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.     

Susceptibility of ponderosa pine, Pinus ponderosa (Dougl. ex Laws.), to mountain pine beetle, Dendroctonus ponderosae Hopkins,attack in uneven-aged stands in the Black Hills of South Dakota and Wyoming USA

Mountain pine beetle, Dendroctonus ponderosae Hopkins can cause extensive tree mortality in ponderosa pine, Pinus ponderosa Dougl. ex Laws., forests in the Black Hills of South Dakota and Wyoming. Most studies that have examined stand susceptibility to mountain pine beetle have been conducted in even-aged stands. Land managers increasingly practice uneven-aged management. We established 84 clusters of four plots, one where bark beetle-caused mortality was present and three uninfested plots. For all plot trees we recorded species, tree diameter, and crown position and for ponderosa pine whether they were killed or infested by mountain pine beetle. Elevation, slope, and aspect were also recorded. We used classification trees to model the likelihood of bark beetle attack based on plot and site variables. The probability of individual tree attack within the infested plots was estimated using logistic regression. Basal area of ponderosa pine in trees ≥25.4 cm in diameter at breast height (dbh) and ponderosa pine stand density index were correlated with mountain pine beetle attack. Regression trees and linear regression indicated that the amount of observed tree mortality was associated with initial ponderosa pine basal area and ponderosa pine stand density index. Infested stands had higher total and ponderosa pine basal area, total and ponderosa pine stand density index, and ponderosa pine basal area in trees ≥25.4 cm dbh. The probability of individual tree attack within infested plots was positively correlated with tree diameter with ponderosa pine stand density index modifying the relationship. A tree of a given size was more likely to be attacked in a denser stand. We conclude that stands with higher ponderosa pine basal area in trees >25.4 cm and ponderosa pine stand density index are correlated with an increased likelihood of mountain pine beetle bark beetle attack. Information form this study will help forest managers in the identification of uneven-aged stands with a higher likelihood of bark beetle attack and expected levels of tree mortality.     

Damage by pathogens and insects to Scots pine and lodgepole pine 25 years after reciprocal plantings in Canada and Sweden

A combined species – provenance – family experiment with Scots pine and lodgepole pine was planted in Canada and Sweden. One aim of the experiment was to evaluate the two species’ sensitivities to pathogens and insects 25 years after establishment in their non-native continents. In Canada, Scots pine had better average survival than lodgepole pine, but survival rates among trees from the best seed-lots were equal. In Canada only western gall rust infected Scots pine to some extent, and mountain pine beetles attacked and killed Scots pine more frequently than lodgepole pine. At one site in Sweden, lodgepole pine had higher survival rates than Scots pine, whether evaluated as an overall average or with data from only the best surviving seed-lots. At the other Swedish site, the species’ survival rates were equal, largely since moose damage was much more frequent on lodgepole pine than on Scots pine. Adaptation to local conditions seemed to be important in the resistance of stem breakage caused by heavy snowfalls. The exotic species generally seemed to resist the new threats, but more serious damage by mountain pine beetle on Scots pine than on lodgepole pine in Canada demonstrates possible drawbacks when using exotic species which are phylogenetically similar to native.     

Moisture and blue stain distribution in mountain pine beetle infested lodgepole pine trees and industrial implications

Healthy lodgepole pine trees and trees at three stages (green, red and grey) of infestation by mountain pine beetles (MPB) were examined for moisture and blue-stain distribution in the radial and longitudinal directions. The sapwood moisture content of trees dropped about 100% from the healthy stage while the heartwood moisture dropped about 10%, regardless of green, red or grey attack stages. Moisture content variation with tree height was unique up to the time since beetle attack. The volume of blue stain increased with increasing time since beetle attack. Maximum discolouration in wood occurred at about 3 m above ground level and expanded rapidly upward. Lumber sawn and tracked from the MPB attacked logs showed a decrease in moisture content with increasing time since beetle attack, indicating the need for specific drying schedules employing lower energy levels. The economic implication of the effect of blue stain was evaluated by the percentage of Japanese grade (J-grade) lumber produced. The J-grade lumber yield decreased with increasing time since beetle attack. Early harvest and conversion of MPB infected trees will positively impact product value and recovery. Part of this research was funded by Forestry Innovation Investment Ltd., Government of British Columbia.     

