Antiaggregation Pheromone Research Articles

Elution rate and spacing of antiaggregation pheromone dispensers for protecting live trees from Dendroctonus pseudotsugae (Coleoptera: Scolytidae).

The antiaggregation pheromone 3-methylcyclohex-2-en-1-one (MCH) is highly effective in preventing the infestation of high-risk trees by Douglas-fir beetle, Dendroctonus pseudotsugae Hopkins. A large portion of the cost of an MCH treatment is related to the time applicators spend walking through an area dispersing the formulated pheromone. Application of fewer MCH dispensers eluting at a higher rate than those currently registered for operational use could potentially reduce treatment costs. Two higher elution rates, 6 and 18 mg/d per dispenser, were compared with the current standard of 2 mg/d per dispenser and an untreated control on 1-ha circular plots. Dispensers were spaced 5,15, and 44 m apart around the plot perimeters eluting 2, 6, and 18 mg/d, respectively. The nominal dose of MCH was 144 mg/ha/d on all plots. Percentages of Douglas-fir trees > or = 20 cm diameter at breast height mass attacked by Douglas-fir beetle were significantly lower on plots treated with dispensers eluting 2 and 6 mg/d and spaced 5 and 15 m apart, respectively, compared with the untreated control. Infestation rate on plots treated with dispensers eluting 18 mg/d and spaced 44 m apart was not significantly different from the control. Douglas-fir beetle abundance and host tree availability were similar on all plots. These results indicate that MCH dispensers eluting 6 mg/d (three times the current standard rate) and spaced 15 m apart (three times existing standard distance) can effectively prevent Douglas-fir beetle infestations.

Protection of lodgepole pines from mass attack by mountain pine beetle, Dendroctonus ponderosae, with nonhost angiosperm volatiles and verbenone

AbstractA number of angiosperm nonhost volatiles (NHVs) and green leaf volatiles (GLVs) were tested alone and as supplements to the antiaggregation pheromone, verbenone, for their ability to disrupt attack by the mountain pine beetle (MPB), Dendroctonus ponderosae Hopkins (Coleoptera: Scolytidae), on lodgepole pine, Pinus contorta Dougl. ex Loud. var. latifolia Engel. Preliminary experiments led to a refined NHV blend [benzyl alcohol, guaiacol, benzaldehyde, nonanal, salicylaldehyde, and conophthorin] and a refined GLV blend [(Z)‐3‐hexen‐1‐ol and (E)‐2‐hexen‐1‐ol]. In a 20‐replicate experiment, NHV and GLV groups both singly, and verbenone alone, significantly reduced MPB mass attack on pheromone‐baited trees and on trees within 5 m of the pheromone‐baited trees. Both blends in combination with verbenone reduced the number of mass attacked, baited trees to three out of 20 compared to 20 out of 20 of the baited controls. Each binary combination was also effective at reducing mass attack. In these experiments, all tested repellents were released from devices stapled to trees at the same point as the pheromone bait, suggesting that the repellency could have been to a point source, rather than to the whole tree. Therefore, in two further experiments bands of release devices were wrapped around the treated trees and the pheromone bait was removed from the treated trees. In one experiment, when the aggregation pheromone bait was suspended between pairs of trees treated with the NHV blend plus GLV blend plus verbenone, only three out of 25 treated pairs had mass attack on at least one member of the pair. In the other 60‐replicate experiment, with no pheromone baits present, attack occurred on 13 untreated and 11 banded trees, all in the path of a large advancing infestation. However, the mean attack density on the banded trees was significantly reduced to a level below the 40 attacks m−2 of bark surface required to kill a healthy lodgepole pine. As a result of these experiments, operational trials are recommended.

Angiosperm bark volatiles disrupt response of Douglas-fir beetle, Dendroctonus pseudotsugae, to attractant-baited traps.

