Nitrogen cycling following mountain pine beetle disturbance in lodgepole pine forests of Greater Yellowstone

Widespread bark beetle outbreaks are currently affecting multiple conifer forest types throughout western North America, yet many ecosystem-level consequences of this disturbance are poorly understood. We quantified the effect of mountain pine beetle (Dendroctonus ponderosae) outbreak on nitrogen (N) cycling through litter, soil, and vegetation in lodgepole pine (Pinus contorta var. latifolia) forests of the Greater Yellowstone Ecosystem (WY, USA) across a 0-30 year chronosequence of time-since-beetle disturbance. Recent (1-4 years) bark beetle disturbance increased total litter depth and N concentration in needle litter relative to undisturbed stands, and soils in recently disturbed stands were cooler with greater rates of net N mineralization and nitrification than undisturbed sites. Thirty years after beetle outbreak, needle litter N concentration remained elevated; however total litter N concentration, total litter mass, and soil N pools and fluxes were not different from undisturbed stands. Canopy N pool size declined 58% in recent outbreaks, and remained 48% lower than undisturbed in 30-year old outbreaks. Foliar N concentrations in unattacked lodgepole pine trees and an understory sedge were positively correlated with net N mineralization in soils across the chronosequence. Bark beetle disturbance altered N cycling through the litter, soil, and vegetation of lodgepole pine forests, but changes in soil N cycling were less severe than those observed following stand replacing fire. Several lines of evidence suggest the potential for N leaching is low following bark beetle disturbance in lodgepole pine.     

Response of advance lodgepole pine regeneration to overstory removal in eastern Idaho

Twenty-two stands of advance lodgepole pine released with overstory removal were sampled to determine height growth response. Tree and site characteristics correlated with release response were identified, and a mathematical model was developed to predict height growth in years 6 through 10 after release as a function of residual overstory basal area, height at release, percent rock cover, five-year cumulative growth prior to release, logging damage, stand elevation, and habitat type. Lodgepole pine responded to release with increased height growth in 97% of the trees sampled. Growth was similar to that of unsuppressed trees. Height growth is best when the entire overstory is removed and logging damage is avoided. Taller trees generally do not respond as well as shorter trees. However, trees growing fast before harvest continue to grow fast regardless of their height at release. Recommendations for selecting stands of advance lodgepole pine to release are provided.     

Growth response and sapwood hydraulic properties of young lodgepole pine following repeated fertilization

We examined how tree growth and hydraulic properties of branches and boles are influenced by periodic (about 6 years) and annual fertilization in two juvenile lodgepole pine (Pinus contorta Dougl. var. latifolia Engelm.) stands in the interior of British Columbia, Canada. Mean basal area (BA), diameter at breast height (DBH) and height increments and percent earlywood and sapwood hydraulic parameters of branches and boles were measured 7 or 8 years after the initial treatments at Sheridan Creek and Kenneth Creek. At Sheridan Creek, fertilization significantly increased BA and DBH increments, but had no effect on height increment. At Kenneth Creek, fertilization increased BA, but fertilized trees had significantly lower height increments than control trees. Sapwood permeability was greater in lower branches of repeatedly fertilized trees than in those of control trees. Sapwood permeabilities of the lower branches of trees in the control, periodic and annual treatments were 0.24 x 10(-12), 0.35 x 10(-12) and 0.45 x 10(-12) m2 at Kenneth Creek; and 0.41 x 10(-12), 0.54 x 10(-12) and 0.65 x 10(-12) m2 at Sheridan Creek, respectively. Annual fertilization tended to increase leaf specific conductivities and Huber values of the lower branches of trees at both study sites. We conclude that, in trees fertilized annually, the higher flow capacity of lower branches may reduce the availability of water to support annual growth of the leader and upper branches.     

Mountain pine beetle scarring of lodgepole pine in south-central Oregon

Three forest disturbance periods, 1973—present, 1922–1929, and 1827–1946 were determined by aging scars on stems of lodgepole pine trees. All of the scars from the 1970's and 1920's, and most of the scars from the 1820's–40's were determined to be caused by the mountain pine beetle. A few scars from the earliest period may be the result of fire. Diagnostic characteristics of beetle scars are: resemblance to fire scars, pitch tubes, beetle emergence holes, blue stain, beetle galleries, retained bark on the scar face, and an orange or red discoloration around healthy sapwood. Beetle attacks tend to have a northeastern aspect and extend approximately two-thirds around tree boles at breast height.By recognizing beetle scars it is possible to accurately age previous mountain pine beetle attacks. Many scars which had been thought to be of fire origin are actually caused by the mountain pine beetle.     

