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.     

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.     

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.     

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.     

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.     

Bark Beetle Demography in a Jeffrey Pine Stand as Influenced by Mechanized Thinning and Prescribed Fire

Forest thinnings implemented with cut-to-length and whole-tree harvesting systems followed by underburning were evaluated for their effects on bark beetle prevalence in pure, uneven-aged Jeffrey pine (Pinus jeffreyi Grev. & Balf.) interspersed with isolated California white fir (Abies concolor var. lowiana [Gord.] Lemm.). Based on pitch tube counts in a stand with a moderate bark beetle population in its pine component, the Jeffrey pine beetle (Dendroctonus jeffreyi Hopkins) generally preferred larger trees before treatment implementation, but after exhibiting mixed pretreatment tendencies concerning stand density demonstrated a posttreatment proclivity toward higher density. Cut-to-length thinning followed by underburning increased the pine beetle population while whole-tree thinning unaccompanied by burning reduced it. Tree mortality was induced by the bark beetle infestation but was not its sole cause. Pitch tube abundance on white fir far exceeded that on Jeffrey pine, and the greatest influence on the fir engraver (Scolytus ventralis LeConte) population was the prevalence of its host tree. The responses presented herein to these thinning and burning practices, which are being increasingly utilized in forest restoration efforts in the western USA, provide natural resource managers insight into potential forest health outcomes when implemented in Jeffrey pine and similar dry site forest types.     

Perpetuating old ponderosa pine

We review current knowledge about the use of management treatments to reduce human-induced threats to old ponderosa pine (Pinus ponderosa) trees. We address the following questions: Are fire-induced damage and mortality greater in old than younger trees? Can management treatments ameliorate the detrimental effects of fire, competition-induced stress, and drought on old trees? Can management increase resistance of old trees to bark beetles? We offer the following recommendations for the use of thinning and burning treatments in old-growth ponderosa pine forests. Treatments should be focused on high-value stands where fire exclusion has increased fuels and competition and where detrimental effects of disturbance during harvesting can be minimized. Fuels should be reduced in the vicinity of old trees prior to prescribed burns to reduce fire intensity, as old trees are often more prone to dying after burning than younger trees. Raking the forest floor beneath old trees prior to burning may not only reduce damage from smoldering combustion under certain conditions but also increase fine-root mortality. Thinning of neighboring trees often increases water and carbon uptake of old trees within 1 year of treatment, and increases radial growth within several years to two decades after treatment. However, stimulation of growth of old trees by thinning can be negated by severe drought. Evidence from young trees suggests that management treatments that cause large increases in carbon allocation to radial xylem growth also increase carbon allocation to constitutive resin defenses against bark beetle attacks, but evidence for old trees is scarce. Prescribed, low-intensity burning may attract bark beetles and increase mortality of old trees from beetle attacks despite a stimulation of bole resin production.     

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.     

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.     

Predicting mortality for five California conifers following wildfire

Fire injury was characterized and survival monitored for 5677 trees >25 cm DBH from five wildfires in California that occurred between 2000 and 2004. Logistic regression models for predicting the probability of mortality 5-years after fire were developed for incense cedar (Calocedrus decurrens (Torr.) Florin), white fir (Abies concolor (Gord. & Glend.) Lindl. ex Hildebr.), sugar pine (Pinus lambertiana Douglas), Jeffrey pine (P. jeffreyi Balf.), and ponderosa pine (P. ponderosa C. Lawson). Differences in crown injury variables were also compared for Jeffrey and ponderosa pine. Most mortality (70–88% depending on species) occurred within 2 years post-wildfire and had stabilized by year 3. Crown length and crown volume injury variables predicted tree mortality equally well; however, the variables were not interchangeable. Crown injury and cambium kill rating was significant in predicting mortality in all models. DBH was only a significant predictor of mortality for white fir and the combined ponderosa and Jeffrey pine models developed from the McNally Fire; these models all predicted increasing mortality with increasing tree size. Red turpentine beetle (Dendroctonus valens) was a significant predictor variable for sugar pine, ponderosa pine, and Jeffrey pine; ambrosia beetle (Trypodendron and Gnathotrichus spp.) was a significant predictor variable for white fir. The mortality models and post-fire tree survival characteristics provide improved prediction of 5-year post-wildfire tree mortality for several California conifers. The models confirm the overall importance of crown injury in predicting post-fire mortality compared to other injury variables for all species. Additional variables such as cambium kill, bark beetles, and tree size improved model accuracies, but likely not enough to justify the added expense of data collection.     

