Efficacy of Beauveria bassiana (Bals.) Vuill. against the spruce bark beetle, Ips typographus L., in the laboratory under various conditions

Abstract:  In laboratory bioassays, the efficacy of the entomopathogenic fungus Beauveria bassiana against the spruce bark beetle, Ips typographus , was tested under various conditions. Four of the tested isolates and the commercial product Boverol^® caused 99–100% mortality when tested at a concentration of 1.0 × 10⁷ conidia/ml at 25°C. Using B. bassiana isolate 138 at a concentration of 1.0 × 10⁶, the median survival time (MST) was 6.1 d and significantly longer compared with the MST of 4.2 and 4.0 d at 1.0 × 10⁷ and 1.0 × 10⁸ conidia/ml, respectively. In the next experiment, the beetles were maintained on spruce bark, filter paper or artificial diet during the bioassay with Boverol^®, and significant differences in the MST of 3.6, 2.5 and 5.3 d, respectively, were noticed. The experiment with Boverol^® at different temperatures showed that the beetles lived significantly longer at 15°C (MST 8.7 d) than at 20, 25, 30 and 35°C. At 25°C, the beetles died most rapidly (MST 3.5 d). At different relative humidities (RH) of 40, 70 and 100%, nearly all beetles were dead after treatment with a suspension of Boverol^® at 1.0 × 10⁷ conidia/ml. At 40% RH, 49% of the untreated beetles died after 7 d. The best effects were achieved with the following bioassay: beetles were fed for three days on artificial diet, then dipped into a solution of 1.0 × 10⁷ conidia/ml and transferred on a piece of spruce bark in Petri dishes at 25°C and 70% RH.     

Influence of age, mating status, sex, quantity of food, and long-term food deprivation on red flour beetle (Coleoptera: Tenebrionidae) flight initiation

Effects of age, sex, presence or absence of food, mating status, quantity of food, and food deprivation on rate of and time of flight initiation of the red flour beetle, Tribolium castaneum (Herbst), were determined. Flight initiation declined with increasing age in both presence and absence of food. However, flight initiation was lower when food was present in the flight chambers than in the absence of food. In the presence of food, both mated and virgin beetles were equally likely to disperse by flight. However, in the absence of food, mated beetles initiated flight more readily that virgin individuals. Flight initiation was greatest when little or no food was present. The presence of varying quantities of food inside the flight chambers impacted the number of progeny produced by females before flight, but not the timing of flight. Rate of flight initiation was higher for beetles deprived of food for short periods of time compared with flight initiation of beetles with food in the flight chamber. Flight initiation decreased with increasing time without food. There were no differences in flight tendencies between males and females in the experiments reported here. Our results suggest that T. castaneum uses flight as a mechanism to disperse to new environments during almost any part of their life span and that this type of dispersion does not fit with the model of the so-called true migratory species that involves an "oogenesis-flight syndrome.     

Multipartite symbioses among fungi, mites, nematodes, and the spruce beetle, Dendroctonus rufipennis

The spruce beetle, Dendroctonus rufipennis, is an eruptive forest pest of significant economic and ecological importance. D. rufipennis has symbiotic associations with a number of microorganisms, especially the ophiostomatoid fungus Leptographium abietinum. The nature of this interaction is only partially understood. Additionally, mite and nematode associates can mediate bark beetle-fungal interactions, but this has not yet been studied for spruce beetles. In this study, we found eight mite species associated with spruce beetles: Tarsonemus ips, T. endophloeus, Histiogaster arborsignis, Dendrolaelaps quadrisetus, Proctolaelaps hytricoides, Trichouropoda alascae, T. n. sp. nr dalarenaensis, and Urobovella n. sp 767. The most prevalent species was H. arborsignis. In addition, 75% of beetles examined carried nematodes, with six species represented. These included a new species of Parasitorhabditis, Ektaphelenchus obtusus, Bursaphelenchus n. sp. 727, Aphelenchoides n. sp., Panagrolaimus sp., and Mykoletzkya ruminis. H. arborsignis showed strong feeding and oviposition preferences for L. abietinum among four fungal species tested in laboratory assays. Information on our attempts to culture the various nematode species collected from D. rufipennis is also provided. Bursaphelenchus were cultured from D. rufipennis nematangia plated on agar containing L. abietinum but not sterile agar. Thus, L. abietinum plays an important role in these gallery communities, affecting the tree-killing bark beetle, its phoretic mites, and nematodes. These data add to our understanding of bark beetle-microorganism interactions.     

