Quantifying spatio-temporal dispersion of bark beetle infestations in epidemic and non-epidemic conditions

Spatio-temporal dispersal of pest species such as bark beetles plays a key role in their population ecology and outbreak dynamics. Understanding the underlying patterns is crucial for applying appropriate management strategies.In contrast to most existing studies which focus on dispersing beetles, we analysed patches of killed trees resulting from bark beetle infestation. The study was based on a 22-year time series of annually captured colour-infrared (CIR) images of the Bavarian Forest National Park (Germany), where Ips typographus L. (Coleoptera, Curculionidae, Scolytinae) propagates undisturbed by human activity. Newly infested patches comprising at least 5 spruce trees were identified in every time step. This investigation of spatio-temporal spread of infestations primarily focused on (i) parameterizing the size and shape of infestation patches, (ii) modelling an infestation gradient and (iii) evaluating the risk of subsequent infestations on landscape scale. We developed a GIS-based distance ring approach to quantify the distance relation of subsequent infestations, including the distribution of potential hosts.Infestation spread was revealed to be strongly distance dependent, following an inverse power law function: on average 65% of new infestations occurred within a 100 m radius of the previous year’s infestations, and 95% within 500 m. ‘Distance’ proved to be a major determinant of I. typographus dispersal on the landscape scale in each time step of the 22-year series we investigated. Infestation distance thus describes the outcome of beetle dispersal very accurately. The time series showed two alternating periods of epidemic and non-epidemic infestation. These gradation stages did not affect the size and shape of infested patches, but epidemics correlated significantly with a higher percentage of infestations within short distances. Additionally, the resulting infestation risk is highly sensitive to the gradation stage, particularly within the first 100 m around source spots where it increases up to 30%.Our study therefore contributes to a better understanding of the outbreak dynamics of I. typographus and suggests concentrating efficient bark beetle management on areas in the close vicinity of previous years’ infestations.     

Prescribed fire effects on bark beetle activity and tree mortality in southwestern ponderosa pine forests

Prescribed fire is an important tool in the management of ponderosa pine (Pinus ponderosa Dougl. ex Laws.) forests, yet effects on bark beetle (Coleoptera: Curculionidae, Scolytinae) activity and tree mortality are poorly understood in the southwestern U.S. We compared bark beetle attacks and tree mortality between paired prescribed-burned and unburned stands at each of four sites in Arizona and New Mexico for three growing seasons after burning (2004–2006). Prescribed burns increased bark beetle attacks on ponderosa pine over the first three post-fire years from 1.5 to 13% of all trees, increased successful, lethal attacks on ponderosa pine from 0.4 to 7.6%, increased mortality of ponderosa pine from all causes from 0.6 to 8.4%, and increased mortality of all tree species with diameter at breast height >13 cm from 0.6 to 9.6%. On a per year basis, prescribed burns increased ponderosa pine mortality from 0.2% per year in unburned stands to 2.8% per year in burned stands. Mortality of ponderosa pine 3 years after burning was best described by a logistic regression model with total crown damage (crown scorch + crown consumption) and bark beetle attack rating (no, partial, or mass attack by bark beetles) as independent variables. Attacks by Dendroctonus spp. did not differ significantly over bole heights, whereas attacks by Ips spp. were greater on the upper bole compared with the lower bole. Three previously published logistic regression models of tree mortality, developed from fires in 1995–1996 in northern Arizona, were moderately successful in predicting broad patterns of tree mortality in our data. The influence of bark beetle attack rating on tree mortality was stronger for our data than for data from the 1995–1996 fires. Our results highlight canopy damage from fire as a strong and consistent predictor of post-fire mortality of ponderosa pine, and bark beetle attacks and bole char rating as less consistent predictors because of temporal variability in their relationship to mortality. The small increase in tree mortality and bark beetle attacks caused by prescribed burning should be acceptable to many forest managers and the public given the resulting reduction in surface fuel and risk of severe wildfire.     

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.     

