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.     

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.     

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.     

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.     

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

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

Bark beetle-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.     

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.     

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.     

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.     

PHENIPS—A comprehensive phenology model of Ips typographus (L.) (Col., Scolytinae) as a tool for hazard rating of bark beetle infestation

We developed the model PHENIPS for spatial and temporal simulation of the seasonal development of Ips typographus at the Kalkalpen National Park in Austria. The model is based on a digital elevation model used for interpolation of temperature and solar radiation to calculate the microclimatic conditions (bark temperature) for the beetles’ development. Additionally, the beetles’ phenology at Kalkalpen National Park was monitored along with air and bark temperature measurements. The onset of host tree infestation in spring was estimated using a lower threshold of 16.5 °C for flight activity and a mean thermal sum of 140 degree-days (dd) from beginning of April 1st onward. Rate of brood development was calculated from accumulated degree-days of hourly temperature data using upper and lower temperature thresholds of 38.9 and 8.3 °C, respectively, and a nonlinear function for calculating effective thermal sums. Re-emergence of parental beetles occurred at a time when 49.7% of the thermal sum for total development (557 dd) was reached. The model includes the discontinuance of the beetle's reproductive activity at a day length <14.5 h. The rate of successful hibernation of established broods is predicted by assessing the developmental stage of initiated generations at the beginning of the cold period. For validation we compared the timing of phenological events in the field with predicted events using both, hourly recorded data at trap trees in the terrain and generated daily topoclimatic data. Using topoclimatic data, the onset of infestation was predicted with a mean absolute error of 1.3 days. The observed onset of emergence of filial beetles in the field was estimated with a mean error of 39 dd. Our PHENIPS explicitly considers the strong effects of regional topography and stand conditions on local air and bark temperature and can be used for precise monitoring of the actual state of bark beetle development at the specific stand/tree level. Using topoclimatic data, PHENIPS simulates the maximum number of generations which is necessary to assess the potential impact of bark beetle outbreaks at regional scale. Further applications of PHENIPS for site-specific hazard rating of bark beetle infestation are discussed.     

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.     

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.     

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.     

Optimizing the joint production of timber and bilberries

Berries and mushrooms are increasingly appreciated products of Finnish forests. Therefore, there is a need to integrate them in silvicultural planning. Bilberry (Vaccinium myrtillus L.) is an economically important wild berry that is widely collected for household consumption and sale in North Karelia, Finland. In this study, bilberry yield models developed recently were included in a stand growth simulator and the joint production of timber and bilberry was optimized by maximizing soil expectation value (SEV) with 3% discounting rate, assuming that 75% of the bilberry yield is harvested. The effect of bilberry production on the optimal stand management increased with increasing bilberry price. With high bilberry prices (4–8 € kg⁻¹) it was optimal to manage the mixed stand of Scots pine, Norway spruce and birch, and the pure stand of Norway spruce so as to promote bilberry production. In the Scots pine stand, where bilberry yields are higher, bilberry production affected optimal stand management already with a price of 2 € kg⁻¹. Compared to timber production, joint production led to longer rotation lengths, higher thinning intensities, more frequent thinnings, and higher share of Scots pine in the mixed stand. The contribution of bilberries to the total SEV increased with increasing bilberry price and discounting rate. In the mixed stand and pine stand the SEV of bilberry production, calculated with 3% discounting rate, exceeded the SEV of timber production when bilberry price was 4 € kg⁻¹.With 4% discounting rate this happened already with bilberry price of 2 € kg⁻¹. It was concluded that forest management which promotes bilberry yields is the most profitable in pine stands where the potential bilberry yields are high.     

Spatial variability in stand structure and density-dependent mortality in newly established post-fire stands of a California closed-cone pine forest

Fire is an important process in California closed-cone pine forests; however spatial variability in post-fire stand dynamics of these forests is poorly understood. The 1995 Vision Fire in Point Reyes National Seashore burned over 5000 ha, initiating vigorous Pinus muricata (bishop pine) regeneration in areas that were forested prior to the fire but also serving as a catalyst for forest expansion into other locales. We examined the post-fire stand structure of P. muricata forest 14 years after fire in newly established stands where the forest has expanded across the burn landscape to determine the important factors driving variability in density, basal area, tree size, and mortality. Additionally, we estimated the self-thinning line at this point in stand development and compared the size-density relationship in this forest to the theorized (−1.605) log-log slope of Reineke’s Rule, which relates maximum stand density to average tree size. Following the fire, post-fire P. muricata density in the expanded forest ranged from 500 to 8900 live stems ha⁻¹ (median density = 1800 ha⁻¹). Post-fire tree density and basal area declined with increasing distance to individual pre-fire trees, but showed little variation with other environmental covariates. Self-thinning (density-dependent mortality) was observed in nearly all stands with post-fire density >1800 stems ha⁻¹, and post-fire P. muricata stands conformed to the size-density relationship predicted by Reineke’s Rule. This study demonstrates broad spatial variability in forest development following stand-replacing fires in California closed-cone pine forests, and highlights the importance of isolated pre-fire trees as drivers of stand establishment and development in serotinous conifers.     

