Stand characteristics and downed woody debris accumulations associated with a mountain pine beetle (Dendroctonus ponderosae Hopkins) outbreak in Colorado

Lodgepole pine (Pinus contorta Dougl. ex Loud.)-dominated ecosystems in north-central Colorado are undergoing rapid and drastic changes associated with overstory tree mortality from a current mountain pine beetle (Dendroctonus ponderosae Hopkins) outbreak. To characterize stand characteristics and downed woody debris loads during the first 7 years of the outbreak, 221 plots (0.02 ha) were randomly established in infested and uninfested stands distributed across the Arapaho National Forest, Colorado. Mountain pine beetle initially attacked stands with higher lodgepole pine basal area, and lower density and basal area of Engelmann spruce (Picea engelmannii [Parry]), and subalpine fir (Abies lasiocarpa (Hook.) Nutt. var. lasiocarpa) compared to uninfested plots. Mountain pine beetle-affected stands had reduced total and lodgepole pine stocking and quadratic mean diameter. The density and basal area of live overstory lodgepole declined by 62% and 71% in infested plots, respectively. The mean diameter of live lodgepole pine was 53% lower than pre-outbreak in infested plots. Downed woody debris loads did not differ between uninfested plots and plots currently infested at the time of sampling to 3 or 4–7 years after initial infestation, but the projected downed coarse wood accumulations when 80% of the mountain pine beetle-killed trees fall indicated a fourfold increase. Depth of the litter layer and maximum height of grass and herbaceous vegetation were greater 4–7 years after initial infestation compared to uninfested plots, though understory plant percent cover was not different. Seedling and sapling density of all species combined was higher in uninfested plots but there was no difference between infested and uninfested plots for lodgepole pine alone. For trees ≥2.5 cm in diameter at breast height, the density of live lodgepole pine trees in mountain pine beetle-affected stands was higher than Engelmann spruce, subalpine fir, and aspen, (Populus tremuloides Michx.), in diameter classes comprised of trees from 2.5 cm to 30 cm in diameter, suggesting that lodgepole pine will remain as a dominant overstory tree after the bark beetle outbreak.     

Effects of mountain pine beetle (Dendroctonus ponderosae Hopkins) infestations on forest stand structure in the southern Canadian Rocky Mountains

Forests in the montane and lower subalpine ecoregions of the southern Canadian Rocky Mountains may have been more open and structurally diverse at the beginning of the 20th Century than today. Today, the area of mature Pinus contorta subsp. latifolia Dougl. forest that is conducive to mountain pine beetle (MPB, Dendroctonus ponderosae Hopkins) infestations, infrequent high severity fires, and herbivory appears to have increased in Banff and Kootenay National Parks. Based on a review of the literature, we hypothesised that MPB infestations increase forest stand structural diversity and tree species diversity. Stand structure parameters were investigated in mesic montane and lower subalpine stands approximately 15, 25, and 65 years after MPB outbreaks. Parameters measured were stand density (number of trees per ha), diameter at breast height, height class, species, age class distribution, and coarse woody debris mass. Influences of fire frequency, time since fire, and fire severity on these parameters were assessed to determine whether fire history had a confounding influence on stand structure. The Shannon–Wiener index indicated higher stand structural diversity 15 years but not 25 and 65 years after MPB infestations. MPB infestations led to general decreases within stands in the number of living trees, small diameter snags, and Pinus tree species and an increase within stands in the number of large diameter snags. Management that allows the occurrence of the natural fire regime of variable severity fires, in addition to some MPB infestations, would provide for more open and diverse stands. MPB infestations have some effects on stand structure that are similar to those of fire. Changes in stand structure resulting from recent declines in burning rates within Banff and Kootenay National Parks can be reversed to some extent by MPB infestations.     

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.     

