Lack of genetic differentiation in aggressive and secondary bark beetles (Coleoptera: Curculionidae, Scolytinae) from Arizona

Bark beetles (Coleoptera: Curculionidae, Scolytinae) play an important role as disturbance agents in ponderosa pine (Pinus ponderosa Douglas ex Lawson) forests of Arizona. However, from 2001 to 2003, elevated bark beetle activity caused unprecedented levels of ponderosa pine mortality. A better understanding of the population structure of these species will facilitate analysis of their dispersal patterns and improve management strategies. Here, we use fluorescently labeled amplified fragment length polymorphism (fAFLP) analysis to resolve genetic variation among and within sampling locations in northcentral Arizona of Ips pini (Say), Dendroctonus brevicomis LeConte, and D. frontalis Zimmermann. We generated genetic fingerprints for >500 beetle specimens and analyzed genetic diversity. For all species, gene flow estimates among sampling locations were high, and significant population subdivision was not discernible across a large portion of ponderosa pine forests in Arizona. However, a weak relationship was detected with I. pini population structure and elevation. Because of the lack of genetic differentiation detected throughout the large study area, our findings suggest these insects are capable of long distance dispersal and exhibit a high degree of gene flow across a broad region. We conclude that our results are consistent with strong dispersal patterns and large population sizes of all three species.     

Impacts of silvicultural thinning treatments on beetle trap captures and tree attacks during low bark beetle populations in ponderosa pine forests of northern Arizona

Our research used a combination of passive traps, funnel traps with lures, baited trees, and surveys of long-term thinning plots to assess the impacts of different levels of stand basal area (BA) on bark beetle tree attack and on trap captures of Ips spp., Dendroctonus spp., and their predators. The study occurred at two sites in ponderosa pine, Pinus ponderosa Dougl. ex Laws., forests, from 2004 to 2007 during low bark beetle populations. Residual stand BA ranged from 9.0 to 37.0 m2/ha. More predators and bark beetles were collected in passive traps in stands of lower BA than in stands of higher BA; however, significance varied by species and site, and total number of beetles collected was low. Height of the clear panel passive traps affected trap catches for some species at some sites and years. When pheromone lures were used with funnel traps [Ips pini (Say) lure: lanierone, +03/-97 ipsdienol], we found no significant difference in trap catches among basal area treatments for bark beetles and their predators. Similarly, when trees were baited (Dendroctonus brevicomis LeConte lure: myrcene, exo-brevicomin and frontalin), we found no significant difference for days to first bark beetle attack. Surveys of long-term thinning treatments found evidence of bark beetle attacks only in unthinned plots (approximately 37 m2/ha basal area). We discuss our results in terms of management implications for bark beetle trapping and control.     

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.     

Evaluation of funnel traps for characterizing the bark beetle (Coleoptera: Scolytidae) communities in ponderosa pine forests of north-central Arizona

Lindgren funnel traps baited with aggregation pheromones are widely used to monitor and manage populations of economically important bark beetles (Coleoptera: Scolytidae). This study was designed to advance our understanding of how funnel trap catches assess bark beetle communities and relative abundance of individual species. In the second year (2005) of a 3-yr study of the bark beetle community structure in north-central Arizona pine (Pinus spp.) forests, we collected data on stand structure, site conditions, and local bark beetle-induced tree mortality at each trap site. We also collected samples of bark from infested (brood) trees near trap sites to identify and determine the population density of bark beetles that were attacking ponderosa pine, Pinus ponderosa Douglas ex Lawson, in the area surrounding the traps. Multiple regression models indicated that the number of Dendroctonus and Ips beetles captured in 2005 was inversely related to elevation of the trap site, and positively associated with the amount of ponderosa pine in the stand surrounding the site. Traps located closer to brood trees also captured more beetles. The relationship between trap catches and host tree mortality was weak and inconsistent in forest stands surrounding the funnel traps, suggesting that trap catches do not provide a good estimate of local beetle-induced tree mortality. However, pheromone-baited funnel trap data and data from gallery identification in bark samples produced statistically similar relative abundance profiles for the five species of bark beetles that we examined, indicating that funnel trap data provided a good assessment of species presence and relative abundance.     