Response of bark beetles and their natural enemies to fire and fire surrogate treatments in mixed-conifer forests in western Montana

Four treatments (control, burn-only, thin-only, and thin-and-burn) were evaluated for their effects on bark beetle-caused mortality in both the short-term (one to four years) and the long-term (seven years) in mixed-conifer forests in western Montana, USA. In addition to assessing bark beetle responses to these treatments, we also measured natural enemy landing rates and resin flow of ponderosa pine (Pinus ponderosa) the season fire treatments were implemented. All bark beetles were present at low population levels (non-outbreak) for the duration of the study. Post-treatment mortality of trees due to bark beetles was lowest in the thin-only and control units and highest in the units receiving burns. Three tree-killing bark beetle species responded positively to fire treatments: Douglas-fir beetle (Dendroctonus pseudotsugae), pine engraver (Ips pini), and western pine beetle (Dendroctonus brevicomis). Red turpentine beetle (Dendroctonus valens) responded positively to fire treatments, but never caused mortality. Three fire damage variables tested (height of crown scorch, percent circumference of the tree bole scorched, or degree of ground char) were significant factors in predicting beetle attack on trees. Douglas-fir beetle and pine engraver responded rapidly to increased availability of resources (fire-damaged trees); however, successful attacks dropped rapidly once these resources were depleted. Movement to green trees by pine engraver was not observed in plots receiving fire treatments, or in thinned plots where slash supported substantial reproduction by this beetle. The fourth tree-killing beetle present at the site, the mountain pine beetle, did not exhibit responses to any treatment. Natural enemies generally arrived at trees the same time as host bark beetles. However, the landing rates of only one, Medetera spp., was affected by treatment. This predator responded positively to thinning treatments. This insect was present in very high numbers indicating a regulatory effect on beetles, at least in the short-term, in thinned stands. Resin flow decreased from June to August. However, resin flow was significantly higher in trees in August than in June in fire treatments. Increased flow in burned trees later in the season did not affect beetle attack success. Overall, responses by beetles to treatments were short-term and limited to fire-damaged trees. Expansions into green trees did not occur. This lack of spread was likely due to a combination of high tree vigor in residual stands and low background populations of bark beetles.     

The effects of mechanical fuel reduction treatments on the activity of bark beetles (Coleoptera: Scolytidae) infesting ponderosa pine

