Lodgepole Pine Research Articles

Lodgepole pine and interior spruce radial growth response to climate and topography in the Canadian Rocky Mountains, Alberta

Climate change may have spatially variable impacts on growth of trees in topographically diverse environments, making generalizing across broad spatial and temporal extents inappropriate. Therefore, topography must be considered when analyzing growth response to climate. We address these topo-climatic relationships in the Canadian Rocky Mountains, focusing on lodgepole pine (Pinus contorta Douglas ex Louden) and interior spruce (Picea glauca (Moench) Voss × Picea engelmannii hybrid Parry) growth response to climate, Palmer drought severity index (PDSI), aspect, and slope angle. Climate variables correlate with older lodgepole pine growth on south- and west-facing slopes, including previous August temperature, winter and spring precipitation, and previous late-summer and current spring PDSI, but younger lodgepole pine were generally less sensitive to climate. Climate variables correlate with interior spruce growth on all slope aspects, with winter temperature and PDSI important for young and old individuals. Numerous monthly growth–climate correlations are not temporally stable, with shifts over the past century, and response differs by slope aspect and angle. Both species are likely to be negatively affected by moisture stress in the future in some, but not all, topographic environments. Results suggest species-specific and site-specific spatiotemporally diverse climate–growth responses, indicating that climate change is likely to have spatially variable impacts on radial growth response in mountainous environments.

Soil carbon balance of afforested peatlands in the maritime temperate climatic zone

Drainage and conversion of natural peatlands into forestry increases soil CO2 emissions through decomposition of peat and modifies the quantity and quality of litter inputs and therefore the soil carbon balance. In organic soils, CO2 net emissions and removals, are reported using carbon emission factors (EF). The choice of specific default Tier 1 EF values from the IPCC 2013 Wetlands supplement depends on land use categories and climate zones. However, Tier 1 EF for afforested peatlands in the temperate maritime climate zone are based on data from eight sites, mainly located in the hemiboreal zone, and the uncertainty associated with these default values is a concern. In addition, moving from Tier 1 to higher-Tier carbon reporting values is highly desirable when large areas are affected by land use changes. In this study, we estimated site-specific soil carbon balance for the development of Tier 2 soil CO2 -C EFs for afforested peatlands. Soil heterotrophic respiration and aboveground litterfall were measured during two years at eight afforested peatland sites in Ireland. In addition, fine roots turnover rate and site-specific fine root biomass were used to quantify belowground litter inputs. Our findings found that draining of peatlands, and planting them with either Sitka spruce or lodgepole pine, resulted in soils being net carbon sources. The soil carbon balance at multi-year sites varied between 63 ± 92 and 309 ± 67 g C m-2 year-1 . Mean CO2 -C EF for afforested peatlands was 1.68 ± 0.33 tCO2 -Cha-1 year-1 . The improved CO2 -C EFs presented here for afforested peatlands are proposed as a basis to update national CO2 -C emissions from this land-use class in Ireland. Furthermore, new data from these sites will significantly contribute to the development of more reliable IPCC default Tier 1 CO2 -C EFs for afforested peatlands in the maritime temperate climate zone.

Bayesian Predictions of Bark Beetle Attack and Mortality of Three Conifer Species During Epidemic and Endemic Population Stages

Bark beetles naturally inhabit forests and can cause large-scale tree mortality when they reach epidemic population numbers. A recent epidemic (1990s–2010s), primarily driven by mountain pine beetles (Dendroctonus ponderosae), was a leading mortality agent in western United States forests. Predictive models of beetle populations and their impact on forests largely depend on host related parameters, such as stand age, basal area, and density. We hypothesized that bark beetle attack patterns are also dependent on inferred beetle population densities: large epidemic populations of beetles will preferentially attack large-diameter trees, and successfully kill them with overwhelming numbers. Conversely, small endemic beetle populations will opportunistically attack stressed and small trees. We tested this hypothesis using 12 years of repeated field observations of three dominant forest species (lodgepole pine Pinus contorta, Engelmann spruce Picea engelmannii, and subalpine fir Abies lasiocarpa) in subalpine forests of southeastern Wyoming paired with a Bayesian modeling approach. The models provide probabilistic predictions of beetle attack patterns that are free of assumptions required by frequentist models that are often violated in these data sets. Furthermore, we assessed seedling/sapling regeneration in response to overstory mortality and hypothesized that higher seedling/sapling establishment occurs in areas with highest overstory mortality because resources are freed from competing trees. Our results indicate that large-diameter trees were more likely to be attacked and killed by bark beetles than small-diameter trees during epidemic years for all species, but there was no shift toward preferentially attacking small-diameter trees in post-epidemic years. However, probabilities of bark beetle attack and mortality increased for small diameter lodgepole pine and Engelmann spruce trees in post-epidemic years compared to epidemic years. We also show an increase in overall understory growth (graminoids, forbs, and shrubs) and seedling/sapling establishment in response to beetle-caused overstory mortality, especially in lodgepole pine dominated stands. Our observations provide evidence of the trajectories of attack and mortality as well as early forest regrowth of three common tree species during the transition from epidemic to post-epidemic stages of bark beetle populations in the field.