Effectiveness of bifenthrin (Onyx) and carbaryl (Sevin SL) for protecting individual, high-value conifers from bark beetle attack (Coleoptera: Curculionidae: Scolytinae) in the Western United States

High-value trees, such as those located in residential, recreational, or administrative sites, are particularly susceptible to bark beetle (Coleoptera: Curculionidae: Scolytinae) attack as a result of increased amounts of stress associated with drought, soil compaction, mechanical injury, or vandalism. Tree losses in these unique environments generally have a substantial impact. The value of these individual trees, cost of removal, and loss of esthetics may justify protection until the main thrust of a bark beetle infestation subsides. This situation emphasizes the need for ensuring that effective insecticides are available for individual tree protection. In this study, we assess the efficacy of bifenthrin (Onyx) and carbaryl (Sevin SL) for protecting: ponderosa pine, Pinus ponderosa Dougl. ex. Laws., from western pine beetle, Dendroctonus brevicomis LeConte, in California; mountain pine beetle, Dendroctonus ponderosae Hopkins in South Dakota; and Ips spp. in Arizona; lodgepole pine, Pinus contorta Dougl. ex Loud., from D. ponderosae in Montana; pinyon, Pinus edulis Engelm. in Colorado and Pinus monophylla Torr. and Frem. in Nevada from pinyon ips, Ips confusus (LeConte); and Engelmann spruce, Picea engelmannii Parry ex. Engelm. from spruce beetle, Dendroctonus rufipennis (Kirby) in Utah. Few trees were attacked by Ips spp. in Arizona and that study was discontinued. Sevin SL (2.0%) was effective for protecting P. ponderosa, P. contorta, and P. monophylla for two field seasons. Estimates of efficacy could not be made during the second field season in P. edulis and P. engelmannii due to insufficient mortality in untreated, baited control trees. Two field seasons of efficacy was demonstrated in P. ponderosa/D. brevicomis and P. monophylla for 0.06% Onyx. We conclude that Onyx is an effective individual tree protection tool, but repeated annual applications may be required in some systems if multiyear control is desired.     

Monogamy in the bark beetle Ips latidens: ecological correlates of an unusual mating system

Abstract. 1. The diversity of mating systems in bark beetles (Coleoptera: Scolytidae) is poorly understood. Ips latidens (LeConte) is a small-bodied species of a harem polygynous genus whose mating system in nature had not been established previously. This study examines the breeding biology of I. latidens breeding in recently dead ponderosa pine ( Pinus ponderosa ) in California. It was expected that both the heterogeneous habitat quality inherent in dead trees and small body size would favour polygyny.
2. Contrary to expectations, I. latidens was found to be monogamous on the basis of excavations of 131 galleries conducted throughout the oviposition period at two wind-felled trees and at four sites composed of freshly-cut logs. Males left the galleries much sooner than females (50% gone by 24 and 36 days, respectively).
3. Individual females made up to four egg gallery arms extending from the male nuptial chamber, with the number of arms and the length of individual arms increasing with the age of the gallery. The relationship between total gallery length and gallery age was consistent among the trees and log sites, and did not depend on the presence of the male.
4. Ips latidens bred in a wide range of tree diameters (10–24 cm), and occurred in both pure aggregations and in aggregations with two other bark beetle species, Dendroctonus brevicomis and I. paraconfusus . The presence of the other species did not appear to influence negatively the density of I. latidens .
5. Monogamy in I. latidens may be due to lower male mortality during dispersal associated with a greater range of suitable breeding habitat than occurs for other Ips species, resulting in an even sex ratio at breeding sites.     

Attraction to monoterpenes and beetle‐produced compounds by syntopic Ips and Dendroctonus bark beetles and their predators

• 1 Bark beetles are significant mortality agents of conifers. Four beetle species, the pine engraver Ips pini, the six‐spined pine engraver Ips calligraphus sub. ponderosae, the southern pine beetle Dendroctonus frontalis, and the western pine beetle Dendroctonus brevicomis, cohabitate pines in Arizona.
• 2 A pheromone trapping study in ponderosa forests of Arizona determined the attraction of beetles to conspecific and heterospecific pheromone components in the presence and absence of host volatiles, and tested whether predators differ in their attraction to combinations of pheromone components and tree monoterpenes.
• 3 All four bark beetle species differed in their responses to heterospecific lures and monoterpenes. Ips calligraphus was the only species that increased in trap catches when heterospecific lures were added. Heterospecific lures did not inhibit the attraction of either Dendroctonus or Ips species. The replacement of myrcene with α‐pinene increased the attraction of Dendroctonus, whereas the addition of α‐pinene had mixed results for Ips. The prominent predators Temnochila chlorodia and Enoclerus lecontei were more attracted to the I. pini lure than the D. brevicomis lure, and the combination of the two lures with α‐pinene was most attractive to both predator species.
• 4 Cross attraction and limited inhibition of bark beetles to heterospecific pheromones suggest that some of these species might use heterospecific compounds to increase successful location and colonization of trees. Predator responses to treatments suggest that tree volatiles are used to locate potential prey and predators are more responsive to Ips than to Dendroctonus pheromone components in Arizona.

