Interspecific inhibition of the response of the bark beetles,Dendroctonus brevicomis andIps paraconfusus,to their pheromones in the field

Ponderosa pine logs infested withIps paraconfusus males inhibited the attraction ofDendroctonus brevicomis in the field to either attractive logs cut from a ponderosa pine tree under attack byD. brevicomis or to their synthetic pheromones,exo-brevicomin, frontalin, and myrcene. Logs cut from trees under attack byD. brevicomis inhibited the response ofI. paraconfusus to logs infested with maleI. paraconfusus.Exo-brevicomin, frontalin, and myrcene did not inhibit their response but verbenone did. Verbenone was found in maleD. brevicomis dissected from attractive logs under attack during the same time the response ofI. paraconfusus was inhibited by these logs.Trans-verbenol andexo-brevicomin were found in femaleD. brevicomis while verbenone,trans-verbenol, and frontalin were found in maleD. brevicomis in relatively large amounts near the beginning of the aggregation phase of host colonization. All of these compounds had decreased at a similar rate 5 days later. This gradual decrease inexo-brevicomin and frontalin probably caused the observed reduction in attraction. The ecological significance of these compounds in relation to termination of the aggregation phase ofD. brevicomis and reduction of interspecific competition is discussed.Coleoptera: Scolytidae. This paper is based in part upon a dissertation submitted by J.A.B. to the University of California, Berkeley, in partial fulfillment of the requirements for the PhD in entomology. These studies were supported in part by the Rockefeller Foundation; U.S. Forest Service; Regional research Project W-110, S.E.A./ U.S.D.A.; and the National Science Foundation and Environmental Protection Agency through a grant (NSF GB-34719/BMS 75-04223) to the University of California.     

Interspecific effects of pheromones on the attraction of the bark beetles,Dendroctonus brevicomis andIps paraconfusus in the laboratory

Dendroctonus brevicomis was attracted to a mixture of theIps paraconfusus pheromones, ipsenol,cis-verbenol, and ipsdienol at 10^–9 g each/l but was not attracted to these pheromones at higher and lower release rates.I. paraconfusus was not attracted to theD. brevicomis pheromonesexo-brevicomin, frontalin, and myrcene at any release rate tested. Increased release rates of a mixture of the three pheromones ofI. paraconfusus inhibited the attraction ofD. brevicomis to its synthetic pheromones. A mixture of ipsenol + ipsdienol orcis-verbenol alone failed to cause inhibition indicating that at least two of theI. paraconfusus pheromones are required to inhibit the response ofD. brevicomis. The pheromones ofD. brevicomis did not inhibit the attraction ofI. paraconfusus to its pheromones; however, verbenone was a potent inhibitor.Coleoptera: Scolytidae. This paper is based in part upon a dissertation submitted by J.A.B. to the University of California, Berkeley, in partial fulfillment of the requirements for the Ph.D. degree in entomology, June 1978. These studies were supported by the Rockefeller Foundation; USDA Forest Service; Regional research project W-1 10, SEA/ USDA; and the National Science Foundation and Environmental Protection Agency through a grant (NSF GB-34719/BMS 75-04223) to the University of California. The findings, opinions, and recommendations are not necessarily those of the University of California or the funding agencies.     

Field responses of the western pine beetle1 and one of its predators to host- and beetle-produced compounds

The responses of the western pine beetle (Dendroctonus brev-icomis LeConte) andTemnochila chlorodia (Mannerheim) to candidate attractants—exo- andendo-brevicomm, frontalin,trans-verbenol, ver-benone, and ponderosa pine turpentine and its major monoterpene components—were quantified by counts of beetles on traps baited with the various attractants, singly and in combinations released simultaneously. Combinations ofexo-brevicomin and frontalin plus a monoterpene or turpentine were the most attractive toD. brevicomis. The responses to these attractant combinations were reduced when verbenone plustrans-verbenol were present. All single compounds and binary mixtures, exceptexo-brevicomin plus frontalin, were much less attractive.exo-Brevicomin was most attractive toT. chlorodia, and this response appeared to decrease when verbenone plustrans-verbenol were present.Coleoptera: Scolytidae.Coleoptera: Trogositidae.This research was supported in part by the Forest Service, U.S. Department of Agriculture, and in part by grants from the Rockefeller Foundation to the University of California. Commercial enterprises and products are mentioned solely for information, and do not imply endorsement by the sponsoring agencies and organizations.     