Susceptibility of sugar pine, Shasta red fir and Sierra lodgepole pine to mountain hemlock dwarf mistletoe (Arceuthobiumtsugense subsp. mertensianae, Viscaceae) in south central Oregon

AbstractIn southern Oregon and northern California, mountain hemlock dwarf mistletoe (MHDM; Arceuthobium tsugense subsp. mertensianae) commonly parasitizes mountain hemlock (Tsuga mertensiana) in multiple mountain ranges such as the Cascades, Sierra Nevada and Klamath Mountains. However, limited information is available on the susceptibility of sugar pine (Pinus lambertiana), Shasta red fir (Abies magnifica var. shastensis) and lodgepole pine (Pinus contorta) to MHDM. From 2015–2019, I found MHDM on sugar pine at multiple sites in northwestern Klamath County, Oregon. To evaluate the susceptibilities of sugar pine, Shasta red fir and Sierra lodgepole pine to MHDM infection, 41 temporary, fixed‐radius plots (area 182.4 m2) were established at five sites in 2018 and 2019 around an overstory tree (western white, sugar and whitebark pines along with one mountain hemlock) severely infected with MHDM in each plot centre. Live trees ≥2.6 cm in diameter at breast height (1.37 m; dbh) were assigned a dwarf mistletoe severity rating (6‐class system) and incidence (% infection) of MHDM on live trees at the tree level were used to estimate host susceptibilities. To confirm the identity of the dwarf mistletoe infecting live trees in each plot, dwarf mistletoe specimens were collected from the severely infected tree at the centre of each plot and from all tree species bearing dwarf mistletoe in each plot. Morphological characters were used to identify the dwarf mistletoe plants on each tree species and included plant heights, length of the third internode and plant colour. Dwarf mistletoe plants were collected from each affected tree species, and morphological characteristics of aerial shoots of dwarf mistletoes were examined in the laboratory. Morphological measurements confirmed the dwarf mistletoe infesting each plot was MHDM. Mountain hemlock dwarf mistletoe was found on sugar pine in all 41 plots. At the tree level after pooling larger sugar pine trees from all plots (total of 164 live trees ≥2.6 cm dbh), incidence was 76%. Morphological characteristics of MHDM aerial shoots, such as colour, shoot height and third internode length, were consistent with previous descriptions of MHDM plants. These data indicate that sugar pine should be classified as a secondary host of MHDM in the Oregon East Cascades, and not as a rare host as previously reported. No dwarf mistletoe plants were found on 98 living Shasta red fir trees ≥2.6 cm dbh in 26 plots. Shasta red fir should be classified as immune to MHDM. No MHDM was found on 573 living lodgepole pines of this size in 41 plots, and the tree species should be classified as immune to MHDM. Western white (Pinus monticola) and whitebark (Pinus albicaulis) pines were severely parasitized by MHDM with 92% and 100% incidence, respectively. However, because the sample sizes for these white pines were relatively small, additional work is needed to evaluate the susceptibility of these pines to MHDM.

Corn Grain and Stover Nutrient Uptake Responses from Sandy Soil Treated with Designer Biochars and Compost

Biochars are used for soil fertility improvement because they may contain certain elements that plants use as nutrients. However, few studies have demonstrated enhanced crop nutrient uptake. Our study examined nutrient uptake responses of corn (Zea Mays L.) grain and stover over 4 years (Y) after a Goldsboro sandy loam (fine-loamy, siliceous, sub-active, thermic Aquic Paleudults) received different designer biochars and a compost. The designer biochars were produced from lodgepole pine (Pinus contorta) chip (PC), poultry litter (PL), blends with switchgrass (SG; Panicum virgatum), and a SG compost alone. Topsoil treated with 100% PL biochar and blended PC:PL biochar had significantly greater Mehlich 1 (M1) extractable P, K and Na contents compared to the control or other treatments. No significant differences were detected in annual grain nutrient concentrations. In the first corn stover harvest (Y1), significantly greater concentrations of P and K were taken up after treatment with 100% PL biochar, with PC:PL blend and with SG when compared to control. By the fourth corn stover harvest (Y4), nutrient uptake between treatments was not significantly different. Biochar impact on corn stover P, K and Na concentrations was time dependent, suggesting that repeated biochar applications may be needed.