Pinus pinaster Ait. tree mortality following wildfire in Spain

Maritime pine (Pinus pinaster Ait.) is the tree species most affected by wildfire in the Iberian Peninsula. Prediction of the probability of fire-injured tree mortality is critical for management of burned areas, evaluation of the ecological and economic impact of wildfire and prescribed fire planning and application. Pine bark beetles (Scolytidae) frequently attack burned maritime pine stands and cause extensive post-fire mortality throughout the Iberian Peninsula. In the present study, maritime pine trees were monitored for three years following 14 wildfires in four ecotypes in Spain (11 fires in Galicia (Galician ecotype - NW Spain), one fire in Portillo (Meseta-Castellana ecotype - Central Spain), one fire in Rodenal (Rodenal ecotype - Central Spain), and one fire in Genalguacil (Sierra Bermeja ecotype - SW Spain)). Data on tree attributes, crown and bole injury, ground fire severity, Ips sp. presence and tree survival were obtained by examining 3085 trees. Logistic regression models for predicting the probability of delayed maritime pine mortality were developed by use of generalized estimated equations (GEE). An ample range of response to fire damage in mortality was evident among the four ecotypes and different models were fitted for each. The most important variables for predicting tree mortality were total crown volume damaged, presence of Ips sp. attack and cambium kill rating. The results highlight the extensive presence of Ips sp. in burned maritime pine forests and its importance in tree mortality process, the ample range of response of P. pinaster, in terms of post-fire mortality, as well as the need to develop site specific mortality models for the different ecotypes of this species following fire.     

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

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

The role of germination microsite in the establishment of sugar pine and Jeffrey pine seedlings

Mature shrubs can provide microhabitats that are beneficial to tree seedling growth and development. Sugar pine trees (Pinus lambertiana) grow in a narrow zone on the eastern slope of the Carson Range in extreme western Nevada, whereas Jeffrey pine (Pinus jeffreyi) is the dominant tree species in the region, an area extensively disturbed by wild fire. This study compares seedling establishment of sugar pine and Jeffrey pine relative to mature shrubs. In the fall of 2002 (cohort 1) and 2003 (cohort 2), 13,600 seeds of both species were planted in wire mesh enclosures, at three sites, under a variety of microhabitat treatments: under shade and in the open, under two species of shrub cover, and with and without plant litter. Seedlings were monitored for survival through two growing seasons. Even though more sugar pine seedlings emerged, more Jeffrey pine seedlings survived, and Jeffrey pine was the more drought tolerant species, better suited for the xeric climate found in the Carson Range. Litter slightly hindered seedling emergence but had no effect on survival and there was no significant species × litter interaction. Supplemental water facilitated survival in all treatments with highest survival in shade treatments. Sugar pine seedlings showed a significant increase in survival over Jeffrey pine seedlings with the addition of water, particularly in open treatments and more of both species survived under manzanita shrubs with water. The highest seedling mortality occurred when shrub canopy was removed, and seedlings experienced the effect of full sun and competition for soil water. For either species, microhabitat is a significant factor in determining success or failure in rehabilitation efforts after disturbance.     

Predisposition to bark beetle attack by root herbivores and associated pathogens: Roles in forest decline,gap formation,and persistence of endemic bark beetle populations

Bark beetles are largely known for their ability to undergo intermittent population eruptions that transform entire landscapes and pose significant economic hardships. However, most species do not undergo outbreaks, and eruptive species usually exert only minor disturbances. Understanding the dynamics of tree-killing noneruptive species can provide insights into how beetles persist at low densities, and how some spatiotemporal patterns of host predisposition may more likely favor breaching eruptive thresholds than others. Elucidating mechanisms behind low-density populations is challenging, however, due to the requirement of long-term monitoring and high degrees of spatial and temporal covariance. We censused more than 2700 trees annually over 7 years, and at the end of 17 years, in a mature red pine plantation. Trees were measured for the presence of bark beetles and wood borers that breed within the primary stem, root weevils that breed in root collars, and bark beetles that breed in basal stems. We quantify the sequence of events that drive this decline syndrome, with the primary emergent pattern being an interaction between below- and above-ground herbivores and their fungal symbionts. This interaction results in an expanding forest gap, with subsequent colonization by early-successional vegetation. Spatial position strongly affects the likelihood of tree mortality. A red pine is initially very likely to avoid attack by tree-killing Ips beetles, but attack becomes increasingly likely as the belowground complex spreads to neighboring trees and eventually make trees susceptible. This system is largely internally driven, as there are strong gap edge, but not stand-edge, effects. Additional stressors, such as drought, can provide an intermittent source of susceptible trees to Ips beetles, and elevated temperature slightly accentuates this effect. New gaps can arise from such trees as they subsequently become epicenters for the full complex of organisms associated with this decline, but this is not common. As Ips populations rise, there is some element of positive feedback, in that the proportion of killed trees that were not first colonized by root organisms increases. This positive feedback is very weak, however, and we propose the slope between beetle population density and reliance on host stress as a quantitative distinction along a gradient from noneruptive through eruptive species. Almost all trees colonized by Ips were subsequently colonized by wood borers, likely a source of negative feedback. We discuss implications to our overall understanding of cross-scale interactions, between-guild interactions, forest declines, and eruptive thresholds.     