Olfactory receptor cell responses ofIps grandicollis (eichhoff) (Coleoptera: Scolytidae) to intra- and interspecific behavioral chemicals

Electrophysiological recordings from antennal olfactory receptor cells were obtained fromIps grandicollis. Recordings were made from olfactory receptor cells from nine regions of the antennae in response to stimulation with the semiochemicals-pinene, frontalin,endo-brevicomin, verbenone,trans-verbenol,cis-verbenol, ipsdienol, and ipsenol. In many cases, up to two cells were recorded concurrently from the same location. When compared to males, females had a greater percentage of cells responsive to the primary pheromones ofDendroctonus frontalis, frontalin andtrans-verbenol, and ofIps spp., ipsdienol and ipsenol. Among females, more cells responded totrans-verbenol and theIps-produced volatiles than to host or otherD. frontalis-produced compounds. Olfactory cells of males responded mostly tocis-verbenol, followed by-pinene, verbenone,trans-verbenol, andendo-brevicomin. Of those cells responsive primarily to one compound, the greatest percentage were responsive totrans-verbenol in females and to verbenone in males. The response of the antennal olfactory receptor cells to semiochemicals used by male and femaleI. grandicollis is consistent with the presence of these compounds during the host colonization period for each sex. Our results, which show a lack of specificity in most pheromone and host odor receptor cells, is in contrast with previously published accounts of olfactory receptor cell specificity in otherIps species.     

Electroantennogram responses by mountain pine beetles,Dendroctonus ponderosae Hopkins,exposed to selected semiochemicals

Electroantennograms (EAGs) were obtained forD. ponderosae to the bark beetle pheromonestrans-verbenol,cis-verbenol,exo-brevicomin,endo-brevicomin, frontalin, verbenone; to the kairomones, -pinene, -pinene, camphene, -3-carene, limonene, myrcene; to a blend (111) oftrans-verbenol,exo-brevicomin, and myrcene; and to diacetone-alcohol. Male and female responses, in general, did not differ significantly over the whole EAG dose-response curves but differed at a few concentrations on many of the curves. There were more differences noted for pheromones than kairomones. The blend yielded among the largest EAGs in both sexes and appeared to show synergism. Responses of females were lower than those for males in most instances. Significant differences in responses by the two sexes were much fewer for the kairomones than the pheromones. EAG recovery rates tested at only one concentration showed significant differences between males and females for three pheromones,trans-verbenol,cis-verbenol, and verbenone, and two kairomones, camphene and -3-carene. Thresholds were quite low for most of the odorants exceptcis-verbenol, camphene, verbenone, and diacetone-alcohol in females, andcis-verbenol, verbenone, -pinene, and diacetone-alcohol in males. The results, using at least one EAG parameter, support behavioral and field studies involvingexo-brevicomin,trans-verbenol, frontalin, and the blend.     

Effects of verbenone andtrans-verbenol on the response ofDendroctonus brevicomis to natural and synthetic attractant in the field

Verbenone andtrans-verbenol were investigated as candidate interruptants for use as tree protectants. Verbenone andtrans-verbenol, pheromones released byDendroctonus brevicomis during host colonization, reduced the trap catch ofD. brevicomis near sources of the attractant composed ofexo-brevicomm, frontalin, and myrcene. Catch reduction at some trap positions was greater at a high release rate than at a low release rate oftrans-verbenol alone and of the combination of verbenone plustrans-verbenol. Verbenone also reduced catches at traps baited with attractive bolts from trees under attack byD. brevicomis. Attempts to use verbenone to protect living trees fromD. brevicomis attack were inconclusive.Coleoptera: Scolytidae.This research was supported in part by the Forest Service, U.S. Department of Agriculture, and in part by grants from the National Science Foundation/Environmental Protection Agency (grant NSF-6B-34718/BMS75-04223), and in part by grants from the Rockefeller Foundation to the University of California. Mention of a proprietary or commercial product does not constitute recommendation or endorsement of the product by the U.S. Department of Agriculture and does not imply its approval to the exclusion of other products that also may may be suitable.     