Antennally active, bark-derived, angiosperm volatiles were tested singly and in groups for their ability to disrupt the response of the Douglas-fir beetle (DFB), Dendroctonus pseudotsugae, to attractant-baited multiple-funnel traps. One compound, conophthorin, was active alone in reducing the response of beetles to the baited traps. Further experiments showed disruptive activity in two aliphatic green-leaf alcohols [1-hexanol and (Z)-3-hexen-1-ol], as well as guaiacol and benzyl alcohol, and three aliphatic aldehydes [nonanal, hexanal, and (E)-2-hexenal] but not in two aromatic aldehydes (benzaldehyde and salicylaldehyde). Every binary combination that included conophthorin or any two of the other groups, except aromatic aldehydes, significantly reduced the response of beetles to baited traps. Various ternary mixtures and the complete mixture of all the groups were generally the most effective treatments. These results provide evidence that DFBs recognize and avoid nonhosts while flying rather than landing on candidate hosts and testing them while in contact with the tree. Nonhost angiosperm bark volatiles may have practical utility on their own or in combination with the antiaggregation pheromone 3-methylcyclohex-3-en-1-one (MCH) to protect single trees, logs, or stands from attack by the DFB.

VISUAL AND SEMIOCHEMICAL DISRUPTION OF HOST FINDING IN THE SOUTHERN PINE BEETLE

The importance of visual silhouettes for host finding by the southern pine beetle (SPB), Dendroctonus frontalis (Coleoptera: Scolytidae), and the potential for disruption of this process using visual deterrents were evaluated with multiple-funnel traps, painted white or black, and with clear, white, or black Plexiglas sticky panels. All traps and panels were baited with the SPB attractant frontalure. The effect of combined semiochemical and visual disruption was evaluated in funnel traps by including the antiaggregation pheromone verbenone, or the repellent/inhibitory host compound 4-allylanisole, in addition to the attractant. Visual treatments had a highly significant effect on catch of SPB and the predatory clerid beetle Thanasimus dubius. In attractant-baited traps, white paint alone reduced the average number of SPB caught by 72% in Florida and 68% in Louisiana. The repellent 4-allylanisole reduced catch of SPB by 56% in Florida and 45% in Louisiana. Verbenone was eluted at 25% of the targeted rate a...

Volatiles from the bark of trembling aspen, Populus tremuloides Michx. (Salicaceae) disrupt secondary attraction by the mountain pine beetle, Dendroctonus ponderosae Hopkins (Coleoptera: Scolytidae)

Coupled gas chromatographic-electroantennographic detection (GC-EAD) analysis of the Porapak Q-captured volatiles from the bark of trembling aspen, Populus tremuloides Michx., revealed four compounds that consistently elicited antennal responses by mountain pine beetles (MPBs), Dendroctonus ponderosae Hopkins. One of these, 1-hexanol, disrupted the capture of MPBs in multiple-funnel traps baited with the aggregation pheromones trans-verbenol and exo-brevicomin and the host kairomone myrcene, a blend of semiochemicals that mediates the secondary attraction response in which beetles mass attack and kill living pines. The other three EAD-active aspen bark volatiles, benzyl alcohol, benzaldehyde and nonanal, were inactive alone, but in binary and ternary combinations contributed to a disruptive effect in an additive and redundant manner when all four aspen bark volatiles were tested in all possible binary and ternary blends. The best ternary blend and the quarternary blend achieved ≥ 80% disruption. The quarternary blend enhanced the disruptive effect of the antiaggregation pheromone verbenone in traps, raising the disruptive effect to 98%, and also enhanced the inhibition of attack on attractant-baited lodgepole pines. This is the first demonstration of specific compounds from the bark of angiosperm trees that disrupt the secondary attraction response of sympatric coniferophagous bark beetles. The results support the hypothesis that such bark beetles are adapted to recognize and avoid non-host angiosperm trees by responding to a broad spectrum of volatiles that can act in various blends with equal effect.

Using Aggregation and Antiaggregation Pheromones of the Douglas-Fir Beetle to Produce Snags for Wildlife Habitat

Abstract Pheromone baits for the Douglas-fir beetle were applied alone and in combination with the antiaggregation pheromone, 3-methylcyclohex-2-en-1-one (MCH), to test potential treatments for creating snags to improve wildlife habitat. All baited trees were attacked by beetles following both treatments. However, more unbaited trees were attacked on plots without MCH than on plots with MCH. One year after treatment, more of the attacked trees were dead on plots without MCH than on plots with MCH. Bark beetle pheromones can be used effectively to create snags for improving wildlife habitat, but to achieve specific objectives treatments must be prescribed based on knowledge of the expected beetle population densities. West. J. Appl. For. 12(2):00-00.