Selective logging, fire suppression, forest succession and climatic changes have resulted in high fire hazards over large areas of the western USA. Federal and state hazardous fuel reduction programs have increased accordingly to reduce the risk, extent and severity of these events, particularly in the wildland–urban interface. In this study, we examined the effects of mechanical fuel reduction treatments on the activity of bark beetles in ponderosa pine, Pinus ponderosa Dougl ex. Laws., forests located in Arizona and California, USA. Treatments were applied in both late spring (April–May) and late summer (August–September) and included: (1) thinned biomass chipped and randomly dispersed within each 0.4 ha plot; (2) thinned biomass chipped, randomly dispersed within each plot and raked 2 m from the base of residual trees; (3) thinned biomass lopped-and-scattered (thinned trees cut into 1–2 m lengths) within each plot; (4) an untreated control. The mean percentage of residual trees attacked by bark beetles ranged from 2.0% (untreated control) to 30.2% (plots thinned in spring with all biomass chipped). A three-fold increase in the percentage of trees attacked by bark beetles was observed in chipped versus lopped-and-scattered plots. Bark beetle colonization of residual trees was higher during spring treatments, which corresponded with peak adult beetle flight periods as measured by funnel trap captures. Raking chips away from the base of residual trees did not significantly affect attack rates. Several bark beetle species were present including the roundheaded pine beetle, Dendroctonus adjunctus Blandford (AZ), western pine beetle, D. brevicomis LeConte (AZ and CA), mountain pine beetle, D. ponderosae Hopkins (CA), red turpentine beetle, D. valens LeConte (AZ and CA), Arizona fivespined ips, Ips lecontei Swaine (AZ), California fivespined ips, I. paraconfusus Lanier (CA) and pine engraver, I. pini (Say) (AZ). Dendroctonus valens was the most common bark beetle infesting residual trees. A significant correlation was found between the number of trees chipped per plot and the percentage of residual trees with D. valens attacks. A significantly higher percentage of residual trees was attacked by D. brevicomis in plots that were chipped in spring compared to the untreated control. In lopped-and-scattered treatments, engraver beetles produced substantial broods in logging debris, but few attacks were observed on standing trees. At present, no significant difference in tree mortality exists among treatments. A few trees appeared to have died solely from D. valens attacks, as no other scolytids were observed in the upper bole. In a laboratory study conducted to provide an explanation for the bark beetle responses observed in this study, monoterpene elution rates from chip piles declined sharply over time, but were relatively constant in lopped-and-piled treatments. The quantities of β-pinene, 3-carene, -pinene and myrcene eluting from chips exceeded those from lopped-and-piled slash during each of 15 sample periods. These laboratory results may, in part, explain the bark beetle response observed in chipping treatments. The implications of these results to sustainable forest management are discussed.     

Aerially applied verbenone-releasing laminated flakes protect Pinus contorta stands from attack by Dendroctonus ponderosae in California and Idaho

We tested a new formulation of verbenone, an antiaggregation pheromone of the mountain pine beetle (Dendroctonus ponderosae Hopkins) (Coleoptera: Curculionidae: Scolytinae), for area-wide protection of lodgepole pine (Pinus contorta Douglas ex Loudon) stands in the western United States. Helicopter applications of verbenone-releasing laminated flakes were made at the rate of 370 g of active ingredient/ha to two sites, one in California and one in Idaho, during summer 2005. Each site consisted of five 20.23-ha treated plots and five matching 20.23-ha untreated plots. We assessed D. ponderosae flight into study plots using traps baited with aggregation pheromones and we tallied D. ponderosae attack rates on P. contorta trees in treated and control plots before and after application. There were no significant differences between numbers of D. ponderosae trapped on treated and control plots. However, a significantly smaller proportion of P. contorta trees was attacked in treated plots than in control plots at both sites; the attack rate in untreated stands was roughly three times that of treated stands in both California and Idaho, even at this low application rate. Furthermore, attack rate in 2004 was a significant explanatory variable for the response in 2005 regardless of treatment in both California and Idaho. There was no significant treatment effect at either site on attraction of Temnochila chlorodia (Coleoptera: Trogositidae) Mannerheim, a key predator of D. ponderosae, to the prey aggregation pheromone.     

Surveying mountain pine beetle damage of forests: A review of remote sensing opportunities

The severity and large spatial extent of the current mountain pine beetle outbreak in western North America has prompted research into methods for surveying the location and extent of beetle impacts using remotely sensed data. New forms of remotely sensed data and methods of image analysis have emerged in recent years, which have the potential to provide information on mountain pine beetle red-attack damage that complements information gathered from existing survey methods. The main objective of this review is to summarize previous and current contributions of remote sensing to the survey of mountain pine beetle impacts. The potential and limits of remotely sensed data for the detection and mapping of mountain pine beetle impacts, over a range of attack stages, are identified and synthesized. The secondary objective of this review is to highlight those methods or data sources that currently have operational potential and that the authors believe warrant further research in support of ongoing planning and management activities. Emphasis is placed on the different information needs associated with the various spatial scales of forest management: regional, landscape, and local. The findings of this review indicate that remotely sensed data is useful for mapping red-attack damage across a range of image data types and scales of inquiry. The review concludes with recommendations for future research, and suggestions for operational standards to guide the use of remotely sensed data for the survey of mountain pine beetle impacts.     

