogen. Foliar treatment did not affect nitrogen transfer by the fungus to shoots, but by day 75, seedling nitrogen contents had dropped by 60% in microcosms where nitrogen had been added to the hyphal compartments. Those seedlings retained only 55% of the nitrogen originally present in the seed. Loss of nitrogen did not occur if water was added or the hyphae were severed. Because of the severing effect, we concluded that S. tomentosus triggered the loss of seedling nitrogen. Nitrogen may have been lost through increased root exudation or transfer to the fungus. Access to nitrogen from nutrient-rich germinants would benefit rhizosphere microorganisms, including ectomycorrhizal fungi colonizing pine from spores after wildfire. Electronic supplementary material The online version of this article (10.1007/s00572-020-00959-7) contains supplementary material, which is available to authorized users.

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.

Disturbances are frequent events across the Canadian boreal forest and can affect both below‐ and above‐ground ecosystem processes. How disturbances change below‐ground soil fungal communities and in‐turn affect pine establishment and performance is poorly understood. Such understanding has become increasingly important in light of observed changes in disturbance regimes in recent years due to climate change. We used a greenhouse experiment to determine how soil inoculum collected from lodgepole pine stands undisturbed (control) or disturbed by fire, mountain pine beetle outbreak, logging and salvage logging affect pine seedling performance in western Canada. We first characterized whether fungal communities of seedling roots change as a function of inoculum source, and then determined whether changes in fungal community composition impact pine seedling performance (biomass and height). Root fungal communities of pine seedlings from logged and salvage logged disturbances differed from their respective paired controls, while soils from natural disturbances (fire and beetle outbreak) did not. Among disturbances, the pine root fungal communities of fire and salvage logged disturbances differed. In parallel to the root fungal communities, seedling performance also decreased when comparing logging and salvage logging disturbances to paired controls. Among disturbance treatments, seedlings from the salvage logged disturbance did not grow as big as seedlings inoculated with soils from burned forests. Synthesis and application. Our findings indicate that anthropogenic disturbances (logging and salvage logging) can have cross‐generational impacts on pine seedling performance, through functional shifts in seedling root fungal community structure. Furthermore, the impacts of soil fungi on pine seedlings appear to be pronounced following salvage logging, stressing the importance of compound disturbance events. These findings may be important to land managers considering clear‐cut logging or salvage logging in pine forests, particularly where soil biotic communities are likely to be one of the predominate factors in pine establishment.

The large-scale introduction of lodgepole pine (Pinus contorta ssp. latifolia, LP) into Swedish forests was initiated around 1970, and currently 520,000 hectares of the forest land are dominated by the species. Even though the tree has mostly serotine cones, adapted to open after forest fires, it has proved able to self-regenerate in Sweden. This study is the first to present, scientifically, the extent of self-regeneration covering the whole current range of LP-forests in Sweden. LP-saplings were recorded for three years, 2015–2017, in 8194 subplots distributed over 214 randomly selected LP-stands from latitude 59.6–66.9°N and altitude 88–710 m asl. Of all subplots, 3% contained LP-saplings, and regeneration was found in 53% of all stands. The probability of finding LP-saplings was significantly dependent on the plots’ distance from the edge of the LP-stand, and 78% of all saplings were found within and up to 15 m beyond the stand edge. Most, 63%, of the plots with LP were found on disturbed ground such as wheel tracks, roadsides and where there had been site preparation. The results show that LP can naturally spread under a range of conditions in the studied region. The regeneration is, however, concentrated in particular stands and should be possible to control with monitoring programmes and measures to eradicate self-dispersed trees in unwanted areas.

