Bark Beetle Attack Research Articles

Oleoresin yield of Pinus elliottii in a subtropical climate: Seasonal variation and effect of auxin and salicylic acid-based stimulant paste

The extraction of induced oleoresin from Pinus elliottii through bark streaking supplies products to the food, pharmaceutical and chemical industries. Resin flow is a typical defense response in conifers and is controled by various environmental factors. Although many studies have focused on oleoresin production in temperate forests, particularly in response to bark beetle attacks, relatively little is known about the physiological regulation of sustainable resin exudation in subtropical planted forests, in which commercial exploitation of this product is favored by the absence of significant pest insect populations. Therefore, the annual oleoresin yield of approximately 2660, 28-year-old trees from different sites was monitored for two consecutive years in southern Brazil. Plain wounding and wounding followed by application of a commercial resin stimulant paste containing the ethylene precursor 2-chloroethylphosphonic acid (CEPA) were used as reference treatments. Different concentrations of auxin, paraquat, yeast extract, and salicylic acid (isolated or in combination with CEPA) were also tested. Higher resin yields were associated with spring and summer, whereas winter had lower but significant yields. Auxin and salicylic acid stimulated the production of oleoresin relative to commercial paste in at least one concentration. Replacement of CEPA with auxin or salicylic acid yielded equivalent oleoresin production, indicating a comparable or complementary effect of these molecules in the process, as well as having implications for new stimulant paste formulations.

A temporal analysis of antioxidative defense responses in the phloem of Picea abies after attack by Ips typographus

The temporal gradation of antioxidants was investigated on the phloem tissue of Norway spruce [Picea abies (L.) Karst.] in response to weather conditions and colonization levels of Ips typographus L. (Col., Scolytidae). Two weeks after pheromone dispensers were placed on trees, the initial reaction of Norway spruce to bark beetle attack resulted in moderately lowered levels of total glutathione (tGSH) and total cysteine. Likewise, the total ascorbic acid dropped slightly below the control levels, whereas the concentration of dehydroascorbic acid increased in comparison to the first sampling date. This transient degradation and oxidation of the glutathione and ascorbate system was accompanied by moderately increased concentrations of total phenolics. One month later, the shift in antioxidant balance after moderate attack differed quantitatively from the reaction after massive attack. An intensification of antioxidant defense occurred within moderately affected bark. Total cysteine and tGSH contents were markedly raised, whereas the concentrations of total ascorbic acid and total phenolics were slightly increased by moderate attack. On the other hand, massive bark beetle colonization caused a strong decrease in tGSH and total phenolics, whereas total cysteine and total ascorbic acid values remained at control level. Dependent upon the intensity and the success of the attack, a progressive degradation of antioxidants was determined at later sampling dates, which was accompanied by an obvious oxidation of the ascorbate and glutathione pools. With an unsuccessful defense upon massive attack, the thiols and total phenolics did not reach a new steady state, but deteriorated until the end of the brood beetles' development. In contrast, the dynamic antioxidative response within the moderately affected trees indicated an acclimation stage in the middle of July. It was characterized by a higher accumulation of tGSH, total ascorbic acid and total phenolics as well as a more reduced redox state of glutathione. A sequence of changes in the endogenous levels of antioxidant defense molecules in the bark beetle-affected Norway spruce showed consistency with the general ecophysiological stress-response concept, and provided important avenues for evaluating the role and effectiveness of antioxidants in systemic acquired resistance against the complex interactive effects of bark beetle attack and environmental factors.

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 m 3of 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 attacked trees. Patches of attacked trees were identified using a kernel estimation procedure coupled with randomization tests. Attacked trees formed patches of 500–700 m 2 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.

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.