Carbon and nitrogen cycling immediately following bark beetle outbreaks in southwestern ponderosa pine forests

Bark beetle infestation is a well-known cause of historical low-level disturbance in southwestern ponderosa pine forests, but recent fire exclusion and increased tree densities have enabled large-scale bark beetle outbreaks with unknown consequences for ecosystem function. Uninfested and beetle-infested plots (n = 10 pairs of plots on two aspects) of ponderosa pine were compared over one growing season in the Sierra Ancha Experimental Forest, AZ to determine whether infestation was correlated with differences in carbon (C) and nitrogen (N) pools and fluxes in aboveground biomass and soils. Infested plots had at least 80% of the overstory ponderosa pine trees attacked by bark beetles within 2 years of our measurements. Both uninfested and infested plots stored ∼9 kg C m⁻² in aboveground tree biomass, but infested plots held 60% of this aboveground tree biomass in dead trees, compared to 5% in uninfested plots. We hypothesized that decreased belowground C allocation following beetle-induced tree mortality would alter soil respiration rates, but this hypothesis was not supported; throughout the growing season, soil respiration in infested plots was similar to uninfested plots. In contrast, several results supported the hypothesis that premature needlefall from infested trees provided a pulse of low C:N needlefall that altered soil N cycling. The C:N mass ratio of pine needlefall in infested plots (∼45) was lower than uninfested plots (∼95) throughout the growing season. Mineral soils from infested plots had greater laboratory net nitrification rates and field resin bag ammonium accumulation than uninfested plots. As bark beetle outbreaks become increasingly prevalent in western landscapes, longer-term biogeochemical studies on interactions with other disturbances (e.g. fire, harvesting, etc.) will be required to predict changes in ecosystem structure and function.     

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.     

Bark beetle-caused mortality in a drought-affected ponderosa pine landscape in Arizona,USA

Extensive ponderosa pine (Pinus ponderosa Dougl. ex Laws.) mortality associated with a widespread severe drought and increased bark beetle (Coleoptera: Curculionidae, Scolytinae) populations occurred in Arizona from 2001 to 2004. A complex of Ips beetles including: the Arizona fivespined ips, Ips lecontei Swaine, the pine engraver beetle, Ips pini (Say), Ips calligraphus (Germar), Ips latidens (LeConte), Ips knausi Swaine and Ips integer (Eichhoff) were the primary bark beetle species associated with ponderosa pine mortality. In this study we examine stand conditions and physiographic factors associated with bark beetle-caused tree mortality in ponderosa pine forests across five National Forests in Arizona. A total of 633 fixed-radius plots were established across five National Forests in Arizona: Apache-Sitgreaves, Coconino, Kaibab, Prescott, and Tonto. Prior to the bark beetle outbreak, plots with mortality had higher tree and stocking compared with plots without pine mortality. Logistic regression modeling found that probability of ponderosa pine mortality caused by bark beetles was positively correlated with tree density and inversely related with elevation and tree diameter. Given the large geographical extent of this study resulting logistic models to estimate the likelihood of bark beetle attack should have wide applicability across similar ponderosa pine forests across the Southwest. This is particularly true of a model driven by tree density and elevation constructed by combining all forests. Tree mortality resulted in significant reductions in basal area, tree density, stand density index, and mean tree diameter for ponderosa pine and for all species combined in these forests. Most of the observed pine mortality was in the 10–35 cm diameter class, which comprise much of the increase in tree density over the past century as a result of fire suppression and grazing practices. Ecological implications of tree mortality are discussed.     

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.     

Multi-scale nest-site selection by black-backed woodpeckers in outbreaks of mountain pine beetles

Areas of mountain pine beetle (Dendroctonus ponderosae Hopkins) outbreaks in the Black Hills can provide habitat for black-backed woodpeckers (Picoides arcticus), a U.S. Forest Service, Region 2 Sensitive Species. These outbreaks are managed through removal of trees infested with mountain pine beetles to control mountain pine beetle populations and salvage timber resources. To minimize impacts to black-backed woodpeckers while meeting management objectives, there is a need to identify characteristics of these areas that support black-backed woodpeckers. We examined the habitat associations of this species nesting in areas of beetle outbreaks in the Black Hills, South Dakota in 2004 and 2005. We used an information theoretic approach and discrete choice models to evaluate nest-site selection of 42 woodpecker nests at 3 spatial scales—territory, nest area, and nest tree. At the territory scale (250 m around nest), availability and distribution of food best explained black-backed woodpecker selection of beetle outbreaks versus the surrounding forest. Selection at the territory scale was positively associated with densities of trees currently infested by mountain pine beetles and indices of wood borer (Cerambycidae and Buprestidae) abundance, and was greatest at distances of 50–100 m from the nearest patch of infestation. At the nest-area scale (12.5 m radius around the nest), densities of snags positively influenced nest-area selection. Finally, at the nest-tree scale, aspen (Populus tremuloides) and 3–5-year-old ponderosa pine (Pinus ponderosa) snags were important resources. The association between abundant wood-boring insects and black-backed woodpeckers creates a difficult challenge for forest managers. In the absence of fire, areas of beetle outbreak might serve as the only substantial source of habitat in the Black Hills. Regulating insect populations via salvage logging will reduce key food resources to black-backed woodpeckers during nesting. Therefore, given the relatively infrequent occurrence of large-scale fire in the Black Hills, management should recognize the importance of beetle-killed forests to the long-term viability of the black-backed woodpecker population in the Black Hills.     