Optimal Dose of an Antiaggregation Pheromone (3-Methylcyclohex-2-en-1-one) for Protecting Live Douglas-Fir from Attack by Dendroctonus pseudotsugae (Coleopera: Scolytidae)

The Douglas-fir beetle, Dendroctonus pseudotsugae Hopkins, anti aggregation pheromone, 3-methylcyclohex-2-en-1-one (MCH), was applied to stands of Douglas-fir, Pseudotsuga menziesii (Mirabel) Franco, at high risk for infestation to determine the lowest effective dose for protecting live trees. MCH was applied at rates of 50, 100, and 150 bubble capsules per hectare (20, 40, and 60 g/ha, respectively) in 1994, and 15, 30, and 50 bubble capsules per hectare (6, 12, and 20 g/ha, respectively) in 1995. Mean release rates throughout the beetle flight periods in 1994 and 1995 were 1.63 and 1.23 milligrams per capsule per day, respectively. For both years, catches of Douglas-fir beetles in pheromone-baited traps located at the plot centers were significantly lower on all MCH-treated plots compared with untreated plots, but there were no differences among the 3 doses of MCH. In contrast, NCR had no effect on the numbers of 3 predators [Thanasimus undatulus (Say), Enoclerus sphegeus F., and Temnochila chlorodia (Mannerheim)] collected in the traps during either year. In 1994, the percentage of Douglas-fir ≥20 cm diameter at breast height (abh) that were mass attacked was significantly lower on MCH-treated plots compared with the untreated control, and all 3 doses were equally effective. In 1995, there were no significant differences in the percentage of mass-attacked trees among the treatments. These results demonstrate that MCH applied at rates as low as 20 g/ha (50 bubble capsules per hectare) can reduce the probability that high-risk Douglas-fir will become infested.

Green leaf volatiles as antiaggregants for the mountain pine beetle,Dendroctonus ponderosae Hopkins (Coleoptera: Scolytidae)

We tested the hypothesis that green leaf volatiles act as antiaggregants for the mountain pine beetle (MPB),Dendroctonus ponderosac Hopkins. In coupled gas chromatographic-electroantennographic detection (GC-EAD) analysis MPB antennae responded to 30 ng doses of all six-carbon green leaf alcohols tested [1-hexanol, (E)-2-hexen-1-ol, (Z)-2-hexen-1-ol, (E)-3-hexen-1-ol, and (Z)-3-hexen-1-ol], but not to the aldehydes, hexanal or (E)-2-hexenal, or to alcohol or aldehyde homologues with more or fewer than six carbon atoms. In field trapping experiments a blend of green leaf alcohols [1-hexanol, (Z)-2-hexen-1-ol, (E)-3-hexen-1-ol and (Z)-3-hexen-1-ol] effectively disrupted the response to attractive semiochemicals; a blend of the aldehydes hexanal and (E)-2-hexenal was inactive. The two best disruptants. (E)-2-hexen-1-ol and (Z)-3-hexen-1-ol, reduced catches of both sexes to levels not significantly different from catches in unbaited control traps. They also reduced the attack on trees baited with attractive MBP pheromones to a level not significantly different from that on unbaited control trees. Neither of the clerid predators captured,Enoclerus sphegeus (F.) norThanasimus undatulus (Say), was repelled by green leaf volatiles. Our results suggest that green leaf alcohols are promising disruptants which may be used to supplement the antiaggregation pheromone, verbenone, in protecting single high-value trees as well as carefully selected stands with low-level populations of MPBs.

Effect of an Inhibitor-Based Suppression Tactic on Abundance and Distribution of the Southern Pine Beetle (Coleoptera: Scolytidae) and Its Natural Enemies

An inhibitor-based tactic can suppress southern pine beetle, Dendroctonus frontalis Zimmermann, infestations using the anti-aggregation pheromone, verbenone. However, it is unknown whether southern pine beetle and its insect natural enemies are impacted by this tactic. We evaluated potential effects by examining mean trap catches, aggregation measures, species diversity and abundance indices, and interspecific covariation between southern pine beetle and its most common insect enemies. Examination of flight and landing trap catches, before and after application of verbenone to infestations, showed that southern pine beetle and its natural enemies remained at the sites at similar densities, continued their associations, and maintained similar spatial distributions. This suggests that verbenone reduces successful colonization of unattacked trees and has no negative effect on the relationship of southern pine beetle with its natural enemies.