Attraction of ambrosia and bark beetles to coast live oaks infected by Phytophthora ramorum

Abstract1 Sudden oak death is caused by the apparently introduced oomycete, Phytophthora ramorum. We investigated the role of bark and ambrosia beetles in disease progression in coast live oaks Quercus agrifolia.2 In two Marin County, California sites, 80 trees were inoculated in July 2002 with P. ramorum and 40 were wounded without inoculation. Half of the trees in each group were sprayed with the insecticide permethrin [cyclopropanecarboxylic acid, 3‐(2,2‐dichloroethenyl)‐2,2‐dimethyl‐(3‐phenoxyphenyl) methyl ester] to prevent ambrosia and bark beetle attacks, and then were sprayed twice per year thereafter. After each treatment, sticky traps were placed on only the permethrin‐treated trees. Beetles were collected periodically in 2003.3 Inoculated trees accounted for 95% of all beetles trapped. The ambrosia beetles Monarthrum scutellare and Xyleborinus saxeseni and the western oak bark beetle Pseudopityophthorus pubipennis were the most abundant of the seven species trapped.4 Permethrin treatment delayed initiation of beetle attacks and significantly reduced the mean number of attacks per tree. Beetles did not attack any wounded or noncankered inoculated trees.5 Trees with larger cankers trapped more beetles early in the disease. Once permethrin lost effectiveness, the number of beetle entrance tunnels was a more reliable predictor of subsequent trap catch than was canker size.6 Beetles were initially attracted to P. ramorum cankers in response to kairomones generated in the host‐pathogen interaction. After beetles attacked the permethrin‐treated trees, aggregation pheromones most probably were the principal factor in beetle colonization behaviour.

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.

Spray Deposition from Ground‐based Applications of Carbaryl to Protect Individual Trees from Bark Beetle Attack

Bark beetles (Coleoptera: Curculionidae, Scolytinae) are recognized as the most important tree mortality agent in western coniferous forests. A common method of protecting trees from bark beetle attack is to saturate the tree bole with carbaryl (1-naphthyl methylcarbamate) using a hydraulic sprayer. In this study, we evaluate the amount of carbaryl drift (ground deposition) occurring at four distances from the tree bole (7.6, 15.2, 22.9, and 38.1 m) during conventional spray applications for protecting individual lodgepole pine (Pinus contorta Dougl. ex Loud.) from mountain pine beetle (Dendroctonus ponderosae Hopkins) attack and Engelmann spruce (Picea engelmannii Parry ex Engelm.) from spruce beetle (D. rufipennis [Kirby]) attack. Mean deposition (carbaryl + alpha-naphthol) did not differ significantly among treatments (nozzle orifices) at any distance from the tree bole. Values ranged from 0.04 +/- 0.02 mg carbaryl m(-2) at 38.1 m to 13.30 +/- 2.54 mg carbaryl m(-2) at 7.6 m. Overall, distance from the tree bole significantly affected the amount of deposition. Deposition was greatest 7.6 m from the tree bole and quickly declined as distance from the tree bole increased. Approximately 97% of total spray deposition occurred within 15.2 m of the tree bole. Application efficiency (i.e., percentage of insecticide applied that is retained on trees) ranged from 80.9 to 87.2%. Based on review of the literature, this amount of drift poses little threat to adjacent aquatic environments. No-spray buffers of 7.6 m should be sufficient to protect freshwater fish, amphibians, crustaceans, bivalves, and most aquatic insects. Buffers >22.9 m appear sufficient to protect the most sensitive aquatic insects (Plecoptera).

Surface temperature change of spruce forest as a result of bark beetle attack: remote sensing and GIS approach

In the 1990s, a bark beetle (Ips typographus [L.]) infection caused the decay of spruce forest (Picea abies [L.] Karst.) in the central part of the Sumava Mountains, the Czech Republic, bordering the Bavarian Forest National Park, Germany, where the bark beetle infection started in the late 1980s. Some areas were left without human intervention and, consequently, the trees around these areas were removed to stop further bark beetle outbreak. The objective of our study was the assessment of surface temperature (ST) change in spruce forest decayed under bark beetle and following clear-cutting. The change detection of ST is based on the comparison of modelled values and thermal satellite data. For this purpose, Landsat scenes from July 11th, 1987 and July 28th, 2002 were used. The models describe the dependence of ST of living spruce forest on topography. The topography effect is based on the Altitude and Hillshade index, which expresses the influence of Aspect and Slope on the relief illumination. Then the modelled ST values were extrapolated for decayed spruce forest and clear-cut areas. In order to increase model accuracy, the forest edge zones (90 m wide) were removed because of their different energy balance; then explained variability value (R2) increased from 0.37 to 0.55. The results of comparing modelled values with satellite ST in the decayed spruce forest and clear-cut areas show an average increase of ST by 5.2 and 3.5°C, respectively. The thermal satellite data from 1987 were used for model validation. This showed that the accuracy of ST modelling using topography was sufficient, because the difference between the modelled ST with and without decayed spruce forest and clear-cut areas was at most only 0.4°C.