Male-specific conversion of the host plant compound,myrcene, to the pheromone, (+)-ipsdienol,in the bark beetle,Dendroctonus brevicomis

When both sexes of the bark beetle,Dendroctonus brevicomis LeConte, were exposed to vapors of myrcene from ponderosa pine, only the male produced (+)-ipsdienol. In the field, racemic ipsdienol significantly reduced the attraction of both sexes in flight to a mixture of myrcene and the aggregation pheromones,exo-brevicomin and frontalin. This suggests that ipsdienol may be involved in regulating colonization density ofD. brevicomis. The implications of the biosynthesis of various enantiomers of ipsdienol byD. brevicomis and the cohabitating bark beetles,Ips paraconfusus andI. pini, in relation to their behavioral responses are discussed in regard to reducing interspecific competition.Coleoptera: Scolytidae.     

Response ofDendroctonus brevicomis Le Conte (Coleoptera: Scolytidae) to different release rates and ratios of aggregation semiochemicals and the inhibitors verbenone and ipsdienol

The response of western pine beetle,Dendroctonus brevicomis Le Conte, to different release rates of the aggregation semiochemicals (attractants)exo-brevicomin, frontalin, and myrcene and the inhibitors verbenone and ipsdienol was investigated. Release rates of verbenone ranging from 0.18 mg/ 24 hr to 1.2 mg/24 hr did not result in significant reductions in mean trap catch ofD. brevicomis. In contrast, very low release rates of ipsdienol (0.02–0.4 mg/24 hr) significantly reduced trap catch compared to controls. The combination of verbenone and ipsdienol, released at rates above 0.09 and 0.02 mg/24 hr, respectively, resulted in significantly lower trap catches ofD. brevicomis in attractant-baited traps. Results of an experiment testing a factorial combination of different release rates of verbenone and attractants suggest that response is not ratio-specific. The response curve ofD. brevicomis to the levels of verbenone was similar across all levels of attractants, while the response to equivalent ratios of attractants to verbenone was not similar, suggesting that the behavior of the beetles is primarily influenced by the absolute release rate of verbenone.     

Electroantennograms by mountain pine beetles,Dendroctonus ponderosae Hopkins,exposed to selected chiral semiochemicals

Electroantennograms (EAGs) were recorded fromD. ponderosae to the enantiomers of the terpenoid bark-beetle pheromonestrans-verbenol,cis-verbenol, verbenone, and the bicyclic ketals frontalin,exo-brevicomin, andendo-brevicomin. Male and female responses to enantiomers of the terpenoids differed significantly only at the two highest concentrations. No sex differences were seen in response to the bicyclic ketals. Significantly different responses to the enantiomers of all the chemicals, except frontalin, were noted over at least part of the dosage-response ranges tested. The negative antipode for all of the terpenoids elicited higher responses, while for the bicyclic ketals, the positive antipode effected the largest responses except for the two highest concentrations ofexo-brevicomin.     

Frontalin in the male mountain pine beetle

Frontalin andexo-brevicomin were identified by GC-MS in air drawn over maleDendroclonus ponderosae Hopk. (MPB) from Oregon that had joined females for 1–2 days in the bark of lodgepole and ponderosa pine logs. Unfed males releasedexo- andendo-brevicomin but not frontalin. These three compounds were not detected in either unfed or fed females. Arrestment of males bytrans-verbenol and terpenes in an olfactory walkway was reduced by the addition of racemic frontalin; production of attractant chirps also diminished. Racemic frontalin also strongly reduced the aggregation of MPB in lodgepole and ponderosa pine stands to sticky traps baited with the aggregation pheromonetrans-verbenol and host terpenes; however, the function of the natural enantiomer of frontalin in MPB is unknown.Research supported by NSF Grant PCN 79-21708. O.S.U. Agricultural Experiment Station Technical Paper No. 6219.     