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.     

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.     

Effects of prescribed fire and season of burn on direct and indirect levels of tree mortality in Ponderosa and Jeffrey Pine Forests in California,USA

Many forests that historically experienced frequent low-intensity wildfires have undergone extensive alterations during the past century. Prescribed fire is now commonly used to restore these fire-adapted forest ecosystems. In this study, we examined the influence of prescribed burn season on levels of tree mortality attributed to prescribed fire effects (direct mortality) and bark beetles (Coleoptera: Curculionidae, Scolytinae) (indirect mortality) in ponderosa pine, Pinusponderosa Dougl. ex Laws., and Jeffrey pine, Pinusjeffreyi Grev. and Balf., forests in California, USA. A total of 816 trees (9.9% of all trees) died during this 3-yr study. Significantly higher levels of tree mortality (all sources) occurred following early and late season burns compared to the untreated control, but no significant difference was observed between burn treatments. The majority (461 trees) of tree deaths were attributed to direct mortality from prescribed burns and was strongly concentrated (391 trees) in the smallest diameter class (<20.2 cm diameter at breast height, dbh). For the largest trees (>50.7 cm dbh), significantly higher levels of tree mortality occurred on early season burns than the untreated control, most of which resulted from indirect mortality attributed to bark beetle attacks, specifically western pine beetle, Dendroctonus brevicomis LeConte, and mountain pine beetle, D. ponderosae Hopkins. Red turpentine beetle, D. valens LeConte, was the most common bark beetle species found colonizing trees, but tree mortality was not attributed to this species. A total of 355 trees (4.3% of all trees) were killed by bark beetles. Dendroctonus brevicomis (67 trees, 18.9%) and D. ponderosae (56 trees, 15.8%), were found colonizing P. ponderosa; and Jeffrey pine beetle, D. jeffreyi Hopkins, was found colonizing P. jeffreyi (seven trees, 2.0%). We also found pine engraver, Ips pini (Say) (137 trees, 38.6%), and, to a much lesser extent, Orthotomicus (=Ips) latidens (LeConte) (85 trees, 23.9%) and emarginate ips, I. emarginatus (LeConte) (3 trees, 0.8%) colonizing P. ponderosa and P. jeffreyi. Few meaningful differences in levels of indirect tree mortality attributed to bark beetle attack were observed between early and late season burns. The incidence of root and root collar pathogens (Leptographium and Sporothrix spp.), including species known to be vectored by bark beetles, was low (18% of trees sampled). The implications of these and other results to management of P. ponderosa and P. jeffreyi forests are discussed in detail.     

Ophiostomatoid fungi associated with bark beetles onAbies veitchii in wave-regenerated forests

Relationships between tree mortality and bark beetle infestation onAbies veitchii at the wave-regenerated forest in Mt. Asahi, Okuchichibu area were investigated. Most of the firs with green needles and newly developed current year’s shoots in the dieback zone were heavily infested by bark beetles before the death of the trees. After heavy infestation of beetles, about half of the infested firs died within the year, and the other half died in the next year. When the species composition of bark beetles and associated ophiostomatoid fungi were investigated in Mt. Asahi and also at a typical wave-regenerated forest in Mt. Shimagare, Yatsugatake area,Cryphalus montanus andC. piceae were dominant beetle species for Mt. Asahi and Mt. Shimagare, respectively.Ophiostoma subalpinum andO. europhioides were dominant fungal species at both wave-generated forests. BecauseO. subalpinum was detected more frequently from deeper areas of sapwood thanO. europhioides, it was suggested that the fungal species may accelerate the death of stressed firs in wave-regenerated forests. Contribution No. 169, Laboratory of Plant Parasitic Mycology, Institute of Agriculture and Forestry, University of Tsukuba.     