EFFICACY OF AN ANTIAGGREGATION PHEROMONE FOR REDUCING DOUGLAS-FIR BEETLE, DENDROCTONUS PSEUDOTSUGAE HOPKINS (COLEOPTERA: SCOLYTIDAE), INFESTATION IN HIGH RISK STANDS

AbstractThe Douglas-fir beetle (Dendroctonus pseudotsugae) antiaggregation pheromone, 3-methylcyclohex-2-en-1-one (MCH), was applied to stands at high risk for infestation. The MCH was applied in a bubble capsule formulation to plots ranging from 2.1 to 2.6 ha in size at rates of 45–76 g/ha. Catches of Douglas-fir beetles in attractant-baited Lindgren funnel traps located at the plot centers were significantly lower on MCH-treated plots compared with untreated plots. In contrast, catches of the most abundant predator, Thanasimus undatulus (Say), were unaffected by the MCH treatment. The percentage of Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco] trees ≥20 cm dbh that were mass attacked was significantly lower on treated plots (0.2%) compared with untreated plots (8.5%). MCH alone was effective in reducing the probability of Douglas-fir beetle infestations occurring in high risk stands.

Reduction of Douglas-fir beetle infestation of high-risk stands by antiaggregation and aggregation pheromones

A combination of antiaggregation and aggregation pheromones was tested for protecting stands of Douglas-fir (Pseudotsugamenziesii (Mirb.) Franco) at high risk for infestation by the Douglas-fir beetle (Dendroctonuspseudotsugae Hopkins). The antiaggregation pheromone, 3-methylcyclohex-2-en-1-one (MCH), was applied in a bubble capsule formulation to the perimeter of 1-ha circular plots at a rate of 60 g/plot. Treated plots also had three or four clusters of four Lindgren funnel traps baited with frontalin, seudenol, 1-methylcyclohex-2-en-1-ol, and ethanol located outside of the plot but within 160 m of the boundary. Mean (±SE) accumulated catches in all traps per plot were 73 658 ± 19 721 Douglas-fir beetles and 12 892 ± 2 513 Thanasimusundatulus (Say), a predator of the Douglas-fir beetle. The mean percentage of Douglas-fir trees ≥20 cm DBH that were mass attacked was reduced by 80% within the treated plots compared with the untreated plots. However, there was an eightfold increase in the percentage of trees mass attacked in the area outside the treated plots in the vicinity of the funnel traps. The net effect of the treatment was to concentrate mass-attacked trees within a limited area outside of the protected stand. Our results indicate that Douglas-fir beetle antiaggregation and aggregation pheromones can be used effectively to reduce the probability of infestation in small, high-value stands.

Photoisomerization of antiaggregation pheromone verbenone: Biological and practical implications with respect to the mountain pine beetle,Dendroctonus ponderosae Hopkins (Coleoptera: Scolytidae)

Release of the antiaggregation pheromone, verbenone, at 3.8 mg/ day from a concentrated source within a multiple-funnel trap completely inhibited response by the mountain pine beetle (MPB),Dendroctonus ponderosae Hopkins, to attractive semiochemical lures. When aerial applications were simulated and verbenone was released at the same rate from beads lying in a 2×2-m area on the forest floor 15-35 cm below a trap, the response of the MPB was inhibited by only 50%. This reduced inhibition may be explained in part by the photoisomerism of verbenone. When exposed to full sunlight on two occasions, the times required for 50% of verbenone vapors to be converted to chrysanthenone were 75 and 100 min, respectively. Trap and tree-baiting experiments indicated no biological activity of chrysanthenone. Rapid photoisomerization could reduce the concentration of verbenone below biologically active levels and would allow the MPB to colonize trees close to already occupied hosts, contributing to the characteristic clumped distribution of MPB attack. The rate of verbenone photoisomerization may vary according to geographic location, stand elevation and density, and should be considered before verbenone is applied to control the MPB and other bark beetles.