     

Survey and phylogenetic analysis of culturable microbes in the oral secretions of three bark beetle species

In a recent study, we reported a previously undescribed behavior in which a bark beetle exuded oral secretions containing bacteria that have antifungal properties, and hence defend their galleries against pervasive antagonistic Hyphomycete fungi. Actinobacteria, a group known for their antibiotic properties, were the most effective against fungi that invade the spruce beetle galleries. In the present study, we describe the isolation and identification of microorganisms from oral secretions of three bark beetles (Coleoptera: Curculionidae: Scolytinae): the spruce beetle, Dendroctonus rufipennis Kirby, the mountain pine beetle, Dendroctonus ponderosae Hopkins, and the pine engraver, Ips pini Say. Bacteria isolated from these three species span the major bacterial classes α-, β-, and γ-Proteobacteria, Firmicutes, Bacteroidetes, and Actinobacteria, except for D. ponderosae , which yielded no α-proteobacteria or Bacteroidetes isolates. Spruce beetles and pine engraver beetles had similar numbers of α-proteobacteria isolates, but pine engravers yielded twice as many Bacteroidetes isolates as spruce beetles. In contrast, mountain pine beetles yielded more isolates in the β- and γ-proteobacteria than spruce beetles and pine engravers. The highest percentage of Actinobacteria was obtained from spruce beetles, followed by pine engravers and mountain pine beetles. All of the fungal isolates obtained from the three beetle species were Ascomycetes. The greatest fungal diversity was obtained in spruce beetles, which had nine species, followed by pine engravers with five, and mountain pine beetles with one.     

外来入侵林业害虫强大小蠹的侵袭以及相关信息化学物质

强大小蠹是入侵我国并造成很大危害的鞘翅目棘胫小蠹科大小蠹属树皮低干森林害虫。它能侵害多种针叶树和松树 ,能被寄主挥发物和其它小蠹虫的外激素所引诱。该文综述了强大小蠹的侵袭习性以及寄主挥发物、其它小蠹虫的外激素对强大小蠹的引诱和驱避作用。     

Extreme ecological stoichiometry of a bark beetle–fungus mutualism

1. Ecological stoichiometry theory was applied to investigate how a consumer contends with an extreme elemental mismatch between its food and its body via symbiotic facilitation. 2. The beetle Dendroctonus brevicomis LeConte develops in bark, a substrate extremely low in nitrogen (N) and phosphorus (P). Its survival there depends on interactions with mutualist and antagonist fungi. 3. This study found that mutualists transfer N and P from sapwood and phloem into bark, where beetles feed, whereas the antagonist moves these elements only to phloem, resulting in starvation of the insect. However, even with mutualists, N and P concentrations remained low in bark, resulting in low N and extremely low P concentrations in the beetle. 4. The N:P ratios found in D. brevicomis larvae were the highest thus far reported for beetles and among the highest for insects and invertebrates. This suggests that the beetle has evolved additional, nutrient‐sparing adaptations.     

Trap type, chirality of alpha-pinene, and geographic region affect sampling efficiency of root and lower stem insects in pine.