Rates, patterns, and drivers of tree reinvasion 15 years after large‐scale meadow‐restoration treatments

Tree encroachment threatens the biodiversity and ecological functioning of grasslands worldwide. Reversing effects of encroachment requires not only removing trees, but limiting subsequent invasions, which can stall grassland reassembly. We quantified rates and patterns of conifer reinvasion of mountain grasslands (meadows) in western Oregon, 7 and 15 years after experimental tree removal with or without burning. We assessed frequency (percentage of 100 m2 subplots) and density of reinvading trees in six 1‐ha plots as a function of burn treatment, abundance in the adjacent forest (a proxy for seed availability), and distance to or orientation of forest edges. After 15 years, frequency and density of colonists varied widely among plots (24–57% of subplots and 60–250 trees/ha, respectively), but burning had no effect on either measure. Abies grandis, which dominated adjacent forests, was the principal invader. Colonization by less‐common Pinus contorta and A. procera was correlated with species' density and basal area in adjacent forests, suggesting that establishment was seed limited. Annualized rates of invasion were similar between early and late sampling intervals (9–10/trees/ha). Establishment was generally greater along the forested edges of plots (0–10 m) than in the core (10–50 m). Prominence of shade‐tolerant A. grandis along northerly‐facing edges suggests strong microclimatic controls on establishment. In sum, tree removal, with or without prescribed fire, is successful in reversing effects of conifer encroachment. Despite abundant seed rain, reinvasion is slow and spatially restricted. Periodic hand removal of colonizing trees, while they are small, can be done efficiently and economically.

Soil compaction and organic matter removal effects on soil properties and tree growth in the Interior Douglas-fir zone of southern British Columbia

Silviculture systems and associated forest harvesting practices that affect the level of soil compaction and organic matter retained on a site will cause changes to soil properties and processes that may affect the growth of planted trees. We present results for soil conditions and tree growth after 15 years for 6 Long-term Soil Productivity Study (LTSP) installations in 2 distinct regions of the Interior Douglas-fir (IDF) biogeoclimatic zone in southern British Columbia (BC). In a 3 × 3 factorial design, three levels of compaction and organic matter removal were replicated in two distinct IDF regions that varied in parent material, but with similar climates. Each treatment was then split and planted with two different species, Douglas-fir and lodgepole pine. In each region, three unique sites (block), all within a radius of 50 km from one other, were installed with the same combination of treatment and species for a total of six replicates, three in each of two regions. Across IDF regions, tree heights on calcareous soils in the Rocky Mountains of south-eastern BC were 36% lower for Douglas-fir and 23% lower for lodgepole pine than on more acidic soils on the Thompson Plateau. There were varying differences across treatments in soil and foliar nutrient concentrations between IDF regions, primarily from regional parent material differences. However, nutrient deficiencies were apparent in both regions and may have limited growth, especially on the more calcareous sites. Increases to soil bulk density (27%) 1 year after compaction did not impact tree survival, height at Year-15, or chemical properties in foliar tissue or soil. Fewer growing season frost events and increased soil water availability during the seedling establishment period was a likely reason for a 25% increase in Douglas-fir survival at Year-15 with the whole tree + forest floor removal (OM3) treatment. However, whole tree + forest floor removal (OM3) also produced a 13–19% reduction in soil N, S, K, and mineralizable N when compared to both stem-only (OM1) and whole tree (OM2) harvesting treatments that kept the forest floor intact. Effects on forest productivity from soil compaction after 15 years appear to be overshadowed by the more prominent effect of organic matter removal treatments during stand establishment and early growth. Future work at these installations focusing on nutrient budget, forest health, and ecosystem carbon stock differences across treatments will provide valuable insight into silviculture and harvesting practices for different species and regions in the dry forests of southern BC.