Determining the vulnerability of Mexican pine forests to bark beetles of the genus Dendroctonus Erichson (Coleoptera: Curculionidae: Scolytinae)

Bark beetles of the genus Dendroctonus are natural inhabitants of forests; under particular conditions some species of this genus can cause large-scale tree mortality. However, only in recent decades has priority been given to the comprehensive study of these insects in México. México possesses high ecological diversity in Dendroctonus–Pinus associations. The geographic coexistence of 12 Dendroctonus species suggests greater vulnerability or threat of tree mortality relative to other areas. We use a biogeographic strategy to identify and rank the areas most vulnerable to tree mortality caused by bark beetles in México. We aim to define the areas that might experience high impact by these insects and also to provide a geographic database useful to forest resource management and conservation policies in México. Using collection records of bark beetles and pines, we develop a quantitative estimate of the threat of beetle infestation of forest areas based on factors including pine and beetle species density, host preference and level of mortality caused by beetle species. A quantitative estimate of forest area vulnerability, the Bark Beetle Threat Index (BBTI) was calculated. Despite the vast area of geographic coincidence of Pinus and Dendroctonus in México, the regions of highest bark beetle pressure are restricted to small zones within some mountain systems. The region that has been most affected by this insect group during the past hundred years is the Transverse Volcanic Belt, followed by the Sierra Madre Occidental and Sierra Madre del Sur. Pine diversity is the major determining factor of BBTI at the regional level, while disturbances from extensive logging and ecosystem change are the key factors behind high BBTIs at the local level.     

Factors affecting the spatio-temporal dispersion of Ips typographus (L.) in Bavarian Forest National Park: A long-term quantitative landscape-level analysis

The relationship between abiotic and biotic factors and the spread of the European spruce bark beetle, Ips typographus (L.), was investigated at a landscape level over a model period of 18 years in the Bavarian Forest National Park in Germany. Deadwood areas - where I. typographus - caused tree mortality of 100% - were photographed annually using Color-infrared aerial photography and digitally recorded in vector form. Thirty-two static and dynamic habitat variables were quantitatively determined using spatial pattern analysis and geostatistics from 1990 to 2007 at the landscape scale. The importance of the presence of deadwood areas for thirty-two habitat variables for the occurrence of the bark beetle was quantitatively recorded using an Ecological Niche Factor Analysis (ENFA).It was shown over a long model period that the intensity of the bark beetle infestation went through different phases over the 18-year study period. No mono-causal correlations could be found between individual habitat factors and the spread of the bark beetle over the entire model period. On the one hand, these findings underline the complexity of the system, on the other hand, this could be interpreted as a possible explanation for conclusions drawn by previous studies that differ from each other.The importance of individual habitat variables and the combinations of variables varied to different extents within these phases. An examination of the cumulative importance of the habitat demonstrated that the biological structural variables such as the distance from the site of the previous year's infestation, the area and the perimeter of the infested areas from the previous year are of great importance for the incidence of the bark beetle, but not across all years. Of equal significance for assessing the size of the area and the distance of the deadwood areas from the sites of the previous year's infestation are the size of the areas, the perimeter of the deadwood areas and the proximity index. An evaluation of the stages of forest succession showed that cumulatively, a short distance between the infested areas and the forest areas with conifers in the early stages of growth was an equally important habitat factor from 1990 to 2007. By quantitatively recording habitat factors that are significant for the spread of the bark beetle it may help predict areas that are at risk and thus to develop suitable management strategies to minimise or stop the spread and the effect of the bark beetle.     

Effects of forest restoration treatments on the abundance of bark beetles in Norway spruce forests of southern Finland

Restoration of protected areas in boreal forests frequently includes creating substantial volumes of dead wood. While this benefits a wide range of dead wood dependent invertebrate species, some of these are regarded as forest pests. Therefore, the risk of elevated levels of tree mortality in surrounding commercial forests must be considered. In a large-scale field experiment in southern Finland, we studied the effects of restoration treatments on the abundance of bark beetles within and in the vicinity of restored areas, in particular focusing on Ips typographus and Pityogenes chalcographus. The treatments applied to managed Norway spruce forests were controlled burning and partial harvesting combined with retaining 5, 30 or 60 m³/ha of cut down wood. We found that the abundance of bark beetles increased by both burning and harvesting with down wood retention, being highest where burning and harvesting had been combined. The actual volume of down wood retention had no significant effect. The effect of burning on the number of bark beetles along host tree boles was negative which suggests that burnt spruces provided a less suitable resource for bark beetles than unburnt dead spruces. The abundance of bark beetles along host trees also decreased with increasing volume of down wood retention. The abundance of P. chalographus was slightly elevated up to 50 m outside restored areas but the abundance was very low compared to that within the areas. The abundance of I. typographus was extremely low outside restored areas. We conclude that restoration treatments increase the abundance of bark beetles via increased availability of resources, but that the effect of burning is likely to be counteracted by decreased resource quality. Thus, burning might be the “safest” way to produce large quantities of dead wood. Furthermore, the fact that only few beetles were collected in adjacent areas suggests that restored areas pose little threat of serving as refugia in which bark beetle populations increase in sufficient numbers to attack live trees in adjacent forests. However, restoration actions repeated at consecutive years within a small area might enable the populations to grow to outbreak levels.     