Response of the clerid predatorThanasimus dubius (F.) to bark beetle pheromones and tree volatiles in a wind tunnel

Tree volatiles and pheromones produced by southern bark beetles were bioassayed for response by the clerid predatorThanasimus dubius (F.). Upwind flights in a laboratory olfactometer, modified from Visser (1976), were used to determine the attractiveness of compounds. Differences in response to a solvent control and pheromone treatment were tested for statistical significance using the Wilcoxon signed ranks test. Both sexes ofT. dubius responded to frontalin, ipsdienol, and -pinene in a dose-dependent manner with different but overlapping concentration ranges. Strong differences between the sexes were observed in response totrans-verbenol, verbenone, andl--pinene. Neither sex responded to ipsenol orendo-brevicomin.Miss. Agric. and For. Exp. Sta. Publication No. 5251.     

Interruption of response ofDendroctonus brevicomis to its attractive pheromone by components of the pheromone

The catch of the western pine beetle,Dendroctonus brevicomis, at an attractive source of racemicexo-brevicomin, racemic frontalin, and myrcene was reduced by surrounding the source with a grid of 48 stations releasing all three compounds together, orexo-brevicomin alone or myrcene alone. Each compound was released at the rate of 2 mg/24 hr/station. The catch at an attractive bolt cut from a tree being colonized byD. brevicomis was not reduced byexo-brevicomin, but was reduced by the combination ofexo-brevicomin, frontalin, and myrcene in one of two tests. When a transect of traps was placed across a 0.81-hectare plot at six of the 48 stations releasing all three compounds, more beetles were caught at outer than at inner traps. More beetles were caught at unbaited traps on trees in a plot when the three compounds were released than when onlyexo-brevicomin or no compounds were released. A few trees were attacked byD. brevicomis in some of the plots. The antiattractant verbenone released from 48 stations at the rate of 4 mg/24 hr/station did not reduce the catch at an attractive tree bolt.Coleoptera: Scolytidae.These studies were supported by the Forest Service, U.S. Department of Agriculture, and in part by grants to the University of California from the U.S. National Science Foundation and U.S. Environmental Protection Agency (NSF GB-34718/BMS 75-04223) and the Rockefeller Foundation. The findings, opinions, and recommendations are not necessarily those of the funding agencies. Mention of commercial enterprises or products is only for information.     

Secondary attraction and field activity of beetle-produced volatiles inDendroctonus terebrans

Controlled infestation ofDendroctonus terebrans (Olivier) on bolts of slash pine,Pinus ellionii Engelm. var.elliottii, elicited greater attraction of wild conspecifics than uninfested bolts. Secondary attraction was not apparent, however, to standing slash pines that had received volunteer attacks, when compared with attraction to unattacked but susceptible trees. Hindguts from in-flight or attacking femaleD. terebrans contained frontalin, and those from in-flight or attacking males containedexo-brevicomin.Trans-pinocarveol,cis-verbenol,trans-verbenol, myrtenal, verbenone, myrtenol, and other compounds were produced by both sexes during gallery construction in host trees. Synthetic frontalin, when deployed with a standard host odor mixture of turpentine and ethanol, was very attractive to maleD. terebrans in field-trapping experiments. The addition of eitherexo-brevicomin orendo-brevicomin to the frontalin-turpentine combination negated the attractive effect of frontalin for males.Trans-verbenol, myrtenol, and verbenone had little effect onD. terebrans behavior. Responses of females did not differ among treatments in any of the 11 field experiments.     