Growth and defense characteristics of whitebark pine (Pinus albicaulis) and lodgepole pine (Pinus contorta var latifolia) in a high-elevation, disturbance-prone mixed-conifer forest in northwestern Montana, USA

Recent, widespread tree mortality in the western U.S. resulting from changes in climate, pathogens, insect activity, and forest management practices has led to concerns for many ecologically and culturally important species. Within conifers, resin-based defenses have long been recognized as a primary defense mechanism against a variety of insects and pathogens. Oleoresin produced by trees contain complex mixtures of terpenoids that have numerous insecticidal and fungicidal properties. Research has also identified links between resin duct characteristics and increased probability of survival during bark beetle outbreaks. Whitebark pine (Pinus albicaulis) is a culturally significant high elevation species that provides numerous ecological services within subalpine and alpine ecosystems. Whitebark pine has co-evolved with a suite of biotic and abiotic disturbances. Individual trees allocate resources towards growth and resin-based defenses, making it a good candidate species to evaluate growth and defense relationships and tradeoffs. In this study we compared constitutive resin chemistry, tree growth and resin duct anatomy between similarly aged whitebark and lodgepole pine (P. contorta var latifolia) growing in proximity within a disturbance-prone, mixed-conifer forest in northwestern Montana. These two host species have varying degrees of historical exposure to mountain pine beetle. Our research yields four important findings. First, we did not find evidence of a tradeoff between tree growth and tree defenses (resin duct morphology and resin chemistry). This suggests that trees growing under favorable field conditions can experience high growth rates and still allocate ample resources towards defense. Second, we found that resin ducts and constitutive mono- and sesqui- terpenes were not correlated in lodgepole pine while duct production and area were positively related to constitutive monoterpenes, and duct size and area were positively related to constitutive sesquiterpenes, in whitebark pine. The lack of distinct, consistent relationships between these defensive features suggests that both whitebark and lodgepole pine trees present beetles with numerous, complex combinations of resin-based defenses. Third, based on constitutive terpene profiles, bark beetles are more likely to enter lodgepole pine but more likely to successfully elicit mass attacks in whitebark pine, which agrees with beetle attack and success patterns observed in the field. Fourth, overstory competition, particularly by Engelmann spruce (Picea engelmannii), can influence tree defenses, specifically by reducing constitutive terpene concentrations in lodgepole and whitebark pine. Competitive tree interactions could lead to altered bark beetle-conifer interactions as host and nonhost species migrate in response to changing climate. Our results suggest that strategies designed to support whitebark pine populations can benefit from better understanding interactions among growth, competition and physical and chemical defenses in response to multiple disturbance.

Evaluating topographic variation as a guide to Cassia crossbill refugia

The Cassia crossbill (Loxia sinesciuris) feeds exclusively on seeds held in serotinous cones of lodgepole pine (Pinus contorta latifolia). Its population, confined to 67 km2 of pine in southern Idaho, declined by ~ 80% between 2003 and 2011 apparently because an increase in the occurrence of hot summer days (≥32 °C) caused the serotinous cones to open and shed their seeds. We hypothesized that the propensity for serotinous cones to open varies across the landscape depending on the maximum temperatures cones experienced. We tested a suite of abiotic variables for predicting the occurrence of open serotinous cones among 1547 trees at 170 sites. The best-supported model indicated that trees with open cones were less frequent on northeastern aspects and as slope increased. Fewer open cones were expected on northeast aspects because they receive less direct solar radiation. However, the decrease in trees with open cones as slope increased on southwest aspects was unexpected. Increased convection on slopes could reduce maximum cone temperatures and account for the decrease of open cones on steeper slopes, regardless of aspects. Although cones are less likely to open on steeper slopes, slopes with southerly aspects are likely the least favorable for pine recruitment and survival. The most favorable sites for both lodgepole pine and the persistence of closed cones are slopes on northeast aspects. Such locations are often dominated by subalpine fir (Abies bifolia). Reforestation of fir-dominated areas with lodgepole pine, therefore, provides a means to expand microclimatic refugia for Cassia crossbills.

The characteristics of insoluble softwood substrates affect fungal morphology, secretome composition, and hydrolytic efficiency of enzymes produced by Trichoderma reesei