Root and lower stem insects cause significant damage to conifers, vector phytopathogenic fungi, and can predispose trees to bark beetle attacks. The development of effective sampling techniques is an important component in managing these cryptic insects. We tested the effects of trap type and stereochemistry of alpha-pinene, in combination with ethanol, on catches of the root colonizing weevils (Coleoptera: Curculionidae) Hylobius spp. [mostly Hylobius pales (Herbst)], and Pachylobius picivorus (Germar), the root colonizing bark beetle (Coleoptera: Scolytidae) Hylastes porculus Erickson, and the lower stem colonizing bark beetle Dendroctonus valens (LeConte). We tested for inter-regional differences by conducting similar field assays in the northern (Wisconsin) and southern (Louisiana) United States. The more effective trap type varied with region. Root weevils were caught primarily in pitfall traps in Wisconsin, whereas they were caught mostly in lower stem flight traps in Louisiana. In Wisconsin, root colonizing bark beetles were also caught primarily in pitfall traps, but lower stem colonizing bark beetles were caught primarily in lower stem flight traps. The root feeding weevils preferred (-) over (+)-alpha-pinene in both regions. Some exceptions relating to trap type or gender occurred in southern populations. The two root and lower stem colonizing bark beetles in Wisconsin showed no preference between (+) and (-)-alpha-pinene in combination with ethanol. No bark beetles were caught in the south. Our results suggest that modifying trap type and enantiomeric ratios of monoterpenes for different insect groups and in different regions can improve sampling efficiency for these important pests.     

Drought and stand susceptibility to attacks by the European spruce bark beetle: A remote sensing approach

1. Several time-series analyses have demonstrated that after extreme summer drought bark beetle damage increased. However, studies predicting stand susceptibility over large spatial extents are limited by technical constraints in obtaining detailed, spatially-explicit data on infestation spot occurrence.
2. Using a unique dataset of georeferenced bark beetle infestation data, we tested whether the spatial variation of local growing conditions of forest stands, topography, and landscape variables modified the local occurrence of Ips typographus infestations after a severe hot drought in Central Europe.
3. Bark beetle infestation occurrence depended on soil-related aridity intensity, elevation, slope, and soil conditions. We showed that elevation interacted with growing conditions and topography. At low elevations, spruce forests growing on flat areas and wetter soils were more sensitive to the infestations. On the contrary, forests on steep slopes and soils with low water availability were rarely attacked. At the landscape scale, bark beetle damage increased with host tree cover but decreased with compositional diversity.
4. Our findings are generally consistent with the growth-differentiation balance hypothesis that predicts that trees growing under chronic dry conditions tend to be more resistant against biotic disturbances.
5. Spruce stands at low elevations located in homogeneous landscapes dominated by spruce were those more exposed to bark beetles in the initial phase of a drought-induced outbreak.

     

不同小蠹类与天牛类引诱剂联用对蛀干甲虫的林间诱捕效果

松墨天牛Monochamus alternatus Hope是传播松材线虫病的主要媒介,引诱剂是抑制松墨天牛种群数量的无公害调控有效方式之一。本研究通过在中国浙江富阳的林间试验测定了6种小蠹类引诱剂与1种天牛引诱剂不同组合联用对松墨天牛成虫以及其他蛀干害虫的诱捕效果。结果表明： 横坑切梢小蠹Tomicus minor引诱剂3-carene-10-ol、云杉八齿小蠹Ips typographus引诱剂2-methyl-3-buten-2-ol、欧洲榆小蠹Scolytus multistriatus引诱剂4-methyl-3-heptanol和纵坑切梢小蠹T. piniperda引诱剂verbenol分别与松墨天牛引诱剂联用后,对松墨天牛的引诱效果无显著性差异(P<0.05)。西部松大小蠹Dendroctonus brevicomis 引诱剂exo-brevicomin或黄杉大小蠹D. pseudotsugae引诱剂1-methylcyclohex-2-en-l-ol与松墨天牛引诱剂联用后对松墨天牛的诱捕效果有减弱的趋势。另外,与单独使用中国天牛引诱剂相比,小蠹类引诱剂与其联用后在诱捕其他蛀干害虫的种类和个体数量上均有所减少。     

Acetophenone superior to verbenone for reducing attraction of western pine beetle Dendroctonus brevicomis to its aggregation pheromone