Change in the forested and developed landscape of the Lake Tahoe basin, California and Nevada, USA, 1940–2002

The current ecological state of the Lake Tahoe basin has been shaped by significant landscape-altering human activity and management practices since the mid-1850s; first through widespread timber harvesting from the 1850s to 1920s followed by urban development from the 1950s to the present. Consequences of landscape change, both from development and forest management practices including fire suppression, have prompted rising levels of concern for the ecological integrity of the region. The impacts from these activities include decreased water quality, degraded biotic communities, and increased fire hazard. To establish an understanding of the Lake Tahoe basin's landscape change in the context of forest management and development we mapped, quantified, and described the spatial and temporal distribution and variability of historical changes in land use and land cover in the southern Lake Tahoe basin (279 km 2) from 1940 to 2002. Our assessment relied on post-classification change detection of multi-temporal land-use/cover and impervious-surface-area data that were derived through manual interpretation, image processing, and GIS data integration for four dates of imagery: 1940, 1969, 1987, and 2002. The most significant land conversion during the 62-year study period was an increase in developed lands with a corresponding decrease in forests, wetlands, and shrublands. Forest stand densities increased throughout the 62-year study period, and modern thinning efforts resulted in localized stand density decreases in the latter part of the study period. Additionally forests were gained from succession, and towards the end of the study period extensive tree mortality occurred. The highest rates of change occurred between 1940 and 1969, corresponding with dramatic development, then rates declined through 2002 for all observed landscape changes except forest density decrease and tree mortality. Causes of landscape change included regional population growth, tourism demands, timber harvest for local use, fire suppression, bark beetle attack, and fuels reduction activities. Results from this study offer land managers within the Lake Tahoe basin and in similar regions a basis for making better informed land-use and management decisions to potentially minimize detrimental ecological impacts of landscape change. The perspective to be gained is based on quantitative retrospection of the effects of human-driven changes and the impacts of management action or inaction to the forested landscape.

Forest management treatments, tree resistance, and bark beetle resource utilization in ponderosa pine forests of northern Arizona

We show experimentally that forest restoration treatments that used thinning to reduce ponderosa pine density near Flagstaff, Arizona decreased the percentage of trees with successful bark beetle attacks induced by beetle community response to a Dendroctonus brevicomis pheromone. Underlying mechanisms for the beneficial effect of restoration treatments on tree resistance to bark beetles included stimulation of resin defenses. Resin flow after phloem wounding was greater in full and partial restoration treatments than the control in the first and second year after treatment. Five bark beetle species, Ips pini, D. brevicomis, D. valens, D. adjunctus, and D. frontalis, colonized and produced brood in the trees. There was a negative correlation between resin volume and number of brood that emerged from the bole. Niche breadth of the bark beetle species over bole positions was widest for I. pini and narrowest for D. brevicomis.

Disturbances on a Wooded Raised Bog—How Windthrow, Bark Beetle and Fire Affect Vegetation and Soil Water Quality?