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

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

Risk rating for mountain pine beetle infestation of lodgepole pine forests over large areas with ordinal regression modelling

The mountain pine beetle Dendroctonus ponderosae Hopkins is endemic to lodgepole pine, Pinus contorta var. latifolia Engelmann, forests in western Canada. However, the current beetle epidemic in this area highlights the challenges faced by forest managers tasked with prioritizing stands for mitigation activities such as salvage harvesting and direct control methods. In western Canada, the operational risk rating system for mountain pine beetle is based on biological knowledge gained from a rich legacy of stand-scale field studies. Due to the large spatial (millions of hectares affected) and temporal (over 10 years) extents of the current epidemic, new research into large-area mountain pine beetle processes has revealed further insights into the landscape-scale characteristics of beetle infested forests. In this paper, we evaluated the potential for this new knowledge to augment an established system for rating the short-term risk of tree mortality in a stand due to mountain pine beetle. New variables explored for utility in risk rating include direct shortwave radiation, site index, diameter at breast height, the temporal trends in local beetle populations, Biogeoclimatic Ecosystem Classification and beetle–host interaction variables. Proportional odds ordinal regression was used to develop a model for the Vanderhoof Forest District in west-central British Columbia. Prediction on independent data was assessed with the area under the receiver operator curve (AUC), indicating good discriminatory power (AUC = 0.84) for predicting levels of mountain pine beetle-caused pine mortality.     

秦岭华山松小蠹生态位研究

通过对秦岭林区海拔１６００～２２００ｍ的中山地带华山松小蠹虫种类和生态位的研究,结果表明：在秦岭林区入侵危害寄主华山松的小蠹虫有１９种,其中能构成竞争和共存的小蠹虫主要有１１种。虽然,这１１种小蠹虫具有各自不同的生态位宽度,且存在不同程度的生态位重叠,但各小蠹虫可依据其对寄主树木营养和空间需求的不同、种群密度的相互制约,以及入侵寄主树木时序的差异达到竞争的平衡和共存。秦岭华山松小蠹生态系统的建立,首先是由华山松大小蠹入侵健康华山松,并通过携带蓝变真菌入侵寄主树木,迅速克服寄主树木抗性系统,使寄主树木树势衰弱;其次松六齿小蠹、暗额星坑小蠹和松十二齿小蠹,作为秦岭华山松的主要次期性小蠹,迅速入侵衰弱的寄主华山松;最后其它次期性小蠹虫入侵寄主树木,利用寄主华山松剩余营养和空间。从而实现秦岭华山松立木小蠹生态系统的动态稳定。     

Tree regeneration and future stand development after bark beetle infestation and harvesting in Colorado lodgepole pine stands

In the southern Rocky Mountains, current mountain pine beetle (Dendroctonus ponderosae Hopkins) outbreaks and associated harvesting have set millions of hectares of lodgepole pine (Pinus contorta var. latifolia Engelm. ex Wats.) forest onto new stand development trajectories. Information about immediate, post-disturbance tree regeneration will provide insight on dynamics of future stand composition and structure. We compared tree regeneration in eight paired harvested and untreated lodgepole pine stands in the Fraser Experimental Forest that experienced more than 70% overstory mortality due to beetles. New seedlings colonized both harvested and untreated stands in the first years after the beetle outbreak. In harvested areas the density of new seedlings, predominantly lodgepole pine and aspen, was four times higher than in untreated stands. Annual height growth of pine and fir advance regeneration (e.g., trees established prior to the onset of the outbreak) has doubled following overstory mortality in untreated stands. Growth simulations based on our regeneration data suggest that stand basal area and stem density will return to pre-beetle levels in untreated and harvested stands within 80-105 years. Furthermore, lodgepole pine will remain the dominant species in harvested stands over the next century, but subalpine fir will become the most abundant species in untreated areas. Owing to terrain, economic and administrative limitations, active management will treat a small fraction (<15%) of the forests killed by pine beetle. Our findings suggest that the long-term consequences of the outbreak will be most dramatic in untreated forests where the shift in tree species composition will influence timber and water production, wildfire behavior, wildlife habitat and other forest attributes.