Quantifying spatio-temporal dispersion of bark beetle infestations in epidemic and non-epidemic conditions

Spatio-temporal dispersal of pest species such as bark beetles plays a key role in their population ecology and outbreak dynamics. Understanding the underlying patterns is crucial for applying appropriate management strategies.In contrast to most existing studies which focus on dispersing beetles, we analysed patches of killed trees resulting from bark beetle infestation. The study was based on a 22-year time series of annually captured colour-infrared (CIR) images of the Bavarian Forest National Park (Germany), where Ips typographus L. (Coleoptera, Curculionidae, Scolytinae) propagates undisturbed by human activity. Newly infested patches comprising at least 5 spruce trees were identified in every time step. This investigation of spatio-temporal spread of infestations primarily focused on (i) parameterizing the size and shape of infestation patches, (ii) modelling an infestation gradient and (iii) evaluating the risk of subsequent infestations on landscape scale. We developed a GIS-based distance ring approach to quantify the distance relation of subsequent infestations, including the distribution of potential hosts.Infestation spread was revealed to be strongly distance dependent, following an inverse power law function: on average 65% of new infestations occurred within a 100 m radius of the previous year’s infestations, and 95% within 500 m. ‘Distance’ proved to be a major determinant of I. typographus dispersal on the landscape scale in each time step of the 22-year series we investigated. Infestation distance thus describes the outcome of beetle dispersal very accurately. The time series showed two alternating periods of epidemic and non-epidemic infestation. These gradation stages did not affect the size and shape of infested patches, but epidemics correlated significantly with a higher percentage of infestations within short distances. Additionally, the resulting infestation risk is highly sensitive to the gradation stage, particularly within the first 100 m around source spots where it increases up to 30%.Our study therefore contributes to a better understanding of the outbreak dynamics of I. typographus and suggests concentrating efficient bark beetle management on areas in the close vicinity of previous years’ infestations.     

Persisting bark beetle outbreak indicates the unsustainability of secondary Norway spruce forests: case study from Central Europe

? Context

Secondary Norway spruce forests in the Western Beskids are among the most damaged forests in Europe. Although spruce bark beetle (Ips typographus) has been recently causing large-scale damage to these forests, our understanding of I. typographus dynamics in this environment is inadequate for evaluating forest sustainability.

? Aim

This study aims to evaluate the patterns of damage caused by I. typographus to spruce forests with compromised ecological stability.

? Methods

Forest infestation by I. typographus was inferred from sanitary felling data collected from 1998 to 2004. Stand and site data were obtained from forest management plans. Spatial-dependence analysis, ordinary kriging and neural network-based regression modelling were used to investigate the patterns of infestation and the casual relationships in the studied ecosystem.

? Results

I. typographus long-distance dispersal substantially decreased with outbreak culmination. The spread of infestation was only weakly related to stand and site parameters. Infestations spread isotropically at the stand and patch level but directionally at the regional scale.

? Conclusions

The large-scale spread of infestation can be explained by the uniform age and species composition of the investigated forests and by the ability of populations to overwhelm suboptimal trees. The observations presented here suggest that secondary spruce forests in Europe may be unsustainable due to unprecedented bark beetle outbreaks, which can be further amplified by changing climate.     

Modelling the predisposition of Norway spruce to <Emphasis Type="Italic">Ips typographus</Emphasis> L. infestation by means of environmental factors in southern Finland

New measures for effective monitoring and controlling of bark beetle infestations are needed as a response to intensified outbreaks caused by the climate change. Various environmental factors affect tree health and susceptibility, as well as stand predisposition to bark beetles. European spruce bark beetle Ips typographus L. abundance and outbreak frequency in Finland has significantly increased during the last decade. The ability to identify sites under a high risk of infestation would facilitate adaptation to this new situation and help target limited forest health management resources. Accordingly, our goal was to investigate the importance of various stand, soil and topographic characteristics in the assessing predisposition of Norway spruce dominated urban forest in southern Finland to I. typographus infestations. Information on the environmental factors was assessed in the field in 2014 and derived from a digital elevation model. Ips typographus infestation intensity was classified into three infestation index classes based on tree-wise symptoms of resin flow, discoloration and defoliation. Cumulative logit link models were utilized for investigating stand-level infestation probability. The best explanatory factors were aspect, slope, site type and soil texture. Models with the highest cumulative probabilities for severe infestation were linked with eastern aspect, moderate steep slope and rich site type fertility (0.72) and eastern aspect, shallow soil and rich site type fertility (0.71). Higher soil C/N ratios with east aspect and rich site type fertility was associated with an increased risk of severe infestation in a third model. The lowest risk was associated with southern and southwestern aspects, fine soil texture, moderate site fertility and gentle slopes.     

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.     

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.     

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.     