Pinus rotundata dominated peatbog (Žofinka Nature Reserve) in the Třeboň Basin, Czech Republic, was affected by “natural” disturbances: wind damage (1984), followed by a bark beetle attack, and fire (1994, 2000). Phytosociological releves were used to document vegetation. Soil water chemistry was compared in three differently affected stands: (1) an undisturbed Pinus rotundata bog forest, (2) a windthrow – bark beetle affected stand and (3) a site burned by wildfire in 2000. The species composition of the windthrow – bark beetle affected sites and the undisturbed P. rotundata bog forest differed mainly in the shrub and tree layers. Burned sites were partly colonized by anemochorous species (e.g. Taraxacum sp. div.) that disappeared within two or three years after colonization. Bare peat was colonized by bryophytes (e.g. Marchantia polymorpha and Funaria hygrometrica) typical of the disturbed sites, and by Polytrichum sp. div. and Aulacomnium palustre. Most plant species characteristic of the P. rotundata bog forest occurred at the burned sites eight years after the fire, but in different abundances. The edificator of the former community—P. rotundata—was mostly absent. Compared with windthrow followed by the bark beetle attack, fire promoted rapid expansion of Molinia caerulea. Soil water in both the undisturbed P. rotundata bog forest and the windthrow – bark beetle affected sites had a similar composition: very low pH values, high P concentrations, low concentrations of cations (Ca2+, Mg2+and K+) and inorganic nitrogen. The concentrations of soluble reactive phosphorus (SRP) and \({\text{NH}}_4^ + - {\text{N}}\) were negatively correlated with the groundwater table. Total P, SRP and \({\text{NH}}_4^ + - {\text{N}}\) concentrations in the soil water at the burned site were by one order of magnitude higher than those in the P. rotundata bog forest, while concentrations of K+, Mg2+ and Ca2+ were only about two times higher. High concentrations of P and N in the soil water found three years after the fire indicated a long-term elevated nutrient content in the soil water.

Nonhost Angiosperm Volatiles and Verbenone Protect Individual Ponderosa Pines from Attack by Western Pine Beetle and Red Turpentine Beetle (Coleoptera: Curculionidae, Scolytinae)

Abstract Nonhost angiosperm volatiles (NAV) and verbenone were tested for their ability to protect individual ponderosa pines, Pinus ponderosa Dougl. ex. Laws., from attack by western pine beetle (WPB), Dendroctonus brevicomis LeConte, and red turpentine beetle (RTB), Dendroctonus valens LeConte (Coleoptera: Curculionidae, Scolytinae). A combination of (−)-verbenone and eight NAVs [benzyl alcohol, benzaldehyde, guaiacol, nonanal, salicylaldehyde, (E)-2-hexenal, (E)-2-hexen-1-ol, and (Z)-2-hexen-1-ol] (NAVV) significantly reduced the density of WPB attacks and WPB successful attacks on attractant-baited trees. A significantly higher percentage of pitchouts (unsuccessful WPB attacks) occurred on NAVV-treated trees during two of three sample dates. In addition, significantly fewer RTB attacks were observed on NAVV-treated trees during all sampling dates. The application of NAVV to individual ponderosa pines significantly reduced tree mortality, with only 4 of 30 attractant-baited trees dying from bark beetle attack while 50% mortality (15/30) was observed in the untreated, baited control. To our knowledge, this is the first report establishing the effectiveness of NAVs and verbenone for protecting individual ponderosa pines from WPB attack.

Methyl jasmonate and oxalic acid treatment of Norway spruce: anatomically based defense responses and increased resistance against fungal infection

To study the effect of chemical pretreatment on conifer resistance, 13-year-old Norway spruce (Picea abies (L.) Karst.) trees were treated with methyl jasmonate (MJ) or oxalic acid (OxA) on the outer bark and inoculated with the pathogenic blue-stain fungus Ceratocystis polonica (Siem.) C. Moreau 4 weeks later. Both chemicals significantly reduced symptoms of fungal infection, but MJ was more effective than OxA (51 versus 18% reduction in length of necrotic lesions in the phloem relative to untreated control trees). Anatomical examination of treated stem tissues showed that MJ induced extensive formation of traumatic resin ducts in the xylem and extra polyphenolic parenchyma (PP) cells in the secondary phloem between the cambium and the regular annual PP cell layer. No traumatic resin ducts were formed after treatment with OxA, and the coverage of extra PP cells in OxA-treated tissues was not significantly higher than in the controls. The anatomically based defense reactions induced by MJ were similar to the reactions observed after pathogen infection, mechanical wounding and bark beetle attack. Neither MJ nor OxA had apparent phytotoxic effects on Norway spruce at the concentrations used, with needle and stem tissues of all trees appearing normal without visible symptoms of toxicity. However, trees treated with MJ had 30% less radial sapwood growth than control trees. In conclusion, MJ treatment of Norway spruce appears to have practical potential as a tool for increasing plant resistance to fungal infection, but with a modest reduction in sapwood growth.