Protection of spruce from colonization by the bark beetle, Ips perturbatus, in Alaska

Field bioassays were conducted in south-central Alaska in a stand of Lutz spruce, Picea × lutzii, to determine whether a semiochemical interruptant (verbenone and trans-conophthorin) and/or a defense-inducing plant hormone (methyl jasmonate, MJ) could be used to protect individual standing trees from bark beetle attack. During two experiments (initiated in May 2004 and 2005, respectively), attacks by Ips perturbatus on standing trees were induced by using a three-component aggregation pheromone (ipsenol, cis-verbenol, and ipsdienol) and prevented by using the interruptant. In 2005, treatments from 2004 were repeated and additional treatments were evaluated by using MJ spray or injection with and without the interruptant. Aggregation began before 3 or 7 June, and attack density was monitored through 3 or 16 August. During both years, tree mortality caused by I. perturbatus was recorded twice (in August, and in May of the following year). In both experiments, attack density was greatest on trees baited with the three-component attractive pheromone, but was significantly reduced by addition of the semiochemical interruptant to trees baited with the attractant. There were no significant differences in attack density between attractant + interruptant-treated trees and unbaited trees. In 2004, mortality was highest among attractant-baited trees, whereas addition of the interruptant significantly reduced the level of initial (10 week post-treatment) and final (54 week post-treatment) mortality. In 2005, no significant reduction in attack density occurred on trees baited with the attractant when MJ was sprayed or injected. The highest initial (10.6 week post-treatment) and final (49.4 week post-treatment) mortality was observed among trees that had been injected with MJ and baited with the attractant. Mortality at the final assessment was significantly lower in all other treatment groups. As in 2004, addition of the interruptant to attractant-baited trees significantly reduced the level of final mortality compared to attractant-baited trees. MJ was not attractive or interruptive to I. perturbatus or associated bark beetles in a flight trapping study. However, MJ-treated trees (sprayed or injected) exuded copious amounts of resin on the bark surface. Anatomical analyses of felled trees from four treatment groups [Tween (solvent)-sprayed, MJ-sprayed, Tween-injected, and MJ-injected + attractant baited] showed that treatment with MJ increased the number and size of resin ducts produced following treatment. These analyses also revealed a reduction in radial growth in MJ-treated trees. Our results show that during both years, treatment with a simple, two-component interruptant system of verbenone and trans-conophthorin significantly reduced I. perturbatus attack density and tree mortality on attractant-baited trees and provided a full year of protection from bark beetle attack.     

Forest thinning and subsequent bark beetle-caused mortality in Northeastern California

The Warner Mountains of northeastern California on the Modoc National Forest experienced a high incidence of tree mortality (2001–2007) that was associated with drought and bark beetle (Coleoptera: Curculionidae, Scolytinae) attack. Various silvicultural thinning treatments were implemented prior to this period of tree mortality to reduce stand density and increase residual tree growth and vigor. Our study: (1) compared bark beetle-caused conifer mortality in forested areas thinned from 1985 to 1998 to similar, non-thinned areas and (2) identified site, stand and individual tree characteristics associated with conifer mortality. We sampled ponderosa pine (Pinus ponderosa var ponderosa Dougl. ex Laws.) and Jeffrey pine (Pinus jeffreyi Grev. and Balf.) trees in pre-commercially thinned and non-thinned plantations and ponderosa pine and white fir (Abies concolor var lowiana Gordon) in mixed conifer forests that were commercially thinned, salvage-thinned, and non-thinned. Clusters of five plots (1/50th ha) and four transects (20.1 × 100.6 m) were sampled to estimate stand, site and tree mortality characteristics. A total of 20 pre-commercially thinned and 13 non-thinned plantation plot clusters as well as 20 commercially thinned, 20 salvage-thinned and 20 non-thinned mixed conifer plot clusters were established. Plantation and mixed conifer data were analyzed separately. In ponderosa pine plantations, mountain pine beetle (Dendroctonus ponderosae Hopkins) (MPB) caused greater density of mortality (trees ha⁻¹ killed) in non-thinned (median 16.1 trees ha⁻¹) compared to the pre-commercially thinned (1.2 trees ha⁻¹) stands. Percent mortality (trees ha⁻¹ killed/trees ha⁻¹ host available) was less in the pre-commercially thinned (median 0.5%) compared to the non-thinned (5.0%) plantation stands. In mixed conifer areas, fir engraver beetles (Scolytus ventralis LeConte) (FEN) caused greater density of white fir mortality in non-thinned (least square mean 44.5 trees ha⁻¹) compared to the commercially thinned (23.8 trees ha⁻¹) and salvage-thinned stands (16.4 trees ha⁻¹). Percent mortality did not differ between commercially thinned (least square mean 12.6%), salvage-thinned (11.0%), and non-thinned (13.1%) mixed conifer stands. Thus, FEN-caused mortality occurred in direct proportion to the density of available white fir. In plantations, density of MPB-caused mortality was associated with treatment and tree density of all species. In mixed conifer areas, density of FEN-caused mortality had a positive association with white fir density and a curvilinear association with elevation.     