The “sun-effect”: microclimatic alterations predispose forest edges to bark beetle infestations

Bark beetle dispersal and host selection behaviour are a complex and poorly understood process, resulting in specific spatio-temporal infestation patterns in forests. Aerial images from the Bavarian Forest National Park (Germany) provide a high-resolution, that is, tree-scale data set for the period 2001–2010, including information about Ips typographus (Col., Curculio., Scolytinae) infestation, the application of sanitary logging, natural forest edges and the area of living spruce susceptible to bark beetle infestation. We combined methods of GIS and image analysis to investigate the infestation probabilities at three types of forest edges under spatial and temporal aspects and compared them to the corresponding probabilities at the stand interior. Our results showed a pronounced infestation predisposition of such edge trees delimiting infestation patches cleared by sanitary logging measures, in particular at the south-facing edge sector. In contrast, edges adjacent to non-cleared infestation were revealed as less attractive for subsequent infestations, but nonetheless more attractive than permanent forest edges or the stand interior. Additionally, we measured near-bark surface air temperature to determine microclimatic differences at those edge- or non-edge sites and related them to predisposition results. Finally, our study emphasized favourable microclimatic conditions—summarized as the “sun-effect”—as a decisive factor enhancing the local infestation probability at recent forest edges in multiple ways. Both insect- and host tree-related reactions to suddenly altered microclimate are supposed to bias arbitrary colonization behaviour at patch and tree level, thereby mainly explaining observed infestation patterns. From the forester’s point of view, our results may contribute to precise bark beetle risk assessment and thus facilitate decision making in forest management.     

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.     

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.     

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.     

Types of bark beetle (Coleoptera: Scolytidae) infestation in spruce forest stands affected by air pollution, bark beetle outbreak and honey fungus (Armillaria mellea)

583 spruce stands in an area affected by air pollution and bark beetle outbreak in Eastern Slovakia were studied in 1996. According to bark beetle infestation of dominant and codominant trees, stands were classified into following types of spruce stand decline:Ips typographus-A,Ips typographus-B,Polygraphus poligraphus, I. typographus/P. poligraphus—A,I. typographus/P. poligraphus—B. The presence of attacked trees in forest edges, bark beetle spots and forest interior was the key important factor for the classification. Data from forest inventory and forest management evidence together with data on types of spruce stands decline were used in further analyses. Results shows that the distribution of forest stands classified into different types or uninfested stands is related mainly to host size and site quality. The percentage of spruce, exposition of stands and stand density showed significant effects. The mechanisms of spreading of studied bark beetle outbreak could be explained by direct effects of stress of trees caused by an abrupt increase of level of solar irradiation and by weakening of trees by the honey fungus.     

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.     

Occurrence of pathogens in associated living bark beetles (Col., Scolytidae) from different spruce stands in Austria

Pathogen occurrence was studied in 16?099 adult specimens of 10 different bark beetle species, which live associated on Norway spruce (Picea abies (L.) Karst.). Beetles (mainly Ips typographus L. and Pityogenes chalcographus L.) were collected from 6 different localities in Austria (4 secondary spruce stands and 2 natural forest type stands, 9 sampling plots in total) in elevations between 400?m and 1600?m. Various viral, protozoan, and fungal pathogens could be diagnosed with a light microscope in the examined beetles. Numerous pathogen species were known from former studies, some pathogens were totally new or could be found in a new host species beside their type host. The most dominant pathogen species were Protozoa, Gregarina cf. typographi, Malamoeba cf. scolyti, and Chytridiopsis cf. typographi. Over the whole investigation period, the highest pathogen diversity with eight pathogen species was found in I. typographus. Differences were observed in the pathogen complex of each beetle species from the different collection sites and in different years of investigation. Several species showed an overlapping in their host range and infected various bark beetle species. Furthermore, pathogen occurrence and prevalence differed in bark beetles from 4 different sampling plots in an area (one locality) within a distance of a few kilometres.     

Damage reduction and performance of mass trapping devices for forest protection against the spruce bark beetle, Ips typographus (Coleoptera Curculionidae Scolytinae)

? The spruce bark beetle, Ips typographus, is one of the main European forest pests, and mass trapping is probably the most common strategy applied to reduce its population density. However, the results concerning the effectiveness of this control system are often controversal, and many studies consider only the trapping performance with no attention to the damage reduction.

? During spring-summer 2005, a control program against I. typographus outbreaks was set up in NE Italy. Twenty-four spruce forests heavily infested by I. typographus were studied: six protected by pheromone slot-traps, six by horizontal trap-logs and six by standing trap-logs; six untreated stands were kept as controls. Trap-logs were baited with a pheromone specific to I. typographus and treated with insecticide. Each type of device was tested at high, medium and low density in relation to the number of trees infested during the previous year. New damage occurring in the investigated stands was later monitored for one year.

? Protected forests showed mean damage about 80% lower in 2005 than in 2004, with no statistical difference among traps, trap-logs or standing trap-logs. Instead, unprotected forests (controls) suffered damage to a similar extent in both years. Trapping devices showed no statistical differences among mean captures. Device densities showed similar results in damage reduction and insect trapping.

? The results support the hypothesis that intensive trapping performed at stand level may be useful for protecting forests against I. typographus, locally reducing population density and tree mortality.

     

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.     