FOREST HEALTH IMPACTS OF BARK BEETLES, DWARF MISTLETOE, AND BLISTER RUST IN A LAKE TAHOE BASIN MIXED CONIFER STAND

Interactions between forest health variables and mensurational characteristics in an uneven-aged eastern Sierra Nevada mixed conifer stand were examined. The stand was located in the Lake Tahoe Basin on a site featuring a coarsely textured granitic soil and numerous rock outcrops. Its composition was dominated by California white fir (Abies concolor var. lowiana [Gord.] Lemm.), with Jeffrey pine (Pinus jeffreyi Grev. & Balf.) and sugar pine (Pinus lambertiana Dougl.) less prominent and incense-cedar (Libocedrus decurrens Torr.) and mountain alder (Alnus tenuifolia Nutt.) the most minor constituents. The majority of saplings and seedlings were white fir. The stand exhibited no evidence that its development had been influenced by fire and, overall, it consisted of numerous small trees accruing little radial growth. Nearly one-quarter of all standing stems pole size or larger were dead, with mortality concentrated in white fir. Forest-floor fuel accumulations were excessive, and coarse debris was especially prominent. A fir engraver beetle (Scolytus ventralis LeConte) epidemic in white fir contrasted against apparent endemic population levels of the Jeffrey pine (Dendroctonus jeffreyi Hopkins) and red turpentine (Dendroctonus valens LeConte) beetles in Jeffrey pine and of the mountain pine beetle (Dendroctonus ponderosae Hopkins) in sugar pine. The severity of fir engraver attack on white fir was weakly related to overall tree size and to the proportion of composition consisting of this host species, while in Jeffrey pine and sugar pine, bark beetle attacks were strongly correlated with the individual proportions of these 2 hosts. Across all species, basal area explained a substantial proportion of the variation in overall attack severity. We found light infestations of true fir dwarf mistletoe (Arceuthobium abietinum Engelm. ex Munz f. sp. concoloris) in white fir and western dwarf mistletoe (Arceuthobium campylopodum Engelm.) in Jeffrey pine, plus an early stage of infection by the white pine blister rust (Cronartium ribicola J.C. Fischer) in sugar pine. Collectively, this case study characterized and quantified many of the conditions, symptoms, and causative agents inherent in a decadent mixed conifer stand in the eastern Sierra Nevada.

Retention or salvage logging of standing trees killed by the spruce bark beetle Ips typographus: Consequences for dead wood dynamics and biodiversity

Standing Norway spruce (Picea abies) trees attacked by the spruce bark beetle are often logged to reduce the risk of future bark beetle attacks. This study focuses on the potential ecological value of such trees. The study was conducted in six reserves in Sweden that were hit by a storm in 1995 and suffered from tree mortality caused by the spruce bark beetle, Ips typographus (L.) (Col.; Curculionidae). The volumes of storm-felled trees and of standing trees killed by the spruce bark beetle were recorded during 5 years after the storm. In the present study five of the six reserves were revisited in 2006. Of the dead standing wood, 81% was created by the spruce bark beetle, while 19% constituted of stumps from trees broken by the storm. Thirty-seven per cent of the beetle-killed trees remained standing in 2006, 8–9 years after tree death. A significantly higher frequency of fruiting bodies of Fomitopsis pinicola on broken snags compared with on intact snags indicates that the rot caused by this fungus contributed to tree fall. Almost 30% of the bark remained on both intact and fallen snags. The ecological value of snags created by the spruce bark beetle is discussed.

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.

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.