Bark beetle (Coleoptera: Scolytidae) diversity in spaced and unmanaged mature lodgepole pine (Pinaceae) in southeastern British Columbia

Variation in the number and diversity of bark beetles in spaced mature lodgepole pine stands in the East Kootenay region of British Columbia was analyzed in relation to location (site), spacing treatment and years following treatment. We analyzed the number of bark beetles and the number of bark beetle species that emerged from stumps or were captured in flight traps in the first five years following spacing. We also investigated the incidence of bark beetle attacks on the remaining trees and the mean dates of emergence from stumps and of capture in flight traps for the common species. Observations were made on three sites, each having three treatments: 4 m × 4 m spacing, 5 m × 5 m spacing, and an untreated control. The mean density of bark beetles emerged from stumps was different among sites and years but not between spacing treatments. There was no statistically significant variation in the number of bark beetle species captured in flight traps by site, spacing treatment, years, or spacing treatment and years. Significantly more bark beetles were captured in the 4 m × 4 m spacing treatment than in the control. The number of bark beetles captured was the highest in the first 2 years following treatment. Up to 26 species of bark beetles, excluding ambrosia beetles, were captured in flight barrier traps. There was no difference in species diversity by site or treatment indicating that species diversity in mature lodgepole pine is relatively stable over large areas. Of the 213 trees that sustained at least 10 attacks by bark beetles on the lower 2 m of the bole, 59.1% occurred in the spaced plots but only 18.2% of those were successful, versus 74.7% success in the infested trees in the control plots. The majority of infested trees contained Ips sp., Dendroctonus valens and D. murrayanae. Of the seven trees attacked by mountain pine beetle (D. ponderosae) only one tree was located in a spaced plot.     

Beetles, trees, and people: Regional economic impact sensitivity and policy considerations related to the mountain pine beetle infestation in British Columbia, Canada

This article investigates the regional economic impact sensitivity to the current mountain pine beetle infestation in five study areas of British Columbia, Canada, using a computable general equilibrium framework. Baseline general equilibrium economic databases are constructed for each region using a hybrid data collection approach involving primary and secondary sources. A computable general equilibrium model is constructed for each region and used to simulate the sensitivity of a suite of economic indicators to changes in forestry sector exports resulting from the impacts of mountain pine beetle on the available timber supply. The computable general equilibrium models constructed for each region provides an indication of economic vulnerability to the infestation and can assist decision-makers with the identification of policy considerations and priority areas for mitigation planning in response to the anticipated fluctuations in timber supply.     

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.     

Differences between provenances of pinus contort a var. latifolia in resistance to attack by gremmeniella abietina

A provenance trial of lodgepole pine (Pinus contorta var. latifolia) planted in 1969 at Matrand, Eidskog, Norway was apparently attacked by Gremmeniella abietina in 1984/85. In late autumn the damage was recorded separately on the lower and upper halves of the trees. The material ranged in latitude from 40.25°N to 56.28°N, and in altitude from 2 850 to 450 m a.s.l., highest in interior western United States and lowest in western Canada. The most heavily attacked sources were those from southern latitudes and from high elevations. The attack frequency changed gradually from south to north. In order to avoid attack by G. abietina, only well‐adapted provenances should be used.     

Stand characteristics and downed woody debris accumulations associated with a mountain pine beetle (Dendroctonus ponderosae Hopkins) outbreak in Colorado