For tree seedlings in boreal forests, ectomycorrhizal (EM) fungal networks may promote, while root competition may impede establishment. Thus, disruption to EM fungal networks may decrease seedling establishment owing to the loss of positive interactions among neighbors. Widespread tree mortality can disrupt EM networks, but it is not clear whether seedling establishment will be limited by the loss of positive interactions or increased by the loss of negative interactions with surrounding roots. Depending upon the relative influence of these mechanisms, widespread tree mortality may have complicated consequences on seedling establishment, and in turn, the composition of future forests. To discern between these possible outcomes and the drivers of seedling establishment, we determined the relative importance of EM fungal networks, root presence, and the bulk soil on the establishment of lodgepole pine and white spruce seedlings along a gradient of beetle-induced tree mortality. We manipulated seedling contact with EM fungal networks and roots through the use of mesh-fabric cylinders installed in soils of lodgepole pine forests experiencing a range of overstorey tree mortality caused by mountain pine beetle. Lodgepole pine seedling survival was higher with access to EM fungal networks in undisturbed pine forests in comparison with that in beetle-killed stands. That is, overstorey tree mortality shifted fungal networks from being a benefit to a cost on seedling survival. In contrast, overstorey tree mortality did not change the relative strength of EM fungal networks, root presence and the bulk soil on survival and biomass of white spruce seedlings. Furthermore, the relative influence of EM fungal networks, root presence, and bulk soils on foliar N and P concentrations was highly contingent on seedling species and overstorey tree mortality. Our results highlight that following large-scale insect outbreak, soil-mediated processes can enable differential population growth of two common conifer species, which may result in species replacement in the future.

Challenges to the mid-term timber supply in interior British Columbia resulting from losses to insect outbreaks and wild-fires have prompted interest in silvicultural interventions to hasten growth of regenerating forests, including high-density(overstocked), height-repressed stands of lodgepole pine (Pinus contorta Dougl. ex Loud. var. latifolia Engelm.). A trialwas established in a dense (4000 to 30 000 trees ha-1) 36-year-old wildfire-origin pine stand in the Cariboo-Chilcotinregion of interior BC to assess the efficacy of thinning, fertilization and a combination of the two treatments for improv-ing tree growth and stand development. Height and diameter of the 36 crop trees in each plot was greater in fertilizedplots then control plots for six years following fertilization, while the height and diameter responses to thinning and thin-ning + fertilization were sustained throughout the 18-year measurement period. Cumulative basal area and volumegrowth of crop trees and all stand trees over the 18-year period was significantly greater in thinned and thinned + fertil-ized treatments than control plots. Projections based on the TASS growth and yield model suggested that total stand vol-umes in thinned plots would surpass that of the control at age 74 and would surpass that of the fertilized treatment at 89years. Unthinned stands (i.e., control and fertilized treatments), did not accumulate significant merchantable volume; inthinned stands (i.e., thinned and thinned + fertilized treatments), the minimum harvest criteria of 80 m3 ha-1 for pine-leading stands in this region was reached at age 102 years. Repeated fertilization would be needed to further hasten standdevelopment. The potential long-term benefits of these silvicultural interventions for timber supply must be weighedagainst treatment costs and risks of mortality related to wildfire, insects and disease. These results highlight the value oflong-term research trials for informing silviculture decisions.

Mixing tree species could be a silviculture model that allows early harvest of short-rotation trees, while longer-rotation crop trees remain in the stand. We examined the effects on growth ...

This study was conducted to quantify growth responses of three major commercial conifer species (lodgepole pine (Pinus contorta Douglas ex Loudon var. latifolia Engelm. ex S. Watson), interior Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco var. glauca (Beissn.) Franco), and spruce (white spruce (Picea glauca (Moench) Voss) and hybrid spruce (Picea engelmannii Parry ex. Engelm. × Picea glauca (Moench) Voss × Picea sitchensis (Bong.) Carrière))) to various fertilizer blends in interior British Columbia, Canada. Over 25 years, growth-response data were repeatedly collected across 46 installations. The fertilizer blends were classified into three groups: nitrogen only; nitrogen and sulfur combined; and nitrogen, sulfur, and boron combined. The growth responses for stand volume, basal area, and top height were calculated through absolute and relative growth rate ratios relative to a controlled group. Fertilizer blend, inverse years since fertilization, site index, stand density at fertilization, and their interactions with the fertilizer blend were used as explanatory variables. The magnitude and significance of volume and basal area growth responses to fertilization differed by species, fertilizer-blend groups, and stand-condition variables (i.e., site index and stand density). In contrast, the response in top height growth did not differ among fertilization blends, with the exception of the nitrogen and sulfur fertilizer subgroup for lodgepole pine. The models developed in this study will be incorporated into the current growth and yield fertilization module (i.e., Table Interpolation Program for Stand Yields (TIPSY)), thereby supporting guidance of fertilization applications in interior forests in British Columbia.