BackgroundOn-site enzyme production using Trichoderma reesei can improve yields and lower the overall cost of lignocellulose saccharification by exploiting the fungal gene regulatory mechanism that enables it to continuously adapt enzyme secretion to the substrate used for cultivation. To harness this, the interrelation between substrate characteristics and fungal response must be understood. However, fungal morphology or gene expression studies often lack structural and chemical substrate characterization. Here, T. reesei QM6a was cultivated on three softwood substrates: northern bleached softwood Kraft pulp (NBSK) and lodgepole pine pretreated either by dilute-acid-catalyzed steam pretreatment (LP-STEX) or mild alkaline oxidation (LP-ALKOX). With different pretreatments of similar starting materials, we presented the fungus with systematically modified substrates. This allowed the elucidation of substrate-induced changes in the fungal response and the testing of the secreted enzymes’ hydrolytic strength towards the same substrates.ResultsEnzyme activity time courses correlated with hemicellulose content and cellulose accessibility. Specifically, increased amounts of side-chain-cleaving hemicellulolytic enzymes in the protein produced on the complex substrates (LP-STEX; LP-ALKOX) was observed by secretome analysis. Confocal laser scanning micrographs showed that fungal micromorphology responded to changes in cellulose accessibility and initial culture viscosity. The latter was caused by surface charge and fiber dimensions, and likely restricted mass transfer, resulting in morphologies of fungi in stress. Supplementing a basic cellulolytic enzyme mixture with concentrated T. reesei supernatant improved saccharification efficiencies of the three substrates, where cellulose, xylan, and mannan conversion was increased by up to 27, 45, and 2800%, respectively. The improvement was most pronounced for proteins produced on LP-STEX and LP-ALKOX on those same substrates, and in the best case, efficiencies reached those of a state-of-the-art commercial enzyme preparation.ConclusionCultivation of T. reesei on LP-STEX and LP-ALKOX produced a protein mixture that increased the hydrolytic strength of a basic cellulase mixture to state-of-the-art performance on softwood substrates. This suggests that the fungal adaptation mechanism can be exploited to achieve enhanced performance in enzymatic hydrolysis without a priori knowledge of specific substrate requirements.

A Methodology for Experimental Quantification of Firebrand Generation from WUI Fuels

Over the past few years, numerous large-scale disasters have occurred due to wildfires at the wildland-urban interface (WUI). In these fires, spread via the transport of firebrands (burning embers) plays a significant role. Several models have been developed to describe the transport of firebrands but few, if any, are available which can provide a quantitative means to generate firebrands at the source of a fire. In this regard, a new methodology is proposed here that uses a wind tunnel to experimentally quantify the generation of firebrands from WUI fuels under different ambient conditions. The setup allows for the collection of all generated solid firebrands and major downstream gaseous species concentrations. Unique firebrand yield correlations can then be generated for each tested fuel, while also accounting for the heat-release rate, providing unique validation targets for numerical simulations. Generation of firebrands from branches of two conifers at a fixed wind speed of 4 m/s are presented to demonstrate the capabilities of this new methodology. A carbon mass balance was utilized to analyze preliminary results and understand how much of the fuel mass transitions to firebrands vs. gases. These results provide a description of the mass burning process and ultimately tie firebrand production to a time-dependent heat-release rate for initialization of firebrand transport in numerical simulations. An average firebrand yield ranging from 3–4% of initial dry mass is ultimately presented for lodgepole pine and Douglas fir. Future work is required with larger fuel sizes pertaining to real wildfire scenarios; however, the presented methodology can provide valuable data needed to initialize numerical simulations of firebrand transport, necessary for reconstruction of WUI fires and to aid in the development of mitigation strategies for the prevention of future disasters.

Microbial response to designer biochar and compost treatments for mining impacted soils.

The Oronogo-Duenweg mining belt is a designated United States Environmental Protection Agency Superfund site due to lead-contaminated soil and groundwater by former mining and smelting operations. Sites that have undergone remediation - in which the O, A, and B horizons have been removed alongside the lead contamination - have an exposed C horizon and are incalcitrant to revegetation efforts. Soils also continue to contain quantifiable Cd and Zn concentrations. In order to improve soil conditions and encourage successful site revegetation, our study employed three biochars, sourced from different feedstocks (poultry litter, beef lot manure, and lodge pole pine), at two rates of application (2.5%, and 5%), coupled with compost (0%, 2.5% and 5% application rates). Two plant species - switchgrass (Panicum virgatum) and buffalograss (Bouteloua dactyloides) - were grown in the amended soils. Amendment of soils with poultry litter biochar applied at 5% resulted in the greatest reduction of soil bioavailable Cd and Zn. Above ground biomass yields were greatest with beef lot manure biochar applied at 2.5% with 5% compost, or with 5% biochar at 2.5% and 5% compost rates. Maximal microbial biomass was achieved with 5% poultry litter biochar and 5% compost, and microbial communities in soils amended with poultry litter biochar distinctly clustered away from all other soil treatments. Additionally, poultry litter biochar amended soils had the highest enzyme activity rates for β-glucosidase, N-acetyl-β-D-glucosaminidase, and esterase. These results suggest that soil reclamation using biochar and compost can improve mine-impacted soil biogeophysical characteristics, and potentially improve future remediation efforts.