Influence of fire and mountain pine beetle on the dynamics of lodgepole pine stands in British Columbia,Canada

An outbreak of the mountain pine beetle (Dendroctonus ponderosae Hopkins; MPB), currently affecting over 10.1 million hectares of lodgepole pine forests (Pinus contorta Dougl.) in British Columbia, Canada, is the largest in recorded history. We examined the dynamics of even-aged lodgepole pine forests in southern British Columbia, which were undergoing this MPB outbreak. Using dendroecology and forest measurements we reconstructed the stand processes of stand initiation, stand disturbances, tree mortality, and regeneration, and explained the current stand structure and the potential MPB impacts in selected stands. Our results indicate that stand-replacing fires initiated even-aged seral lodgepole pine stands in this region. In the absence of fire in the 20th century, multiple MPB disturbances, which each resulted in partial canopy removal, modified the simple one-layer structure of the fire-origin stands by the initiation of post-MPB disturbance regeneration layers, transforming the stands into complex, multi-aged stands. Despite high overstory mortality due to the current MPB outbreak, regeneration layers, which are likely to survive the current outbreak, will provide important ecological legacies and will contribute to mid-term timber supply.     

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.     

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.     

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.     

Impact of bark beetle (Ips typographus L.) disturbance on timber production and carbon sequestration in different management strategies under climate change

The likely environmental changes throughout the next century have the potential to strongly alter forest disturbance regimes which may heavily affect forest functions as well as forest management. Forest stands already poorly adapted to current environmental conditions, such as secondary Norway spruce (Picea abies (L.) Karst.) forests outside their natural range, are expected to be particularly prone to such risks. By means of a simulation study, a secondary Norway spruce forest management unit in Austria was studied under conditions of climatic change with regard to effects of bark beetle disturbance on timber production and carbon sequestration over a time period of 100 years. The modified patch model PICUS v1.41, including a submodule of bark beetle-induced tree mortality, was employed to assess four alternative management strategies: (a) Norway spruce age-class forestry, (b) Norway spruce continuous cover forestry, (c) conversion to mixed species stands, and (d) no management. Two sets of simulations were investigated, one without the consideration of biotic disturbances, the other including possible bark beetle damages. Simulations were conducted for a de-trended baseline climate (1961–1990) as well as for two transient climate change scenarios featuring a distinct increase in temperature. The main objectives were to: (i) estimate the effects of bark beetle damage on timber production and carbon (C) sequestration under climate change; (ii) assess the effects of disregarding bark beetle disturbance in the analysis.Results indicated a strong increase in bark beetle damage under climate change scenarios (up to +219% in terms of timber volume losses) compared to the baseline climate scenario. Furthermore, distinct differences were revealed between the studied management strategies, pointing at considerably lower amounts of salvage in the conversion strategy. In terms of C storage, increased biotic disturbances under climate change reduced C storage in the actively managed strategies (up to −41.0 tC ha⁻¹) over the 100-year simulation period, whereas in the unmanaged control variant some scenarios even resulted in increased C sequestration due to a stand density effect.Comparing the simulation series with and without bark beetle disturbances the main findings were: (i) forest C storage was higher in all actively managed strategies under climate change, when biotic disturbances were disregarded (up to +31.6 tC ha⁻¹ over 100 years); and (ii) in the undisturbed, unmanaged variant C sequestration was lower compared to the simulations with bark beetle disturbance (up to −69.9 tC ha⁻¹ over 100 years). The study highlights the importance of including the full range of ecosystem-specific disturbances by isolating the effect of one important agent on timber production and C sequestration.     

