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.     

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

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

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

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

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.     

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.     

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.     

Physiological responses of ponderosa pine in western Montana to thinning, prescribed fire and burning season

Low-elevation ponderosa pine (Pinus ponderosa Dougl. ex. Laws.) forests of the northern Rocky Mountains historically experienced frequent low-intensity fires that maintained open uneven-aged stands. A century of fire exclusion has contributed to denser ponderosa pine forests with greater competition for resources, higher tree stress and greater risk of insect attack and stand-destroying fire. Active management intended to restore a semblance of the more sustainable historic stand structure and composition includes selective thinning and prescribed fire. However, little is known about the relative effects of these management practices on the physiological performance of ponderosa pine. We measured soil water and nitrogen availability, physiological performance and wood radial increment of second growth ponderosa pine trees at the Lick Creek Experimental Site in the Bitterroot National Forest, Montana, 8 and 9 years after the application of four treatments: thinning only; thinning followed by prescribed fire in the spring; thinning followed by prescribed fire in the fall; and untreated controls. Volumetric soil water content and resin capsule ammonium did not differ among treatments. Resin capsule nitrate in the control treatment was similar to that in all other treatments, although burned treatments had lower nitrate relative to the thinned-only treatment. Trees of similar size and canopy condition in the three thinned treatments (with and without fire) displayed higher leaf-area-based photosynthetic rate, stomatal conductance and mid-morning leaf water potential in June and July, and higher wood radial increment relative to trees in control units. Specific leaf area, mass-based leaf nitrogen content and carbon isotope discrimination did not vary among treatments. Our results suggest that, despite minimal differences in soil resource availability, trees in managed units where basal area was reduced had improved gas exchange and growth compared with trees in unmanaged units. Prescribed fire (either in the spring or in the fall) in addition to thinning, had no measurable effect on the mid-term physiological performance and wood growth of second growth ponderosa pine.     

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.     

Limited response of ponderosa pine bole defenses to wounding and fungi

Tree defense against bark beetles (Curculionidae: Scolytinae) and their associated fungi generally comprises some combination of constitutive (primary) and induced (secondary) defenses. In pines, the primary constitutive defense against bark beetles consists of preformed resin stored in resin ducts. Induced defenses at the wound site (point of beetle entry) in pines may consist of an increase in resin flow and necrotic lesion formation. The quantity and quality of both induced and constitutive defenses can vary by species and season. The inducible defense response in ponderosa pine is not well understood. Our study examined the inducible defense response in ponderosa pine using traumatic mechanical wounding, and wounding with and without fungal inoculations with two different bark beetle-associated fungi (Ophiostoma minus and Grosmannia clavigera). Resin flow did not significantly increase in response to any treatment. In addition, necrotic lesion formation on the bole after fungal inoculation was minimal. Stand thinning, which has been shown to increase water availability, had no, or inconsistent, effects on inducible tree defense. Our results suggest that ponderosa pine bole defense against bark beetles and their associated fungi is primarily constitutive and not induced.     

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.     

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.     

The effects of a thinning treatment on carbon stocks in a northern Arizona ponderosa pine forest

Vast areas of ponderosa pine (Pinus ponderosa Dougl. ex Laws.) forest in the western United States have become unnaturally dense because of relatively recent land management practices that include fire suppression and livestock grazing. In many areas, thinning treatments can re-establish the natural ecological processes and help restore ecosystem structure and function. Precipitous global climate change has focused attention on the carbon storage in forests. An unintended consequence of fire suppression has been the increased storage of carbon in ponderosa stands. Thinning treatments reduce standing carbon stocks while releasing carbon through the combustion of fuel in logging machinery, burning slash, and the decay of logging slash and wood products. These reductions and releases of stored carbon must be compared to the risk of catastrophic fire burning through the stand and releasing large quantities of carbon to the atmosphere to more fully understand the costs and benefits – in carbon terms – of forest restoration strategies.     

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

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

Bark beetle community structure under four ponderosa pine forest stand conditions in northern Arizona

We studied the bark beetle guild (Coleoptera: Scolytidae) in the ponderosa pine forests of northern Arizona to explore if the species assemblages and relative abundance differ between managed and unmanaged stands. Four stand conditions were assessed: (1) unmanaged stands with high tree density, (2) thinned stands, (3) thinned and burned (with prescribed fire) stands and (4) stands that had been burned by stand replacing wildfires. The study was conducted in the ponderosa pine forests of the Coconino Plateau, northern Arizona. For several decades this area has been relatively free of bark beetle outbreaks despite the current overstocked condition of many stands. We found that a similar species assemblage composed of Dendroctonus frontalis, D. brevicomis, D. valens, D. approximatus, D. ponderosae, and Ips pini occurred across all four stand conditions over 3 years of study. The population levels of all these species were endemic across all stand conditions. The non-aggressive D. approximatus and D. valens were indicator species for thinned and unmanaged stands, respectively, but this was not consistent among years. The ambrosia beetle Gnathotrichus sp. and the bark beetle predator Enoclerus sp. consistently indicated stands burned by wildfire. In addition to our field experiment, we analyzed the historical pattern of attacks of bark beetles in our area of study. Our findings suggest that the pattern of attack of D. brevicomis (the only Dendroctonus species for which attacks have been reported) and Ips spp. has been through scattered small infestations in groups of 1–10 trees. Whereas small infestations by Ips spp. are increasing, those for D. brevicomis are decreasing. Although we agree that the high density stands in northern Arizona are in an “unhealthy” condition, our results do not show that they were supporting large bark beetle outbreaks. Our results challenge the theoretical assumptions about the relationship between stand structure, tree resistance and bark beetle performance.     