Fungi Vectored by the Bark Beetle Ips typographus Following Hibernation Under the Bark of Standing Trees and in the Forest Litter

The bark beetle Ips typographus has different hibernation environments, under the bark of standing trees or in the forest litter, which is likely to affect the beetle-associated fungal flora. We isolated fungi from beetles, standing I. typographus-attacked trees, and forest litter below the attacked trees. Fungal identification was done using cultural and molecular methods. The results of the two methods in detecting fungal species were compared. Fungal communities associated with I. typographus differed considerably depending on the hibernation environment. In addition to seven taxa of known ophiostomoid I. typographus-associated fungi, we detected 18 ascomycetes and anamorphic fungi, five wood-decaying basidomycetes, 11 yeasts, and four zygomycetes. Of those, 14 fungal taxa were detected exclusively from beetles that hibernated under bark, and six taxa were detected exclusively from beetles hibernating in forest litter. The spruce pathogen, Ceratocystis polonica, was detected occasionally in bark, while another spruce pathogen, Grosmannia europhioides, was detected more often from beetles hibernating under the bark as compared to litter. The identification method had a significant impact on which taxa were detected. Rapidly growing fungal taxa, e.g. Penicillium, Trichoderma, and Ophiostoma, dominated pure culture isolations; while yeasts dominated the communities detected using molecular methods. The study also demonstrated low frequencies of tree pathogenic fungi carried by I. typographus during its outbreaks and that the beetle does not require them to successfully attack and kill trees.     

Reducing spruce beetle Dendroctonus rufipennis (Kirby) (Coleoptera: Curculionidae) emergence for hibernation in central British Columbia by felling infested trees

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Horizontal Transmission of the Entomopathogenic Fungus Beauveria bassiana among the Spruce Bark Beetle, Ips typographus (Col., Scolytidae) in the Laboratory and under Field Conditions

In laboratory and field cage experiments, the horizontal transmission of the entomopathogenic fungus Beauveria bassiana (Boverol®) between adults of the spruce bark beetle Ips typographus was examined. In a laboratory experiment, conidia transfer between treated and untreated beetles at different ratios resulted in 96% mortality (MST 4.3 days) at a ratio of 1:1 and 90% mortality (MST 4.5 days) at a ratio of 1:2. At ratios of 1:5, 1:10 and 1:20, the mortalities at 7 days were 83, 77 and 75%, respectively. In a second laboratory experiment, the efficacy of B. bassiana was investigated after transmission from contaminated to healthy beetles over a period of 5 days. Two weeks later, the mortality of treated and untreated beetles was 99% (control mortality was 44%), while the rate of mycosis was 79% in the treatment compared to 10% in the control beetles. A single contact between one treated male (2.0×10⁵ conidia/beetle) and an untreated female was found to be sufficient to transmit a lethal dose of B. bassiana of about 1.2×10⁴ conidia. Two experiments in field cages in a spruce stand demonstrated horizontal transmission of B. bassiana (Boverol®) between beetles. In the first experiment the transmission was investigated following the introduction of B. bassiana-inoculated beetles into a population of untreated beetles. Significant reductions were observed in the length of maternal galleries, and the number of larvae and pupae. In the second field cage experiment, natural populations of beetles were lured into a flight cage using a pheromone trap, and treated with B. bassiana or left untreated on alternate days. In this experiment, significant reductions were observed in the number of bore holes and the length of maternal galleries, while no larvae, pupae or juveniles were found under the bark of spruce trunks in the Boverol® transmission treatment. It is concluded that B. bassiana has potential for control of I. typographus, and further field experiments are warranted to investigate its efficacy under more practical conditions.     

Investigating the abundance and flight period of bark beetles (Coleoptera: Curculionidae: Scolytinae) over elevational gradients in Sitka spruce forests

1. A warming climate, as predicted under current climate change projections, is likely to influence the population dynamics of many forest insect species. Numerous bark beetle species in both Europe and North America have already responded to a warming climate by significantly expanding their geographical ranges.
2. The aim of the current study was to investigate how populations of bark beetles within stands of Sitka spruce, a widely planted non-native commercial plantation tree species in the U.K., were likely to respond to a warming climate. Experimental plots were established in stands of Sitka spruce over elevational gradients in two commercial forest plantations, and the abundance and emergence times of key bark beetle species were assessed over a 3-year period using flight interception traps. The air temperature difference between the lowest and highest experimental plot in each forest was consistently >1°C throughout the 3-year period.
3. In general, the abundance of the most dominant bark beetle species (e.g. Trypodendron, Dryocoetes, Hylastes spp.) was higher, and emergence times tended to be earlier in the year at the lower elevation plots, where temperatures were higher, although not all bark beetle species responded in the same manner.
4. The results of the study indicated that, under the projected future climate warming scenarios, monoculture Sitka spruce stands at low elevations may potentially be more vulnerable to significant outbreak events from existing or invasive bark beetle species. Hence, consideration of establishing more resilient forests of Sitka spruce by diversifying the species composition and structure of Sitka spruce stands is discussed.