Effects of different ecological and silvicultural factors on beetle catches in the Turkish fir (Abies bornmülleriana Mattf.) ecosystems

To study the bark beetle attacks on Turkish fir stands and to determine the principle site and stand factors influencing beetle attacks, trap log method was used in the Western Black Sea Region of Turkey. The study was carried out in ten different locations prior to flying time in February 2003. On each experimental plot, three healthy fir trees with 30–50 cm diameter and 200 m apart were cut down with a chainsaw. From each sampling tree, needles were sub-sampled for determination of specific leaf-area and macro-nutrient concentrations. In addition, on each experimental plot, crown closure, basal-area, and stand density were measured. To determine the bulk density and nutrient concentrations of the soil, samples were taken around trees on each plot. The relationship between measured stand variables, the number of beetle species and beetle populations were evaluated using correlation and stepwise multiple regression procedure. Eight different species belonging to four different families from Coleoptera were identified in examining the trap logs. Six of these eight species (Pissodes piceae Illig., Pissodes notatus Fabr., Cryphalus piceae Ratz., Pityophthorus pityographus Ratz., Xyloterus lineatus Oliv., Pityokteines curvidens Germ.) were identified as harmful beetles (HB). The other two (Thanasimus formicarius L. and Rhizophagus dispar Payk.) were identified as predator species. The analysis showed that the total number of HB increased as the site’s slope and needle specific leaf area increased. However, as needle N concentrations increased, HB population decreased. Increased Ca concentration in fir needle reduced total insect attack.

Indirect interactions among dendrophages: Porcupines predispose pinyon pines to bark beetle attack

Discerning linkages among trophic levels and untangling indirect interactions is essential to understanding structuring of communities and ecosystems. Indeed, indirect interactions among disparate taxa are often essential to the functional role of these species. The goal of this research was to test the hypothesis that the relationship between 2 dendrophagous taxa, the North American porcupine ( Erethizon dorsatum) and the pine engraver beetle ( Ips hoppingi), is an asymmetric indirect interaction mediated by a common host. We proposed that damage by the porcupine predisposes the papershell pinyon pine ( Pinus remota) to colonization by pine engraver beetles. We examined porcupine tree selection, pinyon pine physiology and physiognomy, and beetle-pine associations on a study area in the southwestern Edwards Plateau of Texas from June 1997 to August 1999. Although attacks by beetles were evident on both damaged and undamaged trees, successful colonization was greater on pines damaged by porcupines. Intensity of porcupine attack, indexed by number of feeding scars and area of bark removed, also was associated with subsequent colonization by beetles. Porcupines selected pinyon pines over more abundant species ( P < 0.001) and were selective at the level of morphology, whereas pine engraver beetles were selective of tree morphology and physiology. Trees colonized by beetles had phloem with higher concentrations of fructose and glucose and lower percent composition of limonene, sabinene, and terpinolene than uncolonized trees. Our findings supported our hypothesis of an indirect interaction between these dendrophages. We rejected alternative explanations (e.g., that these dendrophages preferred similar trees or that beetles facilitated porcupine damage) for this relationship based on the biology of Ips and their selection of host trees. We propose that release of volatile terpenes as a result of porcupine feeding and reallocation of carbon resources as a response to stress explains the facilitation of beetle colonization in porcupine-damaged trees. Our findings parallel those observed in other systems involving indirect effects and fit within the framework of theories explaining host plant-herbivore interactions.

Air Pollution Distribution Patterns in the San Bernardino Mountains of Southern California: a 40-Year Perspective

Since the mid-1950s, native pines in the San Bernardino Mountains (SBM) in southern California have shown symptoms of decline. Initial studies in 1963 showed that ozone (O3) generated in the upwind Los Angeles Basin was responsible for the injury and decline of sensitive trees. Ambient O3 decreased significantly by the mid-1990s, resulting in decreased O3 injury and improved tree growth. Increased growth of trees may also be attributed to elevated atmospheric nitrogen (N) deposition. Since most of the N deposition to mixed conifer forest stands in the SBM results from dry deposition of nitric acid vapor (HNO3) and ammonia (NH3), characterization of spatial and temporal distribution of these two pollutants has become essential. Although maximum daytime O3 concentrations over last 40 years have significantly decreased (~3-fold), seasonal means have been reduced much less (~1.5-fold), with 2-week long means occasionally exceeding 100 ppb in the western part of the range. In the same area, significantly elevated concentrations of HNO3 and NH3, up to 17.5 and 18.5 μg/m3 as 2-week averages, respectively, have been determined. Elevated levels of O3 and increased N deposition together with long-term drought predispose the SBM forests to massive bark beetle attacks making them susceptible to catastrophic fires.