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.     

Colonisation of native and exotic conifers by indigenous bark beetles (Coleoptera: Scolytinae) in France

Planting exotic conifers offers indigenous forest insects an opportunity to extend their host range and eventually to become significant pests. Knowing the ecological and evolutionary modalities driving the colonisation of exotic tree species by indigenous insects is thus of primary importance. We compared the bark beetle communities (Coleoptera: Curculionidae, Scolytinae) associated with both native and introduced conifers in France. The aim of our study was to estimate the influence of both host- and insect-related factors on the beetles’ likelihood to shift onto new hosts. We considered the influence of host origin (i.e. native vs. exotic), host tree species identity, tree bark thickness and tree taxonomic proximity, as well as insects’ host specificity. A field inventory using trap trees was carried out in two regions in France (Limousin and Jura) during two consecutive years (2006 and 2007) on three European native conifer species [Norway spruce (Picea abies); Scots pine (Pinus sylvestris) and European Silver-fir (Abies alba)] and five North American [Sitka spruce (Picea sitchensis); Eastern white pine (Pinus strobus); Grand fir (Abies grandis); Douglas fir (Pseudotsuga menziesii) and Western red cedar (Thuja plicata)]. A total of 18 indigenous and 2 exotic bark beetle species were collected. All exotic conifer species were colonised by indigenous bark beetle species and no significant difference was observed of the cumulated species richness of the latter between native and exotic tree species (13 vs. 14, P < 0.05). The ability of indigenous bark beetles to shift onto exotic conifers appeared to strongly depend on host species (significantly structuring bark beetle assemblages), the presence of phylogenetically related native conifer species and that of similar resources, in combination with insect host specificity. Host tree species status (native or exotic) also seemed to be involved, but its effect did not seem as essential as that of the previous factors. These findings are discussed in terms of adaptation, plasticity and practical aspects of forest management.     

The “sun-effect”: microclimatic alterations predispose forest edges to bark beetle infestations

Bark beetle dispersal and host selection behaviour are a complex and poorly understood process, resulting in specific spatio-temporal infestation patterns in forests. Aerial images from the Bavarian Forest National Park (Germany) provide a high-resolution, that is, tree-scale data set for the period 2001–2010, including information about Ips typographus (Col., Curculio., Scolytinae) infestation, the application of sanitary logging, natural forest edges and the area of living spruce susceptible to bark beetle infestation. We combined methods of GIS and image analysis to investigate the infestation probabilities at three types of forest edges under spatial and temporal aspects and compared them to the corresponding probabilities at the stand interior. Our results showed a pronounced infestation predisposition of such edge trees delimiting infestation patches cleared by sanitary logging measures, in particular at the south-facing edge sector. In contrast, edges adjacent to non-cleared infestation were revealed as less attractive for subsequent infestations, but nonetheless more attractive than permanent forest edges or the stand interior. Additionally, we measured near-bark surface air temperature to determine microclimatic differences at those edge- or non-edge sites and related them to predisposition results. Finally, our study emphasized favourable microclimatic conditions—summarized as the “sun-effect”—as a decisive factor enhancing the local infestation probability at recent forest edges in multiple ways. Both insect- and host tree-related reactions to suddenly altered microclimate are supposed to bias arbitrary colonization behaviour at patch and tree level, thereby mainly explaining observed infestation patterns. From the forester’s point of view, our results may contribute to precise bark beetle risk assessment and thus facilitate decision making in forest management.