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.     

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.     

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.     

Effects of defoliation by Eucalyptus weevil, Gonipterus scutellatus, and chrysomelid beetles on growth of Eucalyptus globulus in southwestern Australia

Plantations of Eucalyptus globulus in southwestern Australia are defoliated by Eucalyptus weevil, Gonipterus scutellatus, and a complex of chrysomelid and scarab beetles, yet there is no information on the impact of beetle defoliation to tree growth in southwestern Australia. To address this shortcoming, we used insect exclusion trials, to compare growth of insecticide treated (and thus relatively undamaged) trees with untreated (and thus defoliated) trees to determine whether defoliation by G. scutellatus and other beetles reduced the growth and harvest volume of E. globulus trees. Our results showed some evidence of beetle defoliation reducing growth of E. globulus. Mean defoliation levels of the growing tip of untreated trees ranged from 18% to 33% across the duration of the study and were significantly greater than mean defoliation levels of 5–16% on insecticide treated trees. Seasonal peaks in defoliation of 30–80% to the growing tip of untreated trees were recorded between late spring and early autumn. The greatest impact of defoliation on tree growth was evident during the 2.5 year period of insect exclusion, when higher relative growth rates were recorded for insecticide treated trees, which were significantly different from relative growth rates of untreated trees at two of the four plantations. However, our results showed only a limited impact of beetle defoliation on the total volume at harvest. Initially small trees tended to suffer more severe defoliation than initially large trees. Effects of insect exclusion treatment on harvest volume were modified by the initial tree size and the relationship between the initial tree size and levels of defoliation.     

Can prescribed fire be used to maintain fuel treatment effectiveness over time in Black Hills ponderosa pine forests?

We determine the time frame after initial fuel treatment when prescribed fire will be likely to produce high enough mortality rates in ponderosa pine (Pinus ponderosa var. scopulorum Dougl. ex Laws.) regeneration to be successful in maintaining treatment effectiveness in the Black Hills of South Dakota. We measured pine regeneration in disturbed stands and young pine growth rates to estimate the susceptibility of pine regeneration to prescribed fire with time since initial treatment. We also determined surface fuel accumulation rates for stands after prescribed fire to help estimate likely fire behavior in maintenance prescribed fire. Given our estimates of regeneration density and tree size, and likely fire behavior, we then used small pine tree mortality—fire effect relations to estimate the effects of prescribed fire on developing understory pine at specific times since initial treatments.     

Afforestation method affects the isotopic composition of planted Pinus halepensis in a semiarid region of Spain

We used an isotopic approach to evaluate the effects of three afforestation methods on the ecophysiology of an Aleppo pine plantation in semiarid Spain. The site preparation methods tested were excavation of planting holes (H), subsoiling (S), and subsoiling with addition of urban solid refuse to soil (S + USR). Five years after plantation establishment, trees in the S + USR treatment were over three times larger than those in the S treatment, and nearly five-fold larger than those planted in holes. Differences in tree biomass per hectare were even greater due to disparities in initial planting density and pine tree mortality among treatments. Pine trees in the S + USR treatment showed higher foliar P concentration, δ¹³C and δ¹⁵N than those in the S or H treatments. Foliar δ¹⁵N data proved that trees in the S + USR treatment utilized USR as a source of nitrogen. Foliar δ¹³C and δ¹⁸O data suggest that improved nutrient status differentially stimulated photosynthesis over stomatal conductance in the pine trees of the S + USR treatment, thus enhancing water use efficiency and growth. In the spring of 2002, trees in the S + USR treatment exhibited the most negative predawn water potentials of all the treatments, indicating that the rapid early growth induced by USR accelerated the onset of intense intra-specific competition for water. The results of this study have implications for the establishment and management of Aleppo pine plantations on semiarid soils. Planting seedlings at low density and/or early thinning of pine stands are strongly recommended if fast tree growth is to be maintained beyond the first few years after USR addition to soil. Foliar C, O and N isotope measurements can provide much insight into how resource acquisition by trees is affected by afforestation techniques in pine plantations under dry climatic conditions.