Removal of ethanol from lodgepole pine roots

The removal of ethanol from flooded tree roots was examined in three provenances of lodgepole pine (Pinus contorta) Dougl. Less than 0.2% of the ethanol generated by the roots escaped by the gaseous pathway provided by the lenticels. A large proportion, however, was transported from the roots in the transpiration stream. Gas chromatographic detection of ethanol emanating from the lenticels provided a sensitive indicator of oxygen deficits in the roots.     

Decay of fire-damaged lodgepole pine in south-central Oregon

About 150 lodgepole pines, Pinus contorta var. murrayana (Grev. and Balf.), growing in topoedaphic climax stands of south-central Oregon, were identified as having survived fires that occurred over a period from 1839 to 1982. All fires were natural wildfires, except for prescribed burns of 1976, 1980 and 1982. The trees were sampled for bole and root damage to investigate fungal colonization patterns, establishment of decay columns, and diameter growth.Fire-damaged roots were the principal entry point for decay fungi in fire damaged trees. White-rot fungal species and fungi imperfecti rapidly colonized fire-killed root tissues within 2 years after a fire. These fungi were later displaced by slower growing brown-rot Basidiomycetes that eventually extended into the boles of infected trees. Trees with butt rot, principally caused by Poria asiatica, grew significantly slower than uninfected trees.     

Tree regeneration and future stand development after bark beetle infestation and harvesting in Colorado lodgepole pine stands

In the southern Rocky Mountains, current mountain pine beetle (Dendroctonus ponderosae Hopkins) outbreaks and associated harvesting have set millions of hectares of lodgepole pine (Pinus contorta var. latifolia Engelm. ex Wats.) forest onto new stand development trajectories. Information about immediate, post-disturbance tree regeneration will provide insight on dynamics of future stand composition and structure. We compared tree regeneration in eight paired harvested and untreated lodgepole pine stands in the Fraser Experimental Forest that experienced more than 70% overstory mortality due to beetles. New seedlings colonized both harvested and untreated stands in the first years after the beetle outbreak. In harvested areas the density of new seedlings, predominantly lodgepole pine and aspen, was four times higher than in untreated stands. Annual height growth of pine and fir advance regeneration (e.g., trees established prior to the onset of the outbreak) has doubled following overstory mortality in untreated stands. Growth simulations based on our regeneration data suggest that stand basal area and stem density will return to pre-beetle levels in untreated and harvested stands within 80-105 years. Furthermore, lodgepole pine will remain the dominant species in harvested stands over the next century, but subalpine fir will become the most abundant species in untreated areas. Owing to terrain, economic and administrative limitations, active management will treat a small fraction (<15%) of the forests killed by pine beetle. Our findings suggest that the long-term consequences of the outbreak will be most dramatic in untreated forests where the shift in tree species composition will influence timber and water production, wildfire behavior, wildlife habitat and other forest attributes.     

Ponderosa pine snag densities following multiple fires in the Gila Wilderness,New Mexico

Fires create and consume snags (standing dead trees), an important structural and ecological component of ponderosa pine forests. The effects of repeated fires on snag densities in ponderosa pine forests of the southwestern USA have not been studied. Line intercept sampling was used to estimate snag densities in areas of the Gila Wilderness that had burned one to three times under Wildland Fire Use for Resource Benefit (WFU), a fire management policy implemented since 1974 aimed at restoring natural fire regimes. Twenty randomly located transects were measured in areas burned since 1946; six in once-burned areas, six in twice-burned areas and eight in thrice-burned areas. The mean density ± standard errors of large (>47.5 cm dbh) snags for areas that burned once, twice and thrice was 7.0 ± 2.7, 4.4 ± 1.1 and 4.1 ± 1.3 snags/ha, respectively. Differences in snag densities between once- and multiple-burned areas were significant (F-test; p < 0.05). There was no significant difference in density of large snags between twice- and thrice-burned areas. Proportions of type 1 snags (recently created) were higher in once- and twice-burned areas than in areas that burned three times, likely reflecting high tree mortality and snag recruitment resulting from an initial entry fire. Type 3 snags (charred by previous fire) were more abundant in areas that burned multiple times. The lack of differences in snag densities between areas that burned two and three times suggests that repeated fires leave many snags standing. The increasing proportion of type 3 snags with repeated fires supports this conclusion.     