Anion Exchange membrane with High hydroxide ion conductivity and robust tensile strength fabricated from quaternary ammonia functionalized Pinus contorta, Dougl. Chip

The Pinus contorta, Dougl. chip was ammonianized by 3-chloro-2-hydroxypropyltrimethyl ammonium chloride. Subsequently, the functionalized chip was thermally compressed under 30 MPa at 100 °C for 72 h to form the quaternary ammonia functionalized membrane (QAFW). The results revealed that the hydroxide ion conductivity and ion exchange capacity (IEC) of the as-fabricated membrane were up to 15.2 mS cm−1 and 1.86 mmolg−1 at 25 °C under 100 % relative humidity respectively, which is higher than most of the quaternary ammonia-functionalized aromatic polymer membranes. The ratio of hydroxide ion conductivity to methanol permeability of the optimized QAFW membrane is up to 3.86 × 103 S s cm−3. The decomposition temperature of QAFW membrane is ∼252 °C. The tensile strength of the wet QAFW membrane is as high as 60.02 MPa, which is significantly higher than that of the aromatic polymer membranes. Conclusively, a novel method herein characterized with environmentally friendly and economically attractive for the anion exchange membranes fabrication was presented.

Acceleration of Forest Structural Development for Large Trees and Mammals: Restoration in Decades or Centuries?

There is a demand for more progressive restoration directives to regenerate forest ecosystems impacted by harvesting, wildfire, insect outbreaks, and mineral resource extraction. Forest restoration may take many decades and even centuries without active silvicultural intervention to grow large trees that provide suitable habitat for various wildlife species. We tested the hypotheses (H) that, compared with unmanaged (unthinned and old-growth) stands, large-scale precommercial thinning (heavy thinning to <500 stems/ha) of young lodgepole pine (Pinus contorta var. latifolia), at 20–25 years post-treatment, would enhance: (H1) the architecture of large overstory trees (e.g., diameter, height, and crown dimensions); (H2) mean (i) total abundance and species diversity of forest-floor small mammals, (ii) abundance of tree squirrels; and (H3) relative habitat use by mule deer (Odocoileus hemionus). There were three levels of thinning with mean densities of crop trees/ha: 353 (low), 712 (medium) and 1288 (high), an unthinned, and old-growth stand replicated at three areas in south-central British Columbia, Canada. Mammal abundance and habitat use were measured during the period 2013 to 2015. Mean diameter of crop trees was significantly different among stands with the low-density, medium-density, and old-growth stands having diameters larger than the high-density and unthinned stands. Mean height of crop trees was highest in the old-growth stands. Mean crown volume of crop trees was significantly different among stands with the low-density stands 2.1 to 5.8 times higher than the high-density, unthinned, and old-growth stands, and hence partial support for H1. Mean total abundance of forest-floor small mammals was significantly different among stands with the low-density and old-growth stands 1.9 to 2.4 times higher than the other three treatment stands. Mean abundances per stand of the red squirrel (Tamiasciurus hudsonicus) (range of 4.8 to 12.0) and the northern flying squirrel (Glaucomys sabrinus) (range of 3.2 to 4.3) were similar among stands. Mean relative habitat use by mule deer was similar among stands, but variable with counts of pellet-groups/ha in the thinned stands were 3.8 to 4.2 and 2.1 to 2.3 times higher than the unthinned and old-growth stands, respectively. Thus, mean total abundance of forest-floor small mammals of H2 was supported, but species diversity and abundance of tree squirrels was not. Enhanced relative habitat use by mule deer (H3) was not supported. To our knowledge, this is the first concurrent measurement of several mammal species in heavily thinned, unthinned, and old-growth forest across three replicate study areas at 20–25 years post-treatment. Although not all mammal responses were significant, there was a strong indication that restored forests via heavy thinning (<500 trees/ha) produced large overstory trees (at least for diameter and crown dimensions) in stands 33 to 42 years old. Comparable old-growth stands, albeit with crop trees of greater height and merchantable volume, ranged from 120 to 167 years of age. Restored forests with large trees capable of supporting at least these mammal species may be achieved in decades rather than centuries.

Are soil changes responsible for persistent slash pile burn scars in lodgepole pine forests?

Pile burning is the most common method of logging residue disposal in Rocky Mountain forests. Though the high temperatures reached during burning affect numerous soil properties in the short term, the longer-term effects of the practice are less clear. We previously identified a 50-year time series of burn scars created after clear cut harvesting in lodgepole pine stands where we reported sparse tree colonization across the entire chronosequence. Here we analyzed soil nutrients and chemistry and conducted in situ and greenhouse seedling bioassays to determine whether edaphic factors or poor seedling performance explain the pattern. Pile burning had a lasting effect on soil pH, but nutrient availability was 2–3 times higher in burn scars compared to unburned forest soils for many constituents and planted pine seedlings had good survival and growth. However, seedling growth was slightly less in burn scars compared to unburned soils indicating suboptimal soil pH or other belowground factors may contribute to sparse tree colonization of the openings. For example, seedling survival and ectomycorrhizal fungi colonization were both lowest in the most recently created scars where soils were alkaline and improved with time as pH declined, suggesting gradual amelioration of post-fire growing conditions. Survival in burn scars was comparable for unprotected trees and those in protective mesh tubes, indicating that herbivory was not a significant impediment to seedling establishment. However, a preliminary study suggests that seed predation may have contributed to the low tree colonization into the openings. Though large burn pile scars may require soil rehabilitation, and soil changes may have a lasting effect on understory plant composition, we found that they were not a significant barrier to tree establishment in these moderate-size burn scars.