     

Inducibility of chemical defenses in Norway spruce bark is correlated with unsuccessful mass attacks by the spruce bark beetle

Secondary attraction to aggregation pheromones plays a central role in the host colonization behavior of the European spruce bark beetle Ips typographus. However, it is largely unknown how the beetles pioneering an attack locate suitable host trees, and eventually accept or reject them. To find possible biomarkers for host choice by I. typographus, we analyzed the chemistry of 58 Norway spruce (Picea abies) trees that were subsequently either (1) successfully attacked and killed, (2) unsuccessfully attacked, or (3) left unattacked. The trees were sampled before the main beetle flight in a natural Norway spruce-dominated forest. No pheromones were used to attract beetles to the experimental trees. To test the trees' defense potential, each tree was treated in a local area with the defense hormone methyl jasmonate (MeJ), and treated and untreated bark were analyzed for 66 different compounds, including terpenes, phenolics and alkaloids. The chemistry of MeJ-treated bark correlated strongly with the success of I. typographus attack, revealing major chemical differences between killed trees and unsuccessfully attacked trees. Surviving trees produced significantly higher amounts of most of the 39 analyzed mono-, sesqui-, and diterpenes and of 4 of 20 phenolics. Alkaloids showed no clear pattern. Differences in untreated bark were less pronounced, where only 1,8-cineole and (-)-limonene were significantly higher in unsuccessfully attacked trees. Our results show that the potential of individual P. abies trees for inducing defense compounds upon I. typographus attack may partly determine tree resistance to this bark beetle by inhibiting its mass attack.     

Bark beetles as lidar targets and prospects of photonic surveillance

Forestry is raising concern about the outbreaks of European spruce bark beetle, Ips typographus, causing extensive damage to the spruce forest and timber values. Precise monitoring of these beetles is a necessary step towards preventing outbreaks. Current commercial monitoring methods are catch-based and lack in both temporal and spatial resolution. In this work, light scattering from beetles is characterized, and the feasibility of entomological lidar as a tool for long-term monitoring of bark beetles is explored. Laboratory optical properties, wing thickness, and wingbeat frequency of bark beetles are reported, and these parameters can infer target identity in lidar data. Lidar results from a Swedish forest with controlled bark beetle release event are presented. The capability of lidar to simultaneously monitor both insects and a pheromone plume mixed with chemical smoke governing the dispersal of many insects is demonstrated. In conclusion, entomological lidar is a promising tool for monitoring bark beetles.     

The European spruce bark beetle <Emphasis Type="Italic">Ips typographus</Emphasis> in a national park: from pest to keystone species

The influence of natural disturbance on biodiversity is poorly known in the intensively cultivated landscape of Europe. As an example of insect disturbance we studied effects of gaps generated by outbreaks of the spruce bark beetle (Ips typographus) on biodiversity in the area of the National Park “Bavarian Forest” and compared them with openings (e.g. meadows) created by humans in these forests. Insects were sampled using flight interception traps across twelve ecotones between edges of closed forest, six bark beetle gaps and six meadows. The diversity and species density of true bugs and of bees/wasps increased significantly from the closed stand to the edge, and continued to increase inside the openings at interior and exterior edges. Species density in saproxylic beetles also increased significantly from closed forest to opening, but only across ecotones including bark beetle gaps. Similarly, the number of critically endangered saproxylic beetles increased significantly in bark beetle gaps. Using indicator species analysis a total of 60 species were identified as possessing a statistically significant value indicating preference for one of the habitat types along the ecotones: 29 of them preferred gaps, 24 preferred meadows, three were characteristic for edges of meadows, three for edges of bark beetle gaps, but only one was typical of closed forest. Most of our results support the thesis that I. typographus fulfils the majority of criteria for a keystone species, particularly that of maintenance of biodiversity in forests. Our results emphasize the value for the study and conservation of insect diversity of the policy of non-interference with natural processes pursued in some protected areas. As a recommendation to forest management for increasing insect diversity even in commercial forest, we suggest that logging in recent gaps in medium aged mixed montane stands should aim at retention of a part of the dead wood. Planting should be avoided, to lengthen the important phase of sunlit conditions.     

Are bark beetle outbreaks less synchronous than forest Lepidoptera outbreaks?