1 The western pine beetle Dendroctonus brevicomis LeConte (Coleoptera: Scolytidae) is one of the most damaging insect pests of ponderosa pines Pinus ponderosa Douglas ex P. & C. Lawson in Western U.S.A. We compared the effect of verbenone, a well known bark beetle anti‐aggregation pheromone, with that of acetophenone on the attraction of D. brevicomis to its aggregation pheromone in a ponderosa pine forest in northern California. We tested the D. brevicomis aggregation pheromone alone and with three different release ratios of the aggregation pheromone (attractant) to verbenone or acetophenone (1 : 1, 1 : 2 and 1 : 5). 2 All treatments containing acetophenone or verbenone resulted in a significant reduction in the catch of D. brevicomis relative to the aggregation pheromone alone. When beetle responses to the three verbenone or three acetophenone treatments were pooled, the pooled verbenone treatment caught more D. brevicomis than the pooled acetophenone treatment. 3 There was no significant difference in the number of D. brevicomis caught among the three release rates of verbenone. By contrast, the 1 : 2 attractant : acetophenone ratio attracted significantly more D. brevicomis than the traps with the 1 : 5 attractant : acetophenone ratio. 4 Attraction of a major predator, Temnochila chlorodia (Mannerheim) (Coleoptera: Trogositidae), to the aggregation pheromone of D. brevicomis was reduced by verbenone, but not by acetophenone. Moreover, the T. chlorodia : D. brevicomis ratio for the pooled acetophenone treatment was 1.7‐fold greater than that for the attractant alone and two‐fold greater than the ratio for the pooled verbenone treatment, suggesting that acetophenone would not disrupt populations of this natural enemy. The importance of anti‐attractants in the biology of D. brevicomis and other bark beetles is discussed.     

Evaluation of insecticides for protecting Southwestern ponderosa pines from attack by engraver beetles (Coleoptera: Curculionidae: Scolytinae)

Insecticides that might protect pine trees from attack by engraver beetles (Ips spp.) have not been rigorously tested in the southwestern United States. We conducted two field experiments to evaluate the efficacy of several currently and potentially labeled preventative insecticides for protecting high-value ponderosa pine, Pinus ponderosa Dougl ex. Laws., from attack by engraver beetles. Preventative sprays (0.19% permethrin [Permethrin Plus C]; 0.03, 0.06, and 0.12% bifenthrin [Onyx]; and 1.0 and 2.0% carbaryl [Sevin SL] formulations) and systemic implants (0.875 g per capsule acephate [Acecap] and 0.650 g per capsule dinotefuran) were assessed on bolts (sections of logs) as a surrogate for live trees for a period of 13 mo posttreatment. The pine engraver, Ips pini (Say), was the most common bark beetle found attacking control and treated bolts, but sixspined ips, Ips calligraphus (Germar), and Ips lecontei Swain also were present. After approximately 13 mo posttreatment in one experiment, the spray treatments with 2.0% carbaryl, 0.19% permethrin, and 0.06 or 0.12% bifenthrin prevented Ips attack on the bolts at a protection level of > or = 70%. The acephate and dinotefuran systemic insecticides, and the 0.03% bifenthrin spray, provided inadequate (< or = 36%) protection in this experiment. For the other experiment, sprayed applications of 1.0% carbaryl, 0.19% permethrin, and 0.06% bifenthrin prevented beetle attack at protection levels of > or = 90, > or = 80, and > or = 70%, respectively, when bolts were exposed to Ips beetle attack for approximately 9-15 wk posttreatment. The sprays with 0.19% permethrin and 0.06% bifenthrin also provided > or = 90% protection when bolts were exposed for approximately 15-54 wk posttreatment. We concluded that under the conditions tested, 1.0 and 2.0% carbaryl, 0.19% permethrin, and 0.06 and 0.12% binfenthrin were acceptable preventative treatments for protecting ponderosa pine from successful engraver beetle attack for one entire flight season in the U.S. Southwest.     

Predation and bark beetle dynamics

Bark beetle populations may undergo dramatic fluctuations and are often important pests in coniferous forests. Their dynamics are thought to be primarily driven by factors affecting the resistance of the host tree to attack, i.e., bottom-up forces, while natural enemies are usually assigned a minor role in these systems. I present behavioral experiments that suggest that the clerid beetle Thanasimus dubius may be an important source of mortality for the bark beetle Dendroctonus frontalis during attack of the host tree, and determine the nature of the functional response of T. dubius under conditions close to natural. I also examine the numerical response of T. dubius to large-scale fluctuations in D. frontalis density, and the relationship between bark beetle population trends and predator density, and find that beetle populations tend to decline when predator densities are high. Combined with the effects of clerid larvae on bark beetle broods, these results suggest that top-down forces generated by natural enemies could also be an important component of bark beetle dynamics. The implications of these results for bark beetle dynamics are discussed in relation to the prolonged life-cycle of clerid beetles. Received: 23 January 1997 / Accepted: 5 April 1997     