Western gall rust (Cronartium harknessii (J.P. Moore) E. Meinecke) is a pathogen that affects lodgepole pine (Pinus contorta Douglas ex Loudon var. latifolia Engelm. ex S. Watson) and has the potential to reduce lumber product yields derived from stands managed for the commercial production of timber. A dataset containing repeated measurements from 7775 trees located within the province of British Columbia, Canada, was used to develop equations that predict annual rates of stem infection, post-infection mortality, and the location of large stem galls. Results showed that annual rates of infection peaked between 3 and 5 years following stand establishment. Few new infections were predicted to occur beyond the stand age of 15 years. For an individual tree, the probability of first infection increased as its height increased relative to stand top height. The rate of mortality increased with the number of stem infections and was highest among trees infected within the first 3 years following planting. Mortality rates decreased thereafter, with many trees likely to survive to rotation. The equations predicting rust incidence and mortality were added to the Tree and Stand Simulator (TASS), an individual-tree growth model used within the province. A final equation predicting the location of large galls was added and allows TASS to account for losses due to the removal of stem defects during lumber manufacturing.

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.

Global commercial and recreational transport may lead to the unintentional invasion of insect species, which in turn may pose a threat to native organisms. In this study, we aimed to assess whether the economically important pest of Pinus sylvestris L., moth Dendrolimus pini L. (DP), is able to feed on nine other pine species, and how this will affect its survival, performance, growth, and development. We carried out food choice tests and a no-choice laboratory feeding experiment. We found that this insect mostly preferred its prime host, but also Pinus cembra L., Pinus contorta Douglas ex Loudon, Pinus nigra J.F.Arnold, and Pinus ponderosa Douglas ex C.Lawson. The performance test revealed a host-specific response of DP to the host plant. This response was manifested in a large variation in body mass as well as in a decrease or increase in life-history traits, such as fecundity, and wing morphology parameters. However, the larvae’s choice of particular hosts corresponded to the results of the performance test. Larvae more willingly selected food allowing better results in their performance. Larvae achieved better values of growth and development when fed on European and North American pine species or on species with two- and three-needle fascicles. In addition, attractants and repellents in needles of different pine species were chemically analyzed. Variations in the secondary metabolite composition as well as the specific leaf area of different pine species effectively explained the results found in the insects, but the content of sugars and nitrogen remains to be elucidated. We speculate that DP poses a serious threat to large areas of pine forests, if transferred, as it can survive and develop on many economically important tree species in North America and Europe.

Warming temperatures are allowing native insect herbivores to expand into regions that previously exceeded their thermal tolerance, encounter new host species, and pose significant threats to native communities. However, the dynamics of these expansions remain poorly understood, particularly in the extent to which outbreaks remain reliant on emigration from historical hosts or are driven by local reproduction within novel hosts in the expanded range. We tested these non-mutually exclusive hypotheses using spatially explicit data on mountain pine beetle (Dendroctonus ponderosae), which historically undergoes intermittent outbreaks in low-elevation lodgepole pine (Pinus contorta), but is now causing severe mortality in a high-elevation endangered species, whitebark pine (Pinus albicaulis). We compiled data from 2000 to 2019 across British Columbia, Canada, at 1-km2 resolution, and analyzed spatiotemporal patterns of beetle infestations, lodgepole pine distributions, expansion into habitats dominated by whitebark pine, and the likelihood of future outbreaks in all pine communities under simulated conditions. Overall, we found strong support for the hypothesis of emigration from the historical host species continuing to be a major driver of outbreaks in the more recently accessed host. First, beetle population pressure was consistently the best predictor of infestation severity in both lodgepole and whitebark pine, and appeared to be mostly unidirectional from lodgepole to whitebark pine. Second, infestations in lodgepole pine were of a longer duration than those in whitebark pine, which appeared too brief to sustain transitions from endemic to eruptive dynamics. Furthermore, resource depletion appears to drive emigration from lodgepole pine, whereas in whitebark pine drought appears to favor establishment of immigrants although bioclimatic factors and stand structure preclude self-sustaining outbreaks. Finally, we project that most pine in British Columbia will be at risk in the event of a new major outbreak. We describe implications for conserving and protecting whitebark pine and to other climate-driven range expansions.