Comparisons of intraspecific spatial synchrony across multiple epidemic insect species can be useful for generating hypotheses about major determinants of population patterns at larger scales. The present study compares patterns of spatial synchrony in outbreaks of six epidemic bark beetle species in North America and Europe. Spatial synchrony among populations of the Eurasian spruce bark beetle Ips typographus was significantly higher than for the other bark beetle species. The spatial synchrony observed in epidemic bark beetles was also compared with previously published patterns of synchrony in outbreaks of defoliating forest Lepidoptera, revealing a marked difference between these two major insect groups. The bark beetles exhibited a generally lower degree of spatial synchrony than the Lepidoptera, possibly because bark beetles are synchronized by different weather variables that are acting on a smaller scale than those affecting the Lepidoptera, or because inherent differences in their dynamics leads to more cyclic oscillations and more synchronous spatial dynamics in the Lepidoptera.     

Influence of boreal forest succession and dead wood qualities on saproxylic beetles

1 Saproxylic insects, a functional group dominated by beetles, are dependent on dead or moribund trees as habitat elements. 2 Although there are few studies of saproxylic insects from the North American boreal zone, European studies demonstrate that forest harvest can lead to a biologically significant decrease in saproxylic beetle diversity. 3 We studied saproxylic beetles in the North American boreal mixedwood forest using flight intercept traps established on naturally dead and girdled trembling aspen and spruce trees along a successional gradient of undisturbed stands from deciduous to coniferous overstory trees. 4 Composition and diversity of beetle assemblages differed among forest successional types. 5 Snag age class was an important determinant of composition for saproxylic beetle assemblages. 6 Multivariate regression analysis of these data indicated that saproxylic beetles are responding to changes in coarse woody debris, and not to the relative densities of canopy tree species, although these variables are strongly correlated. 7 Coarse woody debris management should be a primary concern in forest management plans seeking to conserve saproxylic organisms and the critical ecosystem functions (i.e. nutrient cycling) in which they participate.     

Influence of elevation on bark beetle (Coleoptera: Curculionidae, Scolytinae) community structure and flight periodicity in ponderosa pine forests of Arizona

We examined abundance and flight periodicity of five Ips and six Dendroctonus species (Coleoptera: Curculionidae, Scolytinae) among three different elevation bands in ponderosa pine (Pinus ponderosa Douglas ex. Lawson) forests of northcentral Arizona. Bark beetle populations were monitored at 10 sites in each of three elevation bands (low: 1,600-1,736 m; middle: 2,058-2,230 m; high: 2,505-2,651 m) for 3 yr (2004-2006) using pheromone-baited Lindgren funnel traps. Trap contents were collected weekly from March to December. We also studied temperature differences among the elevation bands and what role this may play in beetle flight behavior. Bark beetles, regardless of species, showed no consistent elevational trend in abundance among the three bands. The higher abundances of Ips lecontei Swaine, I. calligraphus ponderosae Swaine, Dendroctonus frontalis Zimmerman, and D. brevicomis LeConte at low and middle elevations offset the greater abundance of I. knausi Swaine, D. adjunctus Blandford, D. approximatus Dietz, and D. valens LeConte at high elevations. I. pini (Say) and I. latidens LeConte were found in similar numbers across the three bands. Flight periodicity of several species varied among elevation bands. In general, the flight period shortened as elevation increased; flight initiated later and terminated earlier in the year. The timing, number, and magnitude of peaks in flight activity also varied among the elevation bands. These results suggest that abundance and flight seasonality of several bark beetles are related to elevation and the associated temperature differences. The implications of these results are discussed in relation to bark beetle management and population dynamics.     

Effects of rearing density, age, sex, and food deprivation on flight initiation of the red flour beetle (Coleoptera: Tenebrionidae)

Effects of rearing density, adult density and sex ratio in the flight chamber, adult age, sex, presence or absence of food, and duration of food deprivation on rate of and time to flight initiation of the red flour beetle, Tribolium castaneum (Herbst) (Coleoptera: Tenebrionidae), were studied in the laboratory. Rates of flight initiation were slightly lower at lower rearing densities, but they did not differ with age or sex of adults, presence or absence of food in the flight chamber, or duration of food deprivation. Focal adults were less likely to fly when individuals of the opposite sex were present in flight chambers. Presence of the same sex or mixed sexes and numbers of individuals in flight chambers did not affect tendency to fly of focal individuals. Mean time to flight of older beetles (7-20 d old) was less than that of younger beetles (1-4 d old). No young beetles flew during the first 24 h of flight tests. Mean time to flight did not differ with rearing density, sex, presence or absence of food, or duration of food deprivation. Our results indicate that this species is highly dispersive during the adult stage and that flight does not seem to be associated with just prereproductive or postreproductive dispersal phases, high rearing density, or short to medium periods of food deprivation. Therefore, T. castaneum level of flight activity does not seem to be associated with factors that have been shown in the literature to increase dispersal by walking for this species and to increase flight initiation in other stored-product species.     