Disruption of Secondary Attraction of the Spruce Beetle, Dendroctonus rufipennis, by Pheromones of Two Sympatric Species

Capture of spruce beetles, Dendroctonus rufipennis, in multiple-funnel traps baited with frontalin and -pinene was reduced by up to 42% in the presence of synthetic (+)-exo- and (+)-endo-brevicomin, aggregation pheromones of the sympatric species Dryocoeles affaber. (±)-endo-Brevicomin was inhibitory to spruce beetles in two experiments and (±)-exo-brevicomin was inhibitory in one experiment, reducing spruce beetle trap catches by up to 87% and 75%, respectively. Spruce beetle trap catches were also reduced by 85% by (±)- or (+)-ipsdienol, but not by (–)-ipsdienol. Ips tridens, a second sympatric species, produces both enantiomers of ipsdienol in its pheromone blend. Responses by D. affaber to its own pheromone were significantly enhanced by addition of the spruce beetle lure. Enantiospecific pheromones of secondary competing species, or less costly racemic substitutes, may be useful for managing spruce beetles using competitive displacement or exclusion. Baiting susceptible hosts with pheromones of secondary species may enhance attack by secondary species, while partially repelling spruce beetles.     

Sensitivity of antennae of male and femaleIps paraconfusus (Coleoptera: Scolytidae) to its pheromone and other behavior-modifying chemicals

The antennal sensitivities of both male and femaleIps paraconfusus were found generally to be greatest for conspecific aggregation pheromones (ipsdienol, ipsenol); intermediate for an additional conspecific pheromone (cis-verbenol), an aggregation synergist (2-phenylethanol), and pheromones/allomones of sympatric species (trans-verbenol, verbenone, and frontalin); and lowest for both host terpenes (alpha-pinene and myrcene) and other bark beetle-produced odorants (exo-brevicomin and linalool). Of the enantiomeric compounds tested, antennae of both sexes did not differ in sensitivity between thetrans-verbenol enantiomers at low dosage levels; but at higher dosages, the conspecific-produced enantiomer, (1R,4S,5R)-(+)-trans-verbenol, elicited larger mean EAG responses than its antipode, (1S,4R, 5S)-(?)-trans-verbenol. At the mid-dosage range, female antennae tended to be slightly more responsive to (S)-(?)-verbenone than to (R)-(+)-verbenone, while male antennae were equally responsive to stimulations by either verbenone enantiomer. In field bioassays there was a large and significant reduction in trap catches ofI. paraconfusus on traps where the (S)-(?)- or (R)-(+)-enantiomers of verbenone were evaporated beside logs containing boring conspecific males. Only when the (S)-(?)-enantiomer of verbenone was evaporated beside logs containing boring males did the sex ratio ofI. paraconfusus trapped shift from female-dominated to male-dominated attraction. Thus both physiological and behavioral data suggest a differential chiral sensitivity of female beetles for the verbenone enantiomers. The relative sensitivities between different chiral compounds derived from one or the other of the common precursoral host terpenes, (S)-(?)- and (R)-(+)alpha-pinene or myrcene, are discussed.     

Semiochemicals produced by western balsam bark beetle,Dryocoetes confusus Swaine (Coleoptera: Scolytidae)

The most prominent beetle-produced volatiles identified in the abdominal extracts of maleDryocoetes confusus Swaine after they had bored for 24 hr in logs of subalpine fir,Abies lasiocarpa (Hook.) Nutt. were:exo andendo-brevicomin,trans-verbenol, verbenone, myrtenol,trans-pinocarveol,cis- andtrans-p-menthen-7-ol, 3-caren-10-ol, and several monoterpenes and sesquiterpenes. Myrtenol was the only conspicuous compound in extracts from males that had been exposed toA.lasiocarpa resin volatiles for 24 hr. Laboratory bioassays indicated that both (–)- and (+)-exo-brevicomin were attractive to femaleD. confusus, and that the (–) enantiomer did not inhibit response to its antipode. Results from field trapping experiments indicated that bothero-brevicomin and myrtenol are aggregation pheromones forD. confusus.exo-Brevicomin baits were effective in causing attack byD. confusus on baited and surrounding trees, suggesting that this pheromone may have utility in manipulating populations of the beetle.Research supported by the Natural Sciences and Engineering Research Council of Canada, Operating Grants A3881 and A3706 and Strategic Grant G1039.     