Successful host plant colonization by tree-killing bark beetle-symbiotic fungal complexes depends on host suitability, which is largely determined by host defense metabolites such as monoterpenes. Studies have shown the ability of specific blends of host monoterpenes to influence bark beetles or their fungal symbionts, but how biologically relevant blends of host monoterpenes influence bark beetle-symbiotic fungal interaction is unknown. We tested how interactions between two host species (lodgepole pine or jack pine) and two fungal symbionts of mountain pine beetle (Grosmannia clavigera or Ophiostoma montium) affect the performance of adult female beetles in vitro. Beetles treated with the propagules of G. clavigera or O. montium or not treated (natural fungal load) were introduced into media amended with a blend of the entire monoterpene profile of either host species and beetle performance was compared. Overall, host blends altered beetle performance depending on the fungal species used in the beetle amendment. When beetles were amended with G. clavigera, their performance was superior over beetles amended with O. montium in either host blend. Furthermore, G. clavigera-amended beetles performed better in media amended with host blends than without a host blend; in contrast, O. montium-amended beetles performed better in media without a host blend than with a host blend. Overall, this study showed that host defense metabolites affect host suitability to bark beetles through influencing their fungal symbionts and that different species of fungal symbionts respond differentlly to host defense metabolites.

We present an account of Rhizopogon introduced from plantings of exotic pine plantations in Argentine Patagonia. Nuc rDNA internal transcribed spacer ITS1-5.8S-ITS2 (ITS) and nuc 28S rDNA (28S) sequences were used to identify specimens from Argentina and examine their relationships with geographically different Rhizopogon species. Based on phylogenetic analyses, we confirm that four species of Rhizopogon occur in pine plantations across Patagonia. Several Rhizopogon collections from Pinus ponderosa plantations across different provinces cluster with R. arctostaphyli, a species within R. subg. Amylopogon. The majority of Patagonian Rhizopogon, however, form three different lineages in R. subg. Roseoli. The first of these, R. roseolus sensu Trappe, includes numerous collections from Pinus ponderosa, P. contorta, and P. radiata stands of North American affiliation. The second, R. roseolus sensu Martin and Garcia from P. ponderosa plantations, clusters in clade IIIa of the R. roseolus complex, which also includes the holotype collection of R. mohelnensis from the Czech Republic. The third species in R. subg. Roseoli, and fourth species overall from Patagonia, is R. granuloflavus from Pinus ponderosa plantations. Multiplex polymerase chain reaction (PCR) of numerous Roseoli samples failed to produce an amplicon indicative of either Japanese or New Zealand shoro.

Understory vegetation structure and its relationship with forest canopies and site conditions are important determinants of carbon stocks, wildlife habitat, and fuel loading for wildland fire assessments. Comprehensive studies are needed to assess these relationships through the use of consistently collected field-based data. One approach to achieve this is to make use of preexisting forest inventory data to estimate understory vegetation height and cover from site and overstory attributes. In this study, overstory, understory, and abiotic data describing site conditions were obtained from over 6700 Forest Inventory and Analysis (FIA) fixed radius plots collected between 2000 and 2012 to assess how understory vegetation cover and height vary with overstory attributes and site characteristics. The focus was restricted to four common forest types including lodgepole pine (Pinus contorta var. latifolia), Douglas-fir (Pseudotsuga menziesii), ponderosa pine (Pinus ponderosa), and grand fir (Abies grandis) found on approximately 43 million hectares in the western United States. Random Forest regression classification trees were developed for cover and height of shrub and herb understories as a function of field-measured predictor variables. Separate analyses were undertaken for the Pacific Northwest (PNW) and the Interior West (IW) Forest Inventory and Analysis (FIA) regions. Models developed from the IW data generally performed better and the OOB (out-of-bag) percent variance explained varied from 8.08% for forb height to 39.24% for shrub height. For the PNW data, percent variance explained ranged from 13.82% for forb height to 27.4% for shrub height. Percent variance explained values were higher in all corresponding models for the IW than PNW, except for forb and grass height. Differences in model performance were smallest in the case of forb cover (27.17% vs. 26.15%) and greatest in the case of percent shrub cover (30.92% vs. 15.53%) for IW and PNW models, respectively. Cover models within each dataset performed better, on average, than their associated height models. The most influential variables for predicting understory cover and height were ones representing overstory conditions and conform to ecological expectation corroborated by many studies examining the influence of forest overstories on understory vegetation dynamics. Several variables, including aspect, slope, and stand disturbance and treatment, were not important and contrary to expectation. Predicting understory vegetation attributes to aid assessments of carbon, fuel, and wildlife habitat may be more generalizable across forests of the western U.S. using standardized national inventory data in conjunction with improved measurements.