Lodgepole pine (Pinus contorta Dougl. ex Loud.)-dominated ecosystems in north-central Colorado are undergoing rapid and drastic changes associated with overstory tree mortality from a current mountain pine beetle (Dendroctonus ponderosae Hopkins) outbreak. To characterize stand characteristics and downed woody debris loads during the first 7 years of the outbreak, 221 plots (0.02 ha) were randomly established in infested and uninfested stands distributed across the Arapaho National Forest, Colorado. Mountain pine beetle initially attacked stands with higher lodgepole pine basal area, and lower density and basal area of Engelmann spruce (Picea engelmannii [Parry]), and subalpine fir (Abies lasiocarpa (Hook.) Nutt. var. lasiocarpa) compared to uninfested plots. Mountain pine beetle-affected stands had reduced total and lodgepole pine stocking and quadratic mean diameter. The density and basal area of live overstory lodgepole declined by 62% and 71% in infested plots, respectively. The mean diameter of live lodgepole pine was 53% lower than pre-outbreak in infested plots. Downed woody debris loads did not differ between uninfested plots and plots currently infested at the time of sampling to 3 or 4–7 years after initial infestation, but the projected downed coarse wood accumulations when 80% of the mountain pine beetle-killed trees fall indicated a fourfold increase. Depth of the litter layer and maximum height of grass and herbaceous vegetation were greater 4–7 years after initial infestation compared to uninfested plots, though understory plant percent cover was not different. Seedling and sapling density of all species combined was higher in uninfested plots but there was no difference between infested and uninfested plots for lodgepole pine alone. For trees ≥2.5 cm in diameter at breast height, the density of live lodgepole pine trees in mountain pine beetle-affected stands was higher than Engelmann spruce, subalpine fir, and aspen, (Populus tremuloides Michx.), in diameter classes comprised of trees from 2.5 cm to 30 cm in diameter, suggesting that lodgepole pine will remain as a dominant overstory tree after the bark beetle outbreak.     

Differential effects of plant ontogeny and damage type on phloem and foliage monoterpenes in jack pine (Pinus banksiana)

Coniferous trees have both constitutive and inducible defences that deter or kill herbivores and pathogens. We investigated constitutive and induced monoterpene responses of jack pine (Pinus banksiana Lamb.) to a number of damage types: a fungal associate of the mountain pine beetle (Dendroctonus ponderosae Hopkins), Grosmannia clavigera (Robinson-Jeffrey & R.W. Davidson); two phytohormones, methyl jasmonate (MJ) and methyl salicylate (MS); simulated herbivory; and mechanical wounding. We only included the fungal, MJ and mechanical wounding treatments in the field experiments while all treatments were part of the greenhouse studies. We focused on both constitutive and induced responses between juvenile and mature jack pine trees and differences in defences between phloem and needles. We found that phytohormone applications and fungal inoculation resulted in the greatest increase in monoterpenes in both juvenile and mature trees. Additionally, damage types differentially affected the proportions of individual monoterpenes: MJ-treated mature trees had higher myrcene and β-pinene than fungal-inoculated mature trees, while needles of juveniles inoculated with the fungus contained higher limonene than MJ- or MS-treated juveniles. Although the constitutive monoterpenes were higher in the phloem of juveniles than mature jack pine trees, the phloem of mature trees had a much higher magnitude of induction. Further, induced monoterpene concentrations in juveniles were higher in phloem than in needles. There was no difference in monoterpene concentration between phytohormone applications and G. clavigera inoculation in mature trees, while in juvenile trees MJ was different from both G. clavigera and simulated herbivory in needle monoterpenes, but there was no difference between phytohormone applications and simulated herbivory in the phloem.     

Effects of mass inoculation on induced oleoresin response in intensively managed loblolly pine

Oleoresin flow is an important factor in the resistance of pines to attack by southern pine beetle, Dendroctonus frontalis Zimm., and its associated fungi. Abiotic factors, such as nutrient supply and water relations, have the potential to modify this plant-insect-fungus interaction; however, little is known of the effects of inoculation with beetle-associated fungi on oleoresin flow. We observed that constitutive and induced resin yield in loblolly pine, Pinus taeda L., were affected by either fungal inoculation (with the southern pine beetle-associated fungus Ophiostoma minus (Hedgcock) H. & P. Sydow) or silvicultural treatment. The effects of mass wounding (400 wounds m(-2)) and mass wounding and inoculation with O. minus were assessed by comparison with untreated (control) trees. The treatments were applied to trees in a 2 x 2 factorial combination of fertilizer and irrigation treatments. Fertilization did not significantly affect constitutive resin yield. Even as long as 105 days post-treatment, however, mass-inoculated trees produced higher induced resin yields than control or wounded-only trees, indicating a localized induced response to fungal inoculation. We noted no systemic induction of host defenses against fungal colonization. Although beetles attacking previously attacked trees face a greater resinous response from their host than beetles attacking trees that had not been previously attacked, the effect of an earlier attack may not last more than one flight season. Despite mass inoculations, O. minus did not kill the host trees, suggesting that this fungus is not a virulent plant pathogen.     