Geographic variation in prey preference in bark beetle predators

Abstract 1. Bark beetles and their predators are useful systems for addressing questions concerning diet breadth and prey preference in arthropod natural enemies. These predators use bark beetle pheromones to locate their prey, and the response to different pheromones is a measure of prey preference. 2. Trapping experiments were conducted to examine geographic variation in the response to prey pheromones by two bark beetle predators, Thanasimus dubius and Temnochila virescens. The experiments used pheromones for several Dendroctonus and Ips prey species (frontalin, ipsdienol, and ipsenol) and manipulated visual cues involved in prey location (black vs. white traps). The study sites included regions where the frontalin‐emitter Dendroctonus frontalis was in outbreak vs. endemic or absent. 3. There was significant geographic variation in pheromone preference for T. dubius. This predator strongly preferred a pheromone (frontalin) associated with D. frontalis at outbreak sites, while preference was more even at endemic and absent sites. No geographic variation was found in the response by T. virescens. White traps caught fewer insects than black traps for both predators, suggesting that visual cues are also important in prey location. 4. The overall pattern for T. dubius is consistent with switching or optimal foraging theory, assuming D. frontalis is a higher quality prey than Ips. The two predator species partition the prey pheromones in areas where D. frontalis is abundant, possibly to minimise competition and intraguild predation.     

Integration of visual and olfactory cues of hosts and non-hosts by three bark beetles (Coleoptera: Scolytidae)

Abstract.  1. There has been a long-standing pre-occupation with how phytophagous insects use olfactory cues to discriminate hosts from non-hosts. Foragers, however, should use whatever cues are accurate and easily assessed, including visual cues.
2. It was hypothesised that three bark beetles, the mountain pine beetle (MPB), Dendroctonus ponderosae Hopkins, the Douglas-fir beetle (DFB), D. pseudotsugae Hopkins, and the western balsam bark beetle (WBBB), Dryocoetes confusus Swaine, integrate visual and olfactory information to avoid non-host angiosperms (e.g. paper birch, trembling aspen), that differ in visual and semiochemical profile from their respective host conifers (lodgepole pine, Douglas-fir, interior fir), and tested this hypothesis in a series of field trapping experiments.
3. All three species avoided attractant-baited, white (non-host simulating) multiple-funnel traps, and preferred attractant-baited black (host-simulating) traps. In experiments combining white, non-host traps with non-host angiosperm volatiles, bark beetles were repelled by these stimuli in an additive or redundant manner, confirming that these species could integrate visual and olfactory information to avoid non-host angiosperms while flying.
4. When antiaggregation pheromones were released from white traps, the DFB and MPB were repelled in an additive-redundant manner, suggesting that beetles can integrate diverse and potentially anomalous stimuli.
5. The MPB demonstrated the most consistent visual preferences, suggesting that it may be more of a 'visual specialist' than the DFB or WBBB, for which visual responses may be more contingent on olfactory inputs.     

Predators and competitors of the mountain pine beetle Dendroctonus ponderosae (Coleoptera: Curculionidae) in stands of changing forest composition associated with elevation

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Exogenous application of methyl jasmonate elicits defenses in Norway spruce (Picea abies) and reduces host colonization by the bark beetle Ips typographus

The terpenoid and phenolic constituents of conifers have been implicated in protecting trees from infestation by bark beetles and phytopathogenic fungi, but it has been difficult to prove these defensive roles under natural conditions. We used methyl jasmonate, a well-known inducer of plant defense responses, to manipulate the biochemistry and anatomy of mature Picea abies (Norway spruce) trees and to test their resistance to attack by Ips typographus (the spruce bark beetle). Bark sections of P. abies treated with methyl jasmonate had significantly less I. typographus colonization than bark sections in the controls and exhibited shorter parental galleries and fewer eggs had been deposited. The numbers of beetles that emerged and mean dry weight per beetle were also significantly lower in methyl jasmonate-treated bark. In addition, fewer beetles were attracted to conspecifics tunneling in methyl jasmonate-treated bark. Stem sections of P. abies treated with methyl jasmonate had an increased number of traumatic resin ducts and a higher concentration of terpenes than untreated sections, whereas the concentration of soluble phenolics did not differ between treatments. The increased amount of terpenoid resin present in methyl jasmonate-treated bark could be directly responsible for the observed decrease in I. typographus colonization and reproduction.