Seasonal flight patterns of Ips typographus in southern Sweden and thermal sums required for emergence

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Tree species and position matter: the role of pests for survival of other insects

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Influence of temperature on spring flight initiation for southwestern ponderosa pine bark beetles (Coleoptera: Curculionidae, Scolytinae)

Determination of temperature requirements for many economically important insects is a cornerstone of pest management. For bark beetles (Coleoptera: Curculionidae, Scolytinae), this information can facilitate timing of management strategies. Our goals were to determine temperature predictors for flight initiation of three species of Ips bark beetles, five species of Dendroctonus bark beetles, and two genera of bark beetle predators, Enoclerus spp. (Coleoptera: Cleridae) and Temnochila chlorodia (Mannerheim) (Coleoptera: Ostomidae), in ponderosa pine forests of northcentral Arizona. We quantified beetle flight activity using data loggers and pheromone-baited funnel traps at 18 sites over 4 yr. Ambient air temperature was monitored using temperature data loggers located in close proximity to funnel traps. We analyzed degree-day accumulation and differences between minimum, average, and maximum ambient temperature for the week before and week of first beetle capture to calculate flight temperature thresholds. Degree-day accumulation was not a good predictor for initiation of beetle flight. For all species analyzed other than D. adjunctus Blandford, beetles were captured in traps only when springtime temperatures exceeded 15.0 degrees C. D. adjunctus was collected when maximum temperatures reached only 14.5 degrees C. Once initial flights had begun, beetles were often captured when maximum ambient air temperatures were below initial threshold temperatures. Maximum and average air temperatures were a better predictor for beetle flight initiation than minimum temperature. We establish a temperature range for effective monitoring of bark beetles and their predators, and we discuss the implications of our results under climate change scenarios.     

Attack density and breeding success of bark beetles (Coleoptera, Scolytidae) at different distances from forest-clearcut edge

1 Attack density and breeding success of bark beetles (Coleoptera: Scolytidae) and the abundance of their predators were studied at forest-clearcut edges in southern Finland on Norway spruce, Picea abies, bolts placed at different distances from a forest-clearcut edge. 2 Attack densities of two species, Hylurgops palliatus and H. glabratus, increased markedly towards the forest interior. The breeding success of H. palliatus was increased with increasing distance from the stand edge. None of the recorded 10 bark beetle species was distinctly abundant near the edge, but Pityogenes chalcographus appeared to prefer this zone. 3 There were no significant differences in numbers of bark beetle predators between the forest interior and the edge. Forest-clearcut edge had a significant effect on the desiccation of sample bolts. 4 Thus, forest edge is likely to affect the colonization behaviour and breeding success of bark beetles via host material properties.     

Spatio-temporal impact of climate change on the activity and voltinism of the spruce bark beetle, Ips typographus

The spruce bark beetle Ips typographus is one of the major insect pests of mature Norway spruce forests. In this study, a model describing the temperature-dependent thresholds for swarming activity and temperature requirement for development from egg to adult was driven by transient regional climate scenario data for Sweden, covering the period of 1961–2100 for three future climate change scenarios (SRES A2, A1B and B2). During the 20th century, the weather supported the production of one bark beetle generation per year, except in the north-western mountainous parts of Sweden where the climate conditions were too harsh. A warmer climate may sustain a viable population also in the mountainous part; however, the distributional range of I. typographus may be restricted by the migration speed of Norway spruce. Modelling suggests that an earlier timing of spring swarming and fulfilled development of the first generation will significantly increase the frequency of summer swarming. Model calculations suggest that the spruce bark beetle will be able to initiate a second generation in South Sweden during 50% of the years around the mid century. By the end of the century, when temperatures during the bark beetle activity period are projected to have increased by 2.4–3.8 °C, a second generation will be initiated in South Sweden in 63–81% of the years. The corresponding figures are 16–33% for Mid Sweden, and 1–6% for North Sweden. During the next decades, one to two generations per year are predicted in response to temperature, and the northern distribution limit for the second generation will vary. Our study addresses questions applicable to sustainable forest management, suggesting that adequate countermeasures require monitoring of regional differences in timing of swarming and development of I. typographus , and planning of control operations during summer periods with large populations of bark beetles.