A simplified tree bait for the mountain pine beetle

Seven, split-block experiments throughout British Columbia in 1989 tested the efficacy of binary tree baits containing the pheromones trans-verbenol and exo-brevicomin or ternary baits with the addition of the host tree kairomone myrcene for containing and concentrating infestations of the mountain pine beetle, Dendroctonusponderosae Hopkins, in stands of lodgepole pine, Pinuscontorta var. latifolia Engelm. Attack densities on baited trees, attack frequencies of baited trees and trees within 10 m of the baited trees, and the ratios of newly attacked, green, trees to previously attacked, red, trees were generally statistically equal between sub-blocks containing binary or ternary baits. Where statistically significant differences occurred for one or more of the above criteria in one experiment, they were generally offset by statistically significant differences in the opposite direction in another experiment. Two individual-tree experiments in 1990 that supported the equality of binary and ternary baits indicated that raising the release rate of trans-verbenol in binary baits tended to reduce their efficacy (possibly because of contamination with the antiaggregation pheromone verbenone) and showed that increasing the release rate of exo-brevicomin tended to counteract this effect. When attack frequencies were subdivided by diameter class of available trees attacked, all baits were effective in inducing attack on available trees <30.0 cm diameter at breast height (1.3 m), but attack on baited and control trees ≥30 cm diameter at breast height was equal. Provided that the trans-verbenol in binary baits does not contain or autoxidize to verbenone, myrcene can be deleted from operational tree baits.

Displacement and aggregation of mountain pine beetles, Dendroctonusponderosae (Coleoptera: Scolytidae), in response to their antiaggregation and aggregation pheromones

Deployment of verbenone release devices at 10-m centres significantly reduced the percent of available lodgepole pines (Pinuscontorta Dougl. var. latifolia Engelm.) that were mass attacked by mountain pine beetles (Dendroctonusponderosae Hopkins) in 50 × 150 m subplots flanked on each side by identical subplots, regardless of whether the flanking plots were baited with tree baits or left untreated. Verbenone treatment also increased the percent of unsuccessful attacks (<31.25 attacks/m2) in the central subplot compared with when the central subplot was left untreated. Tree baits, composed of trans-verbenol, exo-brevicomin, and myrcene, applied in flanking subplots did not affect significantly the percent of trees mass attacked nor the attack density. However, assuming a 2:1 expected ratio of attacks in the two flanking subplots over the central subplot, χ2-tests indicated that the attack distribution was consistently altered only when both verbenone and aggregation pheromones were applied. This study suggests that a "push–pull" tactic using aggregation and antiaggregation pheromones may be feasible for managing this destructive pest.

The response of mountain pine beetle (Dendroctonus ponderosae) to lodgepole pine trees baited with verbenone and exo-brevicomin

exo-brevicomin, a multifunctional pheromone of the mountain pine beetle,Dendroctonus ponderosae, was tested at release rates of 0.5 and 2.5 mg/day alone and in combination with the antiaggregation pheromone verbenone against unbaited controls. Significantly more lodgepole pinePinus contorta var.latifolia trees were attacked, and at higher densities, with both release rates ofexo-brevicomin than with all other treatments. Verbenone significantly reduced the response of mountain pine beetles toexo-brevicomin. Verbenone alone did not reduce the number of trees attacked by mountain pine beetle or the attack density when compared to the unbaited controls.

Aerial application of verbenone reduces attack of lodgepole pine by mountain pine beetle

Mountain pine beetle, Dendroctonusponderosae Hopkins, is the primary pest affecting lodgepole pine, Pinuscontorta var. latifolia Engelm., ecosystems in western North America. In 1988, aerial treatments of the antiaggregation pheromone, verbenone, were applied to lodgepole pine stands infested with mountain pine beetle in northwestern Montana. The pheromone was formulated by PHERO TECH Inc. in controlled-release, cylindrical 5 × 5 mm plastic beads and applied without benefit of a sticker at the rate of 54 g verbenone per hectare. There were significantly fewer successfully attacked trees on the treated plots, as evidenced by (i) a fourfold greater incidence of current-year attacked trees per hectare in the untreated check plots and (ii) the significantly lower (α = 0.05) ratio of 1988:1987 attacked trees in the treated plots. Further, the number of trees per hectare resisting attacks (as reflected by number of trees pitching out bark beetles) was higher (α = 0.05) in the treated plots. More pitch outs occurred in treated plots presumably because avoidance of verbenone by beetles reduced the number of beetles below that needed to overcome the natural resistance of attacked trees.