Effect of verbenone on response ofDendroctonus brevicomis toexo-Brevicomin,frontalin, and myrcene

exo-Brevicomin (E), frontalin (F), and myrcene (M) were released at two rates 10-fold apart with verbenone at four rates 10-fold apart, and without verbenone in plots with one trap on a vertical cylinder at the pheromone source and one trap on each of four cylinders 5 m away. Catch of the western pine beetleDendroctonus brevicomis decreased with increasing levels of verbenone at both release rates of EFM, but not all differences in catch were statistically significant. Significantly more beetles were caught at the high rate of EFM than at the low rate, combining all rates of verbenone. The percent of total beetles caught at the center trap tended to decrease with increasing rates of verbenone, but the only statistically significant differences were at the low rate of EFM.Coleoptera: Scolytidae.     

Quantitative estimates of the western pine beetle attractive pheromone components,exo-brevicomin,frontalin, and myrcene in nature

Three parameters are described for estimating the natural occurrence of chemically defined insect pheromones: (1) the rate and duration of release by the insect, (2) the density of the pheromone-emitting insect population in both time and space, and (3) dispersal and degradation rates of the chemicals. Each of these parameters, except dispersal, was estimated for a population ofDendroctonus brevicomis LeC, and its three component attractive pheromones. A single generation of 610,000 beetles, believed to comprise the entire population in a 65-km² forest, was estimated to have released 0.78, 3.7, and 370.5 g of frontalin,exo-brevicomin, and myrcene, respectively, within a 30-day period. Dendroctonus brevicomis Le Conte (Coleoptera: Scolytidae).Parts of this report were prepared for the U.S. Forest Service to accompany a future application to the Environmental Protection Agency for registration of these attractants for use in the suppression ofD. brevicomis populations. The research was supported by grants to the University of California (D.L. Wood) from the National Science Foundation/Environmental Protection Agency (NSF-6B-34718/BMS75-04223), Forest Service, U.S. Department of Agriculture and Rockefeller Foundation. The findings, opinions, and recommendations are those of the authors and not necessarily those of these institutions.     

Pheromone production by axenically rearedDendroctonus ponderosae andIps paraconfusus (Coleoptera: Scolytidae)

Mountain pine beetles,Dendroctonus ponderosae Hopkins, and California five-spined ips,Ips paraconfusus Lanier, were reared axenically from surface-sterilized eggs on aseptic pine phloem. After 24 hr in host logs, axenip femaleD. ponderosae and maleI. paraconfusus produced the aggregation pheromones,trans-verbenol (D. ponderosae), and ipsenol and ipsdienol (I. paraconfusus). Emergent, axenically reared maleD. ponderosae contained normal amounts of the pheromoneexo-brevicomin. Axenic femaleD. ponderosae treated with juvenile hormone or exposed to vapors of -pinene, produced the pheromonetrans-verbenol. By 25–35 days after eclosion, axenic females exposed to -pinene vapors produced over six times as muchtrans-verbenol as wild females, suggesting that while microorganisms in wild females may producetrans-verbenol, they may also inhibit production of the pheromone or use it as a substrate.Research supported in part by Natural Sciences and Engineering Research Council, Canada, Operating Grants A3881 and A3706, and H.R. MacMillan Family Fund Fellowships to J.E. Conn and D.W.A. Hunt.     

Antennal olfactory responsiveness of Douglas-fir beetle,Dendroctonus pseudotsugae Hopkins (Coleoptera: Scolytidae) to pheromones and host odors

Electroantennograms were obtained fromD. pseudotsugae in response to the pheromones 3-methylcyclohex-2-en-1-one (3,2-MCHone), 3-methylcyclohex-2-en-1-ol (3,2-MCHol), frontalin,trans-verbenol, verbenone, and the host terpene hydrocarbons limonene and camphene. Male and female beetles were 10 and 100 times more sensitive to 3,2-MCH-one and 3,2-MCHol than to the other compounds. Of the other compounds, males were most sensitive totrans-verbenol, verbenone, and camphene, while females were most sensitive to frontalin, limonene, and camphene. The results parallel and help explain behavior of individual males and females during host tree selection, aggregation, and colonization.