Assessing forest vulnerability and the potential distribution of pine beetles under current and future climate scenarios in the Interior West of the US

The aim of our study was to estimate forest vulnerability and potential distribution of three bark beetles (Curculionidae: Scolytinae) under current and projected climate conditions for 2020 and 2050. Our study focused on the mountain pine beetle (Dendroctonus ponderosae), western pine beetle (Dendroctonus brevicomis), and pine engraver (Ipspini). This study was conducted across eight states in the Interior West of the US covering approximately 2.2 million km² and encompassing about 95% of the Rocky Mountains in the contiguous US. Our analyses relied on aerial surveys of bark beetle outbreaks that occurred between 1991 and 2008. Occurrence points for each species were generated within polygons created from the aerial surveys. Current and projected climate scenarios were acquired from the WorldClim database and represented by 19 bioclimatic variables. We used Maxent modeling technique fit with occurrence points and current climate data to model potential beetle distributions and forest vulnerability. Three available climate models, each having two emission scenarios, were modeled independently and results averaged to produce two predictions for 2020 and two predictions for 2050 for each analysis. Environmental parameters defined by current climate models were then used to predict conditions under future climate scenarios, and changes in different species’ ranges were calculated. Our results suggested that the potential distribution for bark beetles under current climate conditions is extensive, which coincides with infestation trends observed in the last decade. Our results predicted that suitable habitats for the mountain pine beetle and pine engraver beetle will stabilize or decrease under future climate conditions, while habitat for the western pine beetle will continue to increase over time. The greatest increase in habitat area was for the western pine beetle, where one climate model predicted a 27% increase by 2050. In contrast, the predicted habitat of the mountain pine beetle from another climate model suggested a decrease in habitat areas as great as 46% by 2050. Generally, 2020 and 2050 models that tested the three climate scenarios independently had similar trends, though one climate scenario for the western pine beetle produced contrasting results. Ranges for all three species of bark beetles shifted considerably geographically suggesting that some host species may become more vulnerable to beetle attack in the future, while others may have a reduced risk over time.     

Influence of ozone and nitrogen deposition on bark beetle activity under drought conditions

Four years of severe drought from 1999 through 2003 led to unprecedented bark beetle activity in ponderosa and Jeffrey pine in the San Bernardino and San Jacinto Mountains of southern California. Pines in the San Bernardino Mountains also were heavily impacted by ozone and nitrogenous pollutants originating from urban and agricultural areas in the Los Angeles basin. We studied bark beetle activity and bark beetle associated tree mortality in pines at two drought-impacted sites in the San Bernardino Mountains, one receiving high levels of atmospheric pollutants, and one with more moderate atmospheric input. We also investigated the effects of nitrogen addition treatments of 0, 50 and 150 kg N ha⁻¹ year⁻¹ at each site. Tree mortality and beetle activity were significantly higher at the high pollution site. Differences in beetle activity between sites were significantly associated with ozone injury to pines, while differences in tree mortality between sites were significantly associated with both ozone injury and fertilization level. Tree mortality was 9% higher and beetle activity 50% higher for unfertilized trees at the high pollution site compared to the low pollution site. Tree mortality increased 8% and beetle activity increased 20% under the highest rates of nitrogen additions at the low pollution site. The strong response in beetle activity to nitrogen additions at the low pollution site suggests that atmospheric nitrogen deposition increased tree susceptibility to beetle attack at the high deposition site. While drought conditions throughout the region were a major factor in decreased tree resistance, it appears that both ozone exposure and atmospheric nitrogen deposition further increased pine susceptibility to beetle attack.     