Ophiostomatoid fungi associated with bark beetles onAbies veitchii in wave-regenerated forests

Relationships between tree mortality and bark beetle infestation onAbies veitchii at the wave-regenerated forest in Mt. Asahi, Okuchichibu area were investigated. Most of the firs with green needles and newly developed current year’s shoots in the dieback zone were heavily infested by bark beetles before the death of the trees. After heavy infestation of beetles, about half of the infested firs died within the year, and the other half died in the next year. When the species composition of bark beetles and associated ophiostomatoid fungi were investigated in Mt. Asahi and also at a typical wave-regenerated forest in Mt. Shimagare, Yatsugatake area,Cryphalus montanus andC. piceae were dominant beetle species for Mt. Asahi and Mt. Shimagare, respectively.Ophiostoma subalpinum andO. europhioides were dominant fungal species at both wave-generated forests. BecauseO. subalpinum was detected more frequently from deeper areas of sapwood thanO. europhioides, it was suggested that the fungal species may accelerate the death of stressed firs in wave-regenerated forests. Contribution No. 169, Laboratory of Plant Parasitic Mycology, Institute of Agriculture and Forestry, University of Tsukuba.     

Winter mortality in sub-corticolous populations of Ips typographus (Coleoptera, Scolytidae) and its parasitoids in the south-eastern Alps

A study concerning the winter mortality of a sub-corticolous population of Ips typographus and its parasitoids has been carried out in the south-eastern Alps (Italy) during the winter 1997/98. Three attacked spruce trees were sampled three times (November, February and April) by collecting infested bark disks (1?dm² each). All insects that emerged from the bark or died in the samples before emergence were counted. The mean number of living Ips typographus occurring under the bark decreases by 49?% from November to April. Winter mortality mainly affects larval stages and young adults. The same trend was observed for the parasitoids Coeloides bostrychorum (48.5?%) and Roptrocerus xylophagorum (47.5?%).     

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.     

Fungi isolated from Picea abies infested by the bark beetle Ips typographus in the Białowieża forest in north‐eastern Poland

The assemblage of fungi occurring in the sapwood of Norway spruce (Picea abies) and in bark beetle galleries following attack by the Eurasian spruce bark beetle Ips typographus was investigated in the Bia?owie?a forest in north‐eastern Poland. Fungi were isolated from blue‐stained sapwood of beetle‐infested spruce trees in June 2002, and a few isolates were also obtained from ascospores and conidia taken from perithecia and asexual structures occurring in the gallery systems of the insects. The mycobiota of I. typographus in the Bia?owie?a forest was dominated by ophiostomatoid fungi, which were represented by seven species. Four species, including Ceratocystis polonica, Grosmannia penicillata, Ophiostoma ainoae and Ophiostoma bicolor were isolated at high frequencies, whereas three other taxa, Ceratocystiopsis minuta, Ceratocystiopsis alba and a Pesotum sp. were rare. The anamorphic fungus Graphium fimbriisporum and yeasts also occurred occasionally. In addition, the basidiomycete Gloeocystidium ipidophilum was relatively common. The pathogenic blue‐stain fungus C. polonica was the dominant fungal associate of I. typographus in the Bia?owie?a forest, which is consistent with a previous study at this area in the 1930s. Ceratocystis polonica was the most frequently isolated species at the leading edge of fungal colonization in the sapwood and had on an average penetrated deeper into the wood than other fungal associates. This suggests that it acts as a primary invader into the sapwood after attack by I. typographus in the Bia?owie?a forest, followed by O. bicolor, O. ainoae, G. ipidophilum and G. penicillata. Thus far, the Bia?owie?a forest is one of the few areas in Europe, where C. polonica has been reported as a dominate fungal associate of I. typographus.     

Test von Monoterpenen als Zusatz zu Pheroprax® bzw. Chalcoprax® in Pheromonfallen zum Fang des Buchdruckers,Ips typographus L. bzw. des Kupferstechers,Pityogenes chalcographus L. (Col., Scolytidae)

Experiments on monoterpenes in combination with Pheroprax^® and Chalcoprax^® in pheromone traps for catching the bark beetlesIps typographus L. andPityogenes chalcographus L. (Col., Scolytidae) It is highly evident that monoterpenes of the host tree serve as olfactory stimulants for “pioneer-beetles” colonizing first a tree before any bark-beetle-produced aggregation pheromones can lure those beetles to the host. The findings ofRedemann (1993) about a significant increase of spruce engraver catches by addition of both (?)-alpha-pinen and (+)-limonen simultaneously to pheroprax-baited bark beetle traps (Pheroprax^R being the aggregation pheromone ofIps typographus) induced us to conduct field experiments testing the same host volatiles in comparable bark beetle traps baited with either Pheroprax^® or Chalcoprax^® (aggregation pheromone ofPityogenes chalcographus). Despite we used methods considering the influence of place and time on the bark beetle flight activities, in 3 experiments with 30 repetitions in total no monoterpene-induced enhancement oftypographus-catches could be found; the same was true withP. chalcographus (2 experiments, 18 repetitions in total). With respect to the important role of host volatiles also from a practical point of view, it is strongly recommended to repeat experiments like these under different conditions to reveal the reasons of the different findings.