Aerially applied verbenone-releasing laminated flakes protect Pinus contorta stands from attack by Dendroctonus ponderosae in California and Idaho

We tested a new formulation of verbenone, an antiaggregation pheromone of the mountain pine beetle (Dendroctonus ponderosae Hopkins) (Coleoptera: Curculionidae: Scolytinae), for area-wide protection of lodgepole pine (Pinus contorta Douglas ex Loudon) stands in the western United States. Helicopter applications of verbenone-releasing laminated flakes were made at the rate of 370 g of active ingredient/ha to two sites, one in California and one in Idaho, during summer 2005. Each site consisted of five 20.23-ha treated plots and five matching 20.23-ha untreated plots. We assessed D. ponderosae flight into study plots using traps baited with aggregation pheromones and we tallied D. ponderosae attack rates on P. contorta trees in treated and control plots before and after application. There were no significant differences between numbers of D. ponderosae trapped on treated and control plots. However, a significantly smaller proportion of P. contorta trees was attacked in treated plots than in control plots at both sites; the attack rate in untreated stands was roughly three times that of treated stands in both California and Idaho, even at this low application rate. Furthermore, attack rate in 2004 was a significant explanatory variable for the response in 2005 regardless of treatment in both California and Idaho. There was no significant treatment effect at either site on attraction of Temnochila chlorodia (Coleoptera: Trogositidae) Mannerheim, a key predator of D. ponderosae, to the prey aggregation pheromone.     

Management strategies for a large-scale mountain pine beetle outbreak: Modelling impacts on American martens

We assessed the effectiveness of alternative forest management strategies for maintaining American martens (Martes americana) in a sub-boreal landscape subject to an extensive mountain pine beetle (Dendroctonus ponderosae) outbreak, associated salvage logging, and climate change in north-central British Columbia, Canada. We structured the analysis in a Bayesian network (BN) meta-model that incorporated the results of spatially explicit modelling of landscape conditions (natural and logging disturbance, habitat quality, number of potential territories, and connectedness of territories) with analytical population modelling. The BN meta-model was then used to examine habitat and population size responses (adult females only are presented) to management scenarios, in the context of uncertainty of model parameters, management objectives, and climate change. Status quo management is dominantly clear-cutting with 3–20% of each harvest unit retained as mature patches, with reforestation by planting in the remainder. Management options we examined were: (1) the status quo, (2) varying the total annual timber harvest on the landscape (100%, 80% or 50% of current long-term sustained yield estimates), (3) the protection of understory trees during logging, and (4) 30–70% retention of overstory (partial cutting in distinct patches <1 ha in size) in each harvest unit, for 33% or 50% of the annual timber harvest. We found that marten habitat and population size declined substantively with the beetle outbreak and associated salvage cutting. The choice of management strategy then had a long-term effect on the potential for marten recovery after the beetle outbreak. Partial cutting scenarios had the greatest average long-term marten population levels, followed in order by reduced harvest rates, understory protection, and the status quo. Management scenarios with the best chance of meeting conservation goals without over-protecting habitat (and thus unnecessarily constraining timber management) varied with the population objective chosen. The choice of management strategy will depend on the weighting of marten outcomes against the economic desirability of timber harvest strategies, willingness to gamble on climate change, and the time-frame of interest.     

The influence of mountain pine beetle outbreaks and drought on severe wildfires in northwestern Colorado and southern Wyoming: A look at the past century

Outbreaks of bark beetles and drought both lead to concerns about increased fire risk, but the relative importance of these two factors is the subject of much debate. We examined how mountain pine beetle (MPB) outbreaks and drought have contributed to the fire regime of lodgepole pine forests in northwestern Colorado and adjacent areas of southern Wyoming over the past century. We used dendroecological methods to reconstruct the pre-fire history of MPB outbreaks in twenty lodgepole pine stands that had burned between 1939 and 2006 and in 20 nearby lodgepole pine stands that were otherwise similar but that had not burned. Our data represent c. 80% of all large fires that had occurred in lodgepole pine forests in this study area over the past century. We also compared Palmer Drought Severity Index (PDSI) and actual evapotranspiration (AET) values between fire years and non-fire years. Burned stands were no more likely to have been affected by outbreak prior to fires than were nearby unburned stands. However, PDSI and AET values were both lower during fire years than during non-fire years. This work indicates that climate has been more important than outbreaks to the fire regime of lodgepole pine forests in this region over the past century. Indeed, we found no detectable increase in the occurrence of high-severity fires following MPB outbreaks. Dry conditions, rather than changes in fuels associated with outbreaks, appear to be most limiting to the occurrence of severe fires in these forests.     