Assessing changes in forest fragmentation following infestation using time series Landsat imagery

The current epidemic of mountain pine beetle (Dendroctonus ponderosae Hopkins) in British Columbia, Canada, has impacted an area of over 13 million hectares presenting a considerable challenge to provincial forest resource managers. Remote sensing technologies offer a highly effective tool to monitor this impact due to very large areas involved and its ability to detect dead and dying tree crowns. Conventionally, change detection procedures based upon spectral values have been applied; however, analysis of landscape pattern changes associated with long-time series change detection approaches present opportunities for the generation of unique and ecologically important information. This study is focussed on the detection and monitoring of the shape and area characteristics of lodgepole pine stands during mountain pine beetle infestation to quantify the progression of forest fragmentation and related loss of landscape connectivity. A set of landscape pattern indices were applied to a set of images consisting of six Landsat satellite images spanning the period from 1993 to 2006. Our results indicate that the impacts of the mountain pine beetle infestation on forest spatial pattern consist of an increase in the number of patches, an increase in forest patch shape complexity, a reduction in forest patch size, an increase in forest patch isolation, and a decrease in interspersion. These findings demonstrate the unique information available from long-time series satellite imagery combined with pattern analysis to better understand the combined effects of insect infestation and forest salvage and harvesting.     

Multiscale spatial variation of the bark beetle Ips sexdentatus damage in a pine plantation forest (Landes de Gascogne,Southwestern France)

Bark beetles are notorious pests of natural and planted forests causing extensive damage. These insects depend on dead or weakened trees but can switch to healthy trees during an outbreak as mass-attacks allow the beetle to overwhelm tree defences. Climatic events like windstorms are known to favour bark beetle outbreaks because they create a large number of breeding sites, i.e., weakened trees and for this reason, windthrown timber is generally preventively harvested and removed. In December 1999, the southwest of France was struck by a devastating windstorm that felled more that 27 million m³of timber. This event offered the opportunity to study large-scale spatial pattern of trees attacked by the bark beetle Ips sexdentatus and its relationship with the spatial location of pine logs that were temporally stored in piles along stand edges during the post-storm process of fallen tree removal. The study was undertaken in a pure maritime pine forest of 1300 ha in 2001 and 2002. We developed a landscape approach based on a GIS and a complete inventory of attacked trees. During this study more than 70% of the investigated stands had at least one tree attacked by I. sexdentatus  . Spatial aggregation prevailed in stands with n≥15$n\ge 15$ attacked trees. Patches of attacked trees were identified using a kernel estimation procedure coupled with randomization tests. Attacked trees formed patches of 500–700 m² on average which displayed a clumped spatial distribution. Log piles stemming from the sanitation removals were mainly distributed along the large access roads and showed an aggregated spatial pattern as well. The spatial relationship between patches of attacked trees and log pile storage areas was analyzed by means of the Ripley’s statistic that revealed a strong association at the scale of the studied forest. Our results indicated that bark beetle attacks were facilitated in the vicinity of areas where pine logs were stored. The spatial extent of this relationship was >1000 m. Similar results were obtained in 2001 and 2002 despite differences in the number and spatial distribution of attacked trees. The presence of a strong “facilitation effect” suggests that log piles should be removed quickly in order to prevent outbreaks of bark beetles.     

Breeding efficiency in Tomicus piniperda and shoot damage after late autumn thinning of young Pinus sylvestris stands

The breeding of the pine shoot beetle (Tomicus piniperda) after thinning of young Scots pine stands in late autumn, and subsequent shoot damage, were studied in southern, central and northern Finland during three successive years (1977–79). No distinct increase in the attack density or beetle population was observed on the felled trees during the study period. Shoot damage on the standing trees remained low, and was not considered to have any marked negative effect on tree growth or further development of the pine stand. No noticeable difference was found between the different parts of the country. The economic importance of beetle damage resulting from late autumn thinnings was estimated to be low.