Survival and growth of Siberian larch (SL, Larix sibirica Ledeb.) were compared with those of three conifer species native to Alberta, Canada: lodgepole pine (LP; Pinus contorta var. latifolia Engelm. ex S. Watson), white spruce (WS, Picea glauca (Moench) Voss), and jack pine (JP, Pinus banksiana Lamb.) at 12, 10, and 3 trial locations, respectively. The average data age was 18 years (range: 3 to 27 years). Survival of SL averaged 4.2% and 6.5% worse than LP and WS, respectively, while it was 5% better than JP. SL grew 25%, 94%, and 23% taller than LP, WS, and JP, respectively. Stem forking rates were similar between SL and LP, WS, and JP. The best seed sources for Alberta were mature trees established in Alberta and Saskatchewan of unknown initial provenances. The Russian Altai Mountain source grew well at high elevations, while the Finnish Raivola performed well in the northern, low-elevation area. Open-pollinated progeny tests of 58 families planted in five diverse locations yielded individual-tree narrow-sense heritabilities and family mean heritabilities for height at age 15 of 0.15 and 0.59, respectively. The type B between-site genetic correlation was 0.44, indicating a strong genotype × environment interaction. SL has performed well in Alberta, and its growth can be further improved by selection and breeding from appropriate seed sources.

Seed orchards play a critical role in reforestation, but often struggle to consistently produce enough seed. A mixture of the less polar gibberellins, GA4 and GA7, has been shown to promote flowering in Pinaceae species, but the responses vary significantly based on species, genotype and provenance, application method, timing and dosage. Previous studies with lodgepole pine (Pinus contorta Dougl. var. latifolia Engelm.) suggest that the response to exogenous GA4/7 varied among genotypes. Gibberellins also play a critical role in seed development and germination but it remains unclear how exogenous GA4/7 will affect seed quality. We used 25 lodgepole pine genotypes in a commercial seed orchard in Alberta, Canada to explore the impact of genotype and geographic origin on the efficacy and optimal timing of GA4/7 stem injections to promote female cone bud initiation, cones and seed production. Overall, we found that GA4/7 stem injections promoted female conelets in 40% of genotypes included in this study. For these 10 genotypes, we found that the latitude of their provenance was positively correlated with later effective application times. Of the remaining 15 genotypes, five were hypersensitive to exogenous GA4/7 showing needle discolouration and loss, and 10 genotypes showed no response. In general, stem-injected GA4/7 did not affect the number of seeds per cone while two genotypes showed a reduced seed yield and one genotype showed increased seed yield per cone. Seed weights were not affected by the stem-injected GA4/7, while the July application date resulted in higher germination rates compared to control trees.

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

This study investigated the origins and spread patterns of Heterobasidion root disease in three Pinus contorta Dougl. ex Loudon plantations established on forest and agricultural land and subjected to three different management scenarios. Trees with decline symptoms and stumps remaining from the previous rotation were sampled for fungal isolations. Ten isolates of Heterobasidion parviporum Niemelä & Korhonen and 425 of Heterobasidion annosum (Fr.) Bref. were tested for clonality through somatic compatibility tests. The following conclusions were reached: (i) P. contorta is highly susceptible to H. annosum and H. parviporum and both pathogens cause dieback of P. contorta; (ii) H. annosum from previous-rotation P. sylvestris stumps can effectively transfer to P. contorta; (iii) the pathogens may form constantly expanding territorial clones; (iv) basidiospores of both pathogens colonise stumps of P. contorta (primary infections); (v) H. parviporum clones expanded more slowly than clones of H. annosum; (vi) clonal spread proceeded more quickly from stumps with established secondary infections than from stumps with primary infections; (vii) H. annosum can persist in pine stumps for at least 26 years; and (viii) stump treatment should be considered to control Heterobasidion primary infections.