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.     

Bark beetles,fuels, fires and implications for forest management in the Intermountain West

Bark beetle-caused tree mortality in conifer forests affects the quantity and quality of forest fuels and has long been assumed to increase fire hazard and potential fire behavior. In reality, bark beetles, and their effects on fuel accumulation, and subsequent fire hazard, are poorly understood. We extensively sampled fuels in three bark beetle-affected Intermountain conifer forests and compared these data to existing research on bark beetle/fuels/fire interactions within the context of the disturbance regime. Data were collected in endemic, epidemic and post-epidemic stands of Douglas-fir, lodgepole pine and Engelmann spruce. From these data, we evaluated the influence of bark beetle-caused tree mortality on various fuels characteristics over the course of a bark beetle rotation. The data showed that changes in fuels over time create periods where the potential for high intensity and/or severe fires increases or decreases. The net result of bark beetle epidemics was a substantial change in species composition and a highly altered fuels complex. Early in epidemics there is a net increase in the amount of fine surface fuels when compared to endemic stands. In post-epidemic stands large, dead, woody fuels, and live surface fuels dominate. We then discuss potential fire behavior in bark beetle-affected conifer fuels based on actual and simulated fuels data. Results indicated that for surface fires both rates of fire spread and fireline intensities were higher in the current epidemic stands than in the endemic stands. Rates of spread and fireline intensities were higher in epidemic stands due, however, to decreased vegetative sheltering and its effect on mid-flame wind speed, rather than changes in fuels. Passive crown fires were more likely in post-epidemic stands, but active crown fires were less likely due to decreased aerial fuel continuity. We also discuss the ecological effects of extreme fire behavior. Information is presented on managing forests to reduce the impact of bark beetle outbreaks and the interplay between management, bark beetle populations, fuels and fire hazard and behavior.     

Structure and composition changes following restoration treatments of longleaf pine forests on the Gulf Coastal Plain of Alabama

Longleaf pine (Pinus palustris Mill.) forests of the Gulf Coastal Plain historically burned every 2–4 years with low intensity fires, which maintained open stands with herbaceous dominated understories. During the early and mid 20th century however, reduced fire frequency allowed fuel to accumulate and hardwoods to increase in the midstory and overstory layers, while woody shrubs gained understory dominance. In 2001, a research study was installed in southern Alabama to develop management options that could be used to reduce fuel loads and restore the ecosystem. As part of a nationwide fire and fire surrogates study, treatments included a control (no fire or other disturbance), prescribed burning only, thinning of selected trees, thinning plus prescribed burning, and herbicide plus prescribed burning. After two cycles of prescribed burning, applied biennially during the growing season, there were positive changes in ecosystem composition. Although thinning treatments produced revenue, while reducing midstory hardwoods and encouraging growth of a grassy understory, burning was needed to discourage regrowth of the hardwood midstory and woody understory. Herbicide application followed by burning gave the quickest changes in understory composition, but repeated applications of fire eventually produced the same results at the end of this 8-year study. Burning was found to be a critical component of any restoration treatment for longleaf communities of this region with positive changes in overstory, midstory and understory layers after just three or four burns applied every 2 or 3 years.     

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.     

Postsettlement Changes in Natural Fire Regimes and Forest Structure

Heavy livestock grazing, logging, and fire exclusion associated with Euro-American settlement has brought about substantial changes in forest conditions in western forests. Thus, old-growth definitions based on current forest conditions may not be compatible with the natural conditions prevalent throughout the evolutionary history of western forest types. Detailed analysis of data from two study areas in the southwestern ponderosa pine type suggests that average tree densities have increased from as few as 23 trees per acre in presettlement times to as many as 851 trees per acre today. Associated with these increases in tree density are increases in canopy closure, vertical fuel continuity, and surface fuel loadings resulting in fire hazards over large areas never reached before settlement. In addition, fire exclusion and increased tree density has likely decreased tree vigor (increasing mortality from disease, insect, drought, etc.), herbaceous and shrub production, aesthetic values, water availability and runoff, and nutrient availability, and also changed soil characteristics and altered wildlife habitat. To remedy these problems and restore these forest ecosystems to more nearly natural conditions, and maintain a viable cohort of old age-class trees, it will be necessary to thin out most of the postsettlement trees, manually remove heavy fuels from the base of large, old trees, and reintroduce periodic burning.     

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.