Trade-offs across densities and mixture proportions in lodgepole pine-hybrid spruce plantations

Monocultures tend to yield higher total stand volumes and are simple to manage. Yet, mixed species stands may result in similar stand volumes while providing benefits such as mitigating damage from insects and disease. To understand the effects of stand density and species mixture and their interactions on stand yield, tree size and morphology, and damage in monocultures and mixtures, we analyzed a 25-year-old experiment in interior British Columbia, Canada. The lodgepole pine (Pl)-interior hybrid spruce (Sx) experiment included three densities—1000, 1500, and 2000 stems per hectare (SPH)—and five species mixtures—1:0, 3:1, 1:1, 1:3, and 0:1 Pl:Sx. Results 25 years after stand establishment showed that stand volume was significantly larger with an increasing proportion of Pl across all stand densities. Pl had 10% larger diameters in the 1000 SPH than in the 2000 SPH and when mixed with Sx (1:1). Pl had larger crowns in mixtures regardless of density. Mixture proportion did not affect gall rust incidence or stem form in Pl, but reduced attack in Sx by spruce weevil. Our findings suggest that mixing Pl-Sx and high planting density decrease weevil attacks in Sx, which reduce loss in timber quality. Yet, Pl quality may decrease when mixed with Sx, due to larger Pl crowns. These results may be used to improve the implementation of management strategies that decrease trade-offs between yields, desired market tree sizes, and timber loss from pest and pathogens, while making the stands more resilient to further climate change impact.

Higher temperatures increase growth rates of Rocky Mountain montane tree seedlings

AbstractRecent observational studies report weak or flat temperature − growth relationships for many tree species in temperate forests. In contrast, distribution limits of trees are strongly shaped by temperature, and studies show marked short‐term temperature effects on leaf‐level ecophysiology. To better determine the effects of warming on trees, we planted one‐year‐old seedlings of one lower montane (ponderosa pine), two upper montane (quaking aspen and lodgepole pine), and one subalpine tree species (subalpine fir) in in situ experimental gardens on an elevation gradient in the Rocky Mountains (USA) which span a 6°C range in temperature but have approximately uniform precipitation. Seedlings were lightly watered the first three growing seasons to facilitate establishment, and growth and survivorship were followed for four years. We expected a trade‐off between growth and survivorship, as seedlings in high temperatures grow faster (e.g., with a longer growing season), but have higher mortality from heat stress. Compared to the coldest site, aspen (+256% wider, +337% taller), ponderosa pine (+234% wider, 270% taller), and lodgepole pine (+235% wider, 283% taller) all had strikingly higher cumulative diameter and height growth in the warmest site by the end of the study. Linear models of cumulative and annual growth in the montane species showed strong, positive relationships with growing‐season temperature, but no significant relationships with growing‐season precipitation. In contrast, growth of subalpine fir did not vary significantly with temperature, but increased slightly with higher growing‐season precipitation. Accelerated growth did not come at the expense of survivorship in the montane species: cumulative four‐year survivorship of the montane species remained robust (71.4–94.4%) in high temperatures, but caused complete mortality of subalpine fir. As long as precipitation remains adequate, these results indicate that warming is likely to strongly increase growth in seedlings of montane species with only modest decreases in survivorship despite higher evapotranspiration, especially in cooler and wetter portions of their current distributions where hydric stress is low. In contrast, warming may negatively affect seedling growth and survival in hotter and drier areas of the Rockies, and warming of +3–6°C may endanger the persistence of subalpine fir over much of its current distribution.