ENDO-BREVICOMIN: AN ANTIAGGREGATION PHEROMONE FOR THE WESTERN BALSAM BARK BEETLE, DRYOCOETES CONFUSUS SWAINE (COLEOPTERA: SCOLYTIDAE)

AbstractLaboratory bioassays and field tests demonstrated that the (+) enantiomer of endo-brevicomin is an antiaggregation pheromone for the western balsam bark beetle, Dryocoetes confusus Swaine. (+)-endo-Brevicomin significantly reduced the response of beetles of both sexes to the aggregation pheromone exo-brevicomin in laboratory bioassays and in field experiments using multiple-funnel traps. It also prevented attack on subalpine fir, Abies lasiocarpa (Hook) Nutt., trees baited with exo-brevicomin. The (−) enantiomer of endo-brevicomin was inactive. These results suggest that (+) or (±)-endo-brevicomin has potential for protecting selected trees from attack, or suppressing western balsam bark beetle activity in high hazard or infested stands.

Semiochemicals and bark beetle populations: Exploitation of natural phenomena by pest management strategists

An analysis of semiochemical communication between host trees, bark beetles and commensal or entomophagous insects discloses five principal means by which semiochemicals can influence the population dynamics of bark beetles. These are: mediation of aggregation and mass attack on new hosts, cessation of aggregation and shifting of attack to uninhabited hosts, induction of aggregation by competing species, inhibition of aggregation by competing species, and mediation of host finding by commensal and entomophagous insects. Further analysis suggests major points of natural vulnerability which lead to six fundamental strategies for potential pest management: prevention of production of aggregation pheromones, sabotage of olfactory perception, exploitation of semiochemical‐based secondary attraction, exploitation of antiaggregation pheromones, exploitation of repellent allomones, and exploitation of the kairomonal response by entomophagous insects. Investigations of the many possible tactics arising from these strategies have led to three types of operational pest management programs: prevention of pheromone production by excluding bark beetles from their hosts; suppression of bark beetle populations through the utilization of semiochemical‐baited traps, trees or logs; and the use of antiaggregation pheromones to protect vulnerable hosts from attack.

The role of autoxidation of α-pinene in the production of pheromones of Dendroctonusponderosae (Coleoptera: Scolytidae)

The monoterpene α-pinene, a major component of the terpene composition of Pinus spp., has been reported to act as a host-produced kairomone for a variety of bark beetle species, including the mountain pine beetle, Dendroctonusponderosae Hopkins. However, our experiments indicate that α-pinene autoxidizes under normal temperature and atmospheric conditions to form significant quantities of trans-verbenol, an aggregation pheromone for many species of bark beetles. The quantities of α-pinene present in the resin that can flow from small wounds in pine trees appear to be sufficient to produce trans-verbenol at rates similar to those by female beetles that are actively synthesizing the compound.trans-Verbenol can then autoxidize rapidly to form verbenone, with the content of this compound reaching 8% within 13 weeks of exposure to air. Verbenone is often used by scolytids as an antiaggregation pheromone. Approximately 1.9% of the trans-verbenol and 2.7% of the verbenone found in Porapak Q aerations of phloem with boring spruce beetle, Dendroctonusrufipennis (Kirby), females, as well as 0.8% of the trans-verbenol and 0.8% of the verbenone found in aerations of phloem with boring D. ponderosae females, was due to the autoxidation of α-pinene and (or) the release of oxygenated compounds found in the phloem before bark beetle attack. The natural interconversion of α-pinene, trans-verbenol, and verbenone under ambient conditions suggests that many experiments involving the behavioral activity of these compounds require re-evaluation.

Terpene alcohol pheromone production byDendroctonus ponderosae andIps paraconfusus (Coleoptera: Scolytidae) in the absence of readily culturable microorganisms

Dendroctonus ponderosae Hopkins andIps paraconfusus Lanier of both sexes produced most of their complement of terpene alcohols at normal to elevated levels in the absence of readily culturable microorganisms. However, there was some evidence that microbial involvement may be required by maleI. paraconfusus to produce ipsenol and ipsdienol at normal levels. Increased levels of certain terpene alcohols found in axenically reared or streptomycin-fed beetles suggest that symbiotic microorganisms may be responsible for breaking down pheromones and other terpene alcohols. There was also evidence for microbial involvement in the production of the antiaggregation pheromone verbenone inD. ponderosae. This compound was not produced in quantifiable levels by axenically reared or streptomycin-fed beetles exposed to α-pinene as vapors or through feeding, but was found in wildD. ponderosae exposed to α-pinene through feeding on bolts of lodgepole pine,Pinus contorta var.latifolia Engelmann.