Woodpeckers increase in abundance but maintain fecundity in response to an outbreak of mountain pine bark beetles

Many temperate woodpecker species are thought to be highly conservative in their fecundity with little response to fluctuations in availability of resources. In a 15-year field study in interior British Columbia, we evaluated responses in abundance and fecundity of six species of resident and migrant woodpeckers (downy woodpecker [Picoides pubescens], hairy woodpecker [Picoides villosus], American three-toed woodpecker [Picoides dorsalis], pileated woodpecker [Drycopus pileatus], northern flicker [Colaptes auratus], and red-naped sapsucker [Sphyrapicus nuchalis]) to a large-scale outbreak of mountain pine bark beetles that resulted in a strong positive pulse in food supply. Population densities of woodpecker species increased during outbreak years. Despite the year-round multi-annual increase in food resources, and in contrast to the strong increases in fecundity shown by nuthatches and chickadees, annual fecundity (as indicated by clutch size and number of nestlings that fledged) did not change for any woodpecker species over the study. Similarly, we found no changes in fecundity in response to selective forest harvesting despite numerical increases for woodpeckers at these sites. Our study confirms that these woodpecker species are conservative in their reproductive investment patterns even during strong multi-annual increases in food. Our findings indicate woodpecker populations are regulated numerically through variable survival and/or greater immigration rates, which can result in higher breeding densities temporarily during resource pulses.     

基于马尾松反转录转座子序列的IRAP分子标记开发及应用

[目的]为了丰富马尾松遗传信息,开发更多适用于马尾松的新型分子标记。[方法]依据马尾松Ty1-copia类型和Ty3-gypsy类型反转录转座子RT序列的保守区域设计引物,建立了马尾松IRAP-PCR技术体系并以12个基因型个体为材料进行验证。[结果]42条引物中筛选出多态性丰富、重复性好的29条进行PCR扩增,共获得227条谱带,其中多态性条带207个,多态性比例为91.19%,平均观测等位基因数(Na)为1.911 9±0.284 1,有效等位基因数(Ne)为1.468 0±0.288 2,Nei’s基因多样性指数(H)为0.291 1±0.144 9,Shannon’s信息指数(I)为0.447 2±0.195 3;利用引物P-12、P-15或R-1,可以将栽培种与无性系两类区分开;P-2可作为核心引物,能将12份供试材料有效区分开来;采用IRAP标记构建了供试种质的DNA指纹图谱;供试种质的遗传相似系数为0.46 0.69,以相似系数为基础,进行UPGMA聚类分析,以0.57为阈值可将供试种质分为三类,其中无性系内不同材料间也存在较大的遗传变异。[结论]IRAP分子标记能有效地用于马尾松种质的鉴别与亲缘关系分析等相关研究。     

Effects of crown-pruning on seed and pollen cone production in two lodgepole pine seed orchards in British Columbia

The effects of top pruning on subsequent seed and pollen cone production in lodgepole pine (Pinus contorta Dougl.) grafted clones in two seed orchards were evaluated. In the older orchard (14 years old), three pruning regimes were applied: an untreated control, a moderate pruning where 25% of the crown was removed and a severe pruning where 40% of the crown was removed. In the younger orchard (ten years old), only a control pruning and a severe pruning (50% of crown removed) were applied. Five-year seed cone production was lower in treated ramets compared to controls in both orchards. However, when cumulative seed cone yields were adjusted for reduced tree heights after pruning, differences were no longer significant in the older orchard. Strong clonal variation was observed in seed cone yields in both orchards. Pollen cone development was less variable among treatments in the older orchard. In the younger orchard, controls consistently produced heavier pollen crops. Pruning recommendations for similar orchards are made and results discussed for reducing the impacts of losses in seed cone yields.Deceased     