Effects of compaction and water content on lodgepole pine seedling growth

Soil disturbance by heavy machinery usually causes a decrease in porosity and an increase in soil strength, which may limit access to nutrients and compromise seedling survival and growth. This study used a soil strength and a greenhouse experiment to assess the impact of compaction on lodgepole pine (Pinus contorta Dougl. Ex. Loud. var. latifolia Engelm.) seedling growth and the degree to which soil water influences the effects of compaction. A silt loam soil was collected from a forest landing in the central interior of British Columbia (BC) in the Sub-Boreal Spruce Biogeoclimatic zone. The silt loam soil was used in a soil strength experiment where soil with four water content levels (0.10, 0.18, 0.27, and 0.36 cm³ cm⁻³) was packed into 0.21 cm³ cores with three levels of compaction (74, 79, and 84% of maximum bulk density (MBD)). Soil strength was strongly affected by compaction and water content. In the greenhouse experiment, three water content levels (0.10–0.15, 0.20–0.30, and 0.30–0.35 cm³ cm⁻³) and three levels of compaction (67, 72, and 76% of MBD) were applied to soil in pots and 1-year old lodgepole pine seedlings were grown in the pots. Soil strength was highest (1275 kPa) for the high compaction and dry water content treatment in the greenhouse experiment. Though the soil strength for this treatment did not exceed 2500 kPa, the effect of compaction on growth was noticeable, with a decrease in diameter growth, total shoot mass, and new root mass as compaction increased at the dry water content. At dry water content and high compaction, foliar nutrient concentrations were greatest. Generally, water content had a greater impact on seedling growth than did compaction, at the levels of compaction used in this study. This study indicates that if there is a critical value for mechanical impedance of the conifer roots, it likely occurs below 2500 kPa. Our results are consistent with the explanation that soil strength incrementally affects root growth below 2500 kPa for this soil type. Expensive rehabilitation techniques may not be needed on lightly disturbed soils similar to that used in this study if soil water content is high enough throughout the conifer growing season to alleviate the effects of compaction on soil strength.     

Shoot dieback of lodgepole pine in Scotland caused by Ramichloridium pini

Research conducted on shoot dieback of lodgepole pine, Pinus contorta, during the early 1980's in west Scotland is summarised. The dieback symptoms and their stages of development are described. Ramichloridium pini de Hoog and Rahman was regularly isolated from bark and xylem tissue of dieased shoots. Artificial inoculations established that this fungus was the cause of the disease.     

Dynamics of mediterranean pine forests reforested after fires

Forest fires are frequent under a Mediterranean climate and have shaped the landscape of the region but are currently altered by human action and climate change.Fires have historically conditioned the presence of pine forests,depending on severity and forest regeneration.Regeneration of Mediterranean pine forests is not always successful,and a transition to shrublands or stands of resprouting species can occur,even after reforestation.This study analyses vegetation changes in two Mediterranean p...     

Influence of fire and mountain pine beetle on the dynamics of lodgepole pine stands in British Columbia,Canada

An outbreak of the mountain pine beetle (Dendroctonus ponderosae Hopkins; MPB), currently affecting over 10.1 million hectares of lodgepole pine forests (Pinus contorta Dougl.) in British Columbia, Canada, is the largest in recorded history. We examined the dynamics of even-aged lodgepole pine forests in southern British Columbia, which were undergoing this MPB outbreak. Using dendroecology and forest measurements we reconstructed the stand processes of stand initiation, stand disturbances, tree mortality, and regeneration, and explained the current stand structure and the potential MPB impacts in selected stands. Our results indicate that stand-replacing fires initiated even-aged seral lodgepole pine stands in this region. In the absence of fire in the 20th century, multiple MPB disturbances, which each resulted in partial canopy removal, modified the simple one-layer structure of the fire-origin stands by the initiation of post-MPB disturbance regeneration layers, transforming the stands into complex, multi-aged stands. Despite high overstory mortality due to the current MPB outbreak, regeneration layers, which are likely to survive the current outbreak, will provide important ecological legacies and will contribute to mid-term timber supply.     

Variation and heritability of resistance of lodgepole pine to Western gall rust

Families of pine seedlings from 40 open-pollinated trees (mother trees) from the interior of British Columbia (B. C.) were inoculated with two spore mixtures, one from the coast and the other from the interior of B. C. The relative susceptibilities of the 40 families to each spore mixture was assessed and narrow sense heritability with respect to disease resistance was estimated. The results are presented and discussed.     

Parameters affecting shoot production and rooting of cuttings from lodgepole pine hedges

By propagating lodgepole pine (Pinus contorta) cuttings in vivo, we obtained after 7 growing cycles (ca 3.5 years) in a greenhouse, sufficient number of cuttings from most families to establish clonal progeny tests. Twenty-one full-sib families with approximately 20 clones per family were studied for five years. Years when cuttings were set, families within latitude and clones within families differed significantly in rooting percentages, with the variance components 4.2%, 8.2% and 9.5%, respectively. One way to get a frequent and uniform rooting is to take cuttings from non-leading shoots since they have higher rooting percentage than leading shoots. Neither total length of the cuttings nor length of the primary needles were significantly correlated to rooting percentage. With appropriate management of the ortets and the cuttings during rooting, most clones could be included in a cutting propagation program.     