Tree mortality in western U.S. forests forecasted using forest inventory and Random Forest classification

AbstractClimate change is projected to significantly affect the vulnerability of forests across the western United States to wildfires, insects, disease, and droughts. Here, we provide recent mortality estimates for large trees for 53 species across 48 ecological sections using an analysis of 23,215 Forest Inventory plots and a Random Forest classification model. Models were also used to predict mortality in future FIA inventories under the RCP 4.5 emissions scenario. Model performance indicated species identity as the most important predictor of mortality under both current and future scenarios, with contributions from climate and soil variables. Our results show relatively high levels of recent mortality in the Middle and Southern Rocky Mountains driven by high mortality in Populus tremuloides, Pinus contorta, Pinus albicaulis, and Abies lasiocarpa. Low levels of mortality were observed in several species, with <1% annual mortality observed throughout all other sections. Future mortality was predicted to increase significantly for most species and correlated well with recent mortality at the species level, but not at the plot level. These results suggest that future attempts to model or predict widespread forest mortality would benefit from more research on vulnerable species and that significant mortality events in some species may not be important for dynamics across all systems.

Primary and Secondary Metabolite Profiles of Lodgepole Pine Trees Change with Elevation, but Not with Latitude.

Climate change has a large influence on plant functional and phenotypic traits including plant primary and secondary metabolites. One well-established approach to investigating the variation in plant metabolites involves studying plant populations along elevation and latitude gradients. We considered how two space-for-time climate change gradients (elevation and latitude) influence carbohydrate reserves (soluble sugars, starches) and secondary metabolites (monoterpenes, diterpene resin acids) of lodgepole pine trees in western Canada. We were particularly interested in the relationship of terpenes and carbohydrates with a wide range of tree, site, and climatic factors. We found that only elevation had a strong influence on the expression of both terpenes and carbohydrates of trees. Specifically, as elevation increased, concentrations of monoterpenes and diterpenes generally increased and soluble sugars (glucose, sucrose, total sugars) decreased. In contrast, latitude had no impact on either of terpenes or carbohydrates. Furthermore, we found a positive relationship between concentrations of starch and total terpenes and diterpenes in the elevation study; whereas neither starches nor sugars were correlated to terpenes in the latitude study. Similarly, both terpenes and carbohydrates had a much greater number of significant correlations to site characteristics such as slope, basal area index, and sand basal area, in the elevational than in the latitude study. Overall, these results support the conclusion that both biotic and abiotic factors likely drive the patterns of primary and secondary metabolite profiles of lodgepole pine along geographical gradients. Also, presence of a positive relationship between terpenes and starches suggests an interaction between primary ad secondary metabolites of lodgepole pine trees.

Legacy of bark beetles (Dendroctonus spp.) on soil carbon and nitrogen cycling seven years after forest infestation

Between years 2007 and 2013, an outbreak of pine- and spruce-bark beetles (Dendroctonus spp.) has decimated coniferous forests of the northwestern United States. Beetles introduced blue stain fungus to the tree xylem that caused blockage and discontinuation of plant water conductance. One of the immediate disease symptoms was premature and immediate drop-off of all needles rich in nitrogen (N). Soil surface-accumulated needle litter in combination with elevated soil water levels, have led to accelerated N and carbon (C) mineralization. Understanding of the soil C and N cycling during forest transition from “standing dead trees” to “snagfall and forest regrowth” is still unclear. Specifically, the potential lasting legacy of beetle-caused surface-deposited tree litter is of importance in order to assess long-term forest health. The main objective of this study was to assess whether soil C and N mineralization continues to be affected by litter chemistry in this transitional period of post beetle disturbance ecosystem recovery. Mineral soil (0–10 cm) and corresponding surface tree litter were obtained from beneath dead and live tree clusters in three forest types in Medicine Bow-Routt National Forest in Wyoming, USA. All forests were located in the same watershed, grew on similar soils, experienced up to 35% of forest mortality and consisted of: Engelmann spruce (Picea engelmannii), subalpine fir (Abies lasiocarpa) mixed with lodgepole pine (Pinus contorta), and lodgepole pine alone. Results suggest that tree litter uniformly affected mineral soils across all forest types regardless of beetle infestation. In laboratory incubations, the presence of litter doubled mineral soil heterotrophic respiration. While litter had the most uniform effect on soils, beetle infestation and forest type resulted in a number of differences. Soils beneath dead tree clusters continued to have elevated inorganic N concentrations when normalized by total N, which was twice as high compared to soils beneath live tree clusters across all forest types. The lowest NH4+ concentrations were observed in soils beneath lodgepole pine trees, which was accompanied by the highest C to N ratio of microbially available substrates. In conclusion, the legacy of forest die-offs caused by bark beetle infestation was still observable seven years later during the early transitional period. This confirms ecosystem modeling simulations that, while entering decadal stage of forest recovery, forest structure may differ depending on the forest type, and that the forest types that contain lodgepole pines may encounter an earlier onset of potential N limitation.