Somatic embryo production and plant regeneration of Japanese black pine (Pinus thunbergii)

Somatic embryogenesis in Pinus thunbergii was initiated from megagametophytes containing immature zygotic embryos. Embryogenic cultures were maintained and proliferated in medium supplemented with 2,4-dichlorophenoxyacetic acid and 6-benzylaminopurine. High maturation frequencies of somatic embryos were obtained on maturation media containing maltose, activated charcoal, abscisic acid, and polyethylene glycol as osmotic agent. The best result among the cell lines tested was achieved with the cell line T-205-3. More than 900 somatic embryos per petri dish, on average, were obtained after about 8 weeks of culture on maturation medium. Sixty percent of somatic embryos tested germinated after transfer to plant growth regulator-free medium and then 85% of them converted into plantlets.     

Changes in the spatial structure of oak carbon-based secondary compounds after pine harvesting

In natural plant populations, leaf polyphenols show high intraspecific variation that occurs both temporally and spatially. Leaf phenolics may be induced by diverse ecological factors such as light, nitrogen availability or herbivory attack. Both light and nitrogen availability can show spatial structure in forested stands, meaning that they each have a high degree of autocorrelation, which can determine the appearance of spatial structure in leaf polyphenols. However, the availability of these resources may be drastically changed by forest disturbance, and little is known about the effect of forest disturbance on the spatial pattern and scale of leaf secondary compounds. We hypothesise that the spatial structure of leaf polyphenols in understory vegetation will disappear due to forest harvesting, because these compounds depend on light availability, yet it will remain unaltered for those compounds that either depend on the availability of other resources or are under major genetic control. The study was performed in young pedunculate oak (Quercus robur) populations growing either under a pine canopy (Pinus pinaster) stand or in a pine harvested stand in NW Spain. The spatial structures of green and senescent leaf polyphenols, tannins, non-tannin polyphenols and nitrogen were analysed in both stands using geostatistical analysis. The spatial structures observed for green and senescent leaf polyphenols and tannins in the forested stand disappeared in the harvested stand. However, non-tannin polyphenols, as well as nitrogen, showed spatial structure in both stands. Understanding these changes may be important for the successful recovery of native oak populations growing under pine forests in NW Spain, one of the priorities of the local government. Our results showed that changes in the concentration of leaf secondary compounds after disturbance may be accompanied by differences in their spatial properties, which may have important consequences for ecosystem function.     

寄主树木、寄主和环境因子对松褐天牛深沟茧蜂寄生率的影响研究

[目的]松材线虫病是我国对森林危害和威胁最严重的病害,控制其传播媒介昆虫松褐天牛是防治松材线虫病的主要手段。作者发现的松褐天牛深沟茧蜂(Iphiaulax monochamusi Yang)是寄生松褐天牛中老龄幼虫的重要天敌。为明确松褐天牛深沟茧蜂的寄生率与寄主树木、寄主和环境因子的关系,开展了本研究。[方法]通过解剖46株松褐天牛危害致死的马尾松,调查了松褐天牛深沟茧蜂寄生率与马尾松、松褐天牛和环境因子之间的关系。[结果]调查研究表明:寄主树木的高度、胸径和树龄对松褐天牛深沟茧蜂寄生率没有显著影响,松褐天牛的数量对松褐天牛深沟茧蜂寄生率亦没有显著影响,而松褐天牛幼虫龄期和松褐天牛幼虫在树干上的位置显著影响松褐天牛深沟茧蜂寄生率。松褐天牛深沟茧蜂寄生率与松褐天牛龄期呈显著正相关,其偏好寄生3~5龄幼虫;就天牛在其危害寄主树木上的位置而言,松褐天牛深沟茧蜂偏好寄生马尾松主干上部和在韧皮部危害的松褐天牛幼虫,其对位于马尾松树干上部的寄主幼虫寄生率最高,达27.38%,对在韧皮部生活、危害的寄主幼虫寄生率为20.18%,显著高于位于木质部生活的寄主幼虫寄生率(5.46%)。逐步回归分析表明:影响松褐天牛深沟茧蜂寄生率的关键环境因子是寄主树木的坡位和其它天敌寄生率,其中,坡位与松褐天牛深沟茧蜂呈显著正相关,其它天敌寄生率与松褐天牛深沟茧蜂寄生率呈显著负相关。[结论]以上研究初步明确了松褐天牛深沟茧蜂的寄生率与寄主树木、寄主害虫和天敌之间的关系,为今后利用该重要天敌控制松褐天牛打下了基础。     