Structure and abundance of arachnid communities in Scots and lodgepole pine plantations

Between April and September 1994, six plots within Aberfoyle Forest in Scotland, three of Scots (Pinus sylvestris) and three of lodgepole pine (Pinus contorta), were surveyed for spiders and harvestmen, as these are predators of several forest invertebrate pests.A total of 64 species of spider were trapped in Scots and lodgepole pine; 56 in Scots pine and 32 in Lodgepole pine. The most abundant spiders, both in Scots and Lodgepole pine, were Linyphiidae.Scots pine plots were floristically more diverse and structurally more complex than lodgepole pine plots. More species of spiders and harvestmen (midsummer only) were found at ground level in Scots pine than lodgepole pine plots. More species and a greater numbers of spiders were found in the canopy of Scots pine than in that of lodgepole pine. However, no differences in the abundance of harvestmen were found between the canopies of Scots and lodgepole pine. Spider species diversity was greatest in Scots pine.     

Risk rating for mountain pine beetle infestation of lodgepole pine forests over large areas with ordinal regression modelling

The mountain pine beetle Dendroctonus ponderosae Hopkins is endemic to lodgepole pine, Pinus contorta var. latifolia Engelmann, forests in western Canada. However, the current beetle epidemic in this area highlights the challenges faced by forest managers tasked with prioritizing stands for mitigation activities such as salvage harvesting and direct control methods. In western Canada, the operational risk rating system for mountain pine beetle is based on biological knowledge gained from a rich legacy of stand-scale field studies. Due to the large spatial (millions of hectares affected) and temporal (over 10 years) extents of the current epidemic, new research into large-area mountain pine beetle processes has revealed further insights into the landscape-scale characteristics of beetle infested forests. In this paper, we evaluated the potential for this new knowledge to augment an established system for rating the short-term risk of tree mortality in a stand due to mountain pine beetle. New variables explored for utility in risk rating include direct shortwave radiation, site index, diameter at breast height, the temporal trends in local beetle populations, Biogeoclimatic Ecosystem Classification and beetle–host interaction variables. Proportional odds ordinal regression was used to develop a model for the Vanderhoof Forest District in west-central British Columbia. Prediction on independent data was assessed with the area under the receiver operator curve (AUC), indicating good discriminatory power (AUC = 0.84) for predicting levels of mountain pine beetle-caused pine mortality.     

Patterns of nitrogen mineralization in 23-year old pine forest following nitrogen fertilizing

Application of nitrogen fertilizer to 23-year old Pinus radiata D. Don at high rates induced considerable change in patterns of nitrogen mineralization in the soil. Increased net mineralization, nitrification, root Nitrate Reductase Activity, and foliar N levels were measured within 12 months of fertilizer application. Fertilizing did not, however, affect growth significantly over the 30 months of measurement. Significant differences in inorganic-N and patterns of mineralization between fertilized plots and controls were not maintained and all of the fertilized plots had levels of inorganic-N and patterns of mineralization approaching those of the control within 30 months. N added to these young aggrading forests of high productivity therefore appears to be quickly immobilized. Within normal forest rotations, it is apparent that the C/N ratio of the soil acts as a strong buffer to perturbation of mineralization patterns.     

Fertilization of lodgepole pine trees increased diameter growth but reduced root carbohydrate concentrations

Tree growth variables and more complex derived variables such as vigor index have all been linked to the ability of lodgepole pine (Pinus contorta var latifolia) trees to defend against insect herbivores, particularly mountain pine beetle (Dendroctonus ponderosae Hopkins, Coleoptera: Curculionidae). These variables are considered indirect measures of carbohydrate reserves. Trees with high vigor index values have high growth rates and are assumed to have high carbohydrate reserves. However, this critical assumption was untested for lodgepole pine. In this paper we evaluated the relationship between carbohydrate concentration and tree characteristics. We measured concentrations of root starch in 10 pure lodgepole pine stands of different ages in the Canadian Rockies which had been fertilized, thinned, fertilized and thinned or left untreated in a 2 × 2 factorial design. Both fertilization and thinning increased basal area increment while only fertilization increased lateral branch growth in the crown. Contrary to expectations, fertilization decreased root starch reserves although it increased basal area increment; thinning had no effect on root carbohydrates. Root starch reserves were positively related to basal area increment and vigor index in both fertilized and unfertilized plots, however, the best predictors of starch reserves were height-to-live-crown and cambial surface area below the live crown; starch reserves declined as both of these variables increased. These data suggest that large carbon sinks associated with long boles below the crown, and rapid growth of the crown and stem as a result of fertilization are detrimental to building starch reserves in the roots.     

Clones of Armillaria ostoyae and the pattern of infected juvenile lodgepole pine in Alberta,Canada

Small patches of dead trees were common among 7- to 15-year-old Pinns contorta var. latifolia in Alberta, Canada. The sizes of clones of Armillaria ostoyae, determined by pairing diploid isolates, was larger than these patches. Pairing diploid isolates on strips of Betula papyrifera provided a clear indication of clonal relationships.