Regeneration of Rocky Mountain bristlecone pine (Pinus aristata) and limber pine (Pinus flexilis) three decades after stand-replacing fires

Rocky Mountain bristlecone pine (Pinus aristata) and limber pine (Pinus flexilis) are important high-elevation pines of the southern Rockies that are forecast to decline due to the recent spread of white pine blister rust (Cronartium ribicola) into this region. Proactive management strategies to promote the evolution of rust resistance and maintain ecosystem function require an improved understanding of the role of disturbance on the population dynamics of both species and environmental conditions that favor seedling establishment. We examined patterns of bristlecone and limber pine regeneration across the perimeters of three, 29-year-old, high-severity burns in northern, central, and southern Colorado: Ouzel, Badger Mountain, and Maes Creek, respectively. Both species exhibited a very protracted regeneration response to these fires. Bristlecone pine regeneration was concentrated near burn edges and beneath surviving seed sources. This spatial pattern is consistent with limitations incurred by wind-dispersal, also borne out by the low occurrence of seedling clusters. Relative to unburned stands, the absolute abundance of bristlecone pine generally increased only on plots retaining some surviving trees. Limber pine regeneration pattern varied between sites: high in the burn interior at Ouzel, concentrated at burn edges at Badger, and mostly in unburned stands at Maes. Clark’s Nutcracker dispersal of limber pine in each study area was indicated by high seedling distance from possible seed sources and high frequencies of clustered stems. Except at Ouzel, the absolute abundance of limber pine decreased in burns. Across sites, establishment by both species was boosted by nearby nurse objects (rocks, fallen logs, and standing tree trunks), a relationship that extended out at least as far as the closest three such objects, usually found within 50 cm. Fire decreased the frequency of Pedicularis but increased Castilleja and Ribes species (alternate hosts of white pine blister rust), though only one species, R. cereum, was positively associated with either pine species. We conclude that regeneration of bristlecone and limber pine may benefit from natural disturbance or proactive management creating appropriately sized openings and microtopographic structure (e.g., abundant fallen logs); however, beneficial responses may require many decades to be achieved.     

Pronounced fluctuations of spruce bark beetle (Scolytinae: Ips typographus) populations do not invoke genetic differentiation

The dynamics of a recent outbreak of the spruce bark beetle (Ips typographus) in Switzerland was ruled by a devastating winter storm in 1999 and the drought and heat of the summer 2003. Starting from a similar level of population sizes, estimated as the rate of infested growing stock, beetle populations increased differently in magnitude and time among different regions in Switzerland. Accordingly, we expected local or regional genetic differentiation as a result of such repeated population expansion/breakdown dynamics. We analyzed 5 nuclear microsatellites of spruce bark beetles sampled from pheromone traps at 30 locations distributed over Switzerland. Our genetic results did not indicate any sign of population differentiation, structure, isolation by distance, or recent bottlenecks. This complete lack of genetic structure suggests that spruce bark beetles are highly mobile, precluding the formation of a spatial structure at neutral molecular markers. Thus, this molecular–genetic approach does not allow us to discriminate among regional gene pools and to identify the origin of expanding beetle populations.     

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

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