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.     

Ipsenol: an aggregation pheromone forIps latidens (Leconte) (Coleoptera: Scolytidae)

Ipsenol was identified from the frass of male, but not female,Ips latidens from British Columbia, feeding in phloem tissue of lodgepole pine,Pinus contorta var.latifolia. The responses ofJ. latidens to sources of ips-enol andcis-verbenol were determined with multiple-funnel traps in stands of lodgepole pine in British Columbia. Ipsenol attracted both male and femaleI. latidens, verifying that it is a pheromone for this species. MaleI. latidens showed a slight preference for (S)-(–)-ipsenol.cis-Verbenol was not produced by beetles of either sex and, in contrast to an earlier report, both enantiomers inhibited attraction to ipsenol-baited traps. The predators,Enoclerus sphegeus andThanasimus undatulus (Cleridae), were attracted to traps baited withcis-verbenol and ipsenol.     

Inhibition of the attractant pheromone response inIps pini andI. paraconfusus (Coleoptera: Scolytidae): Field evaluation of ipsenol and linalool

Linalool, a compound from maleI. pini, previously suggested as an inhibitor forI. paraconfusus, has no obvious effect on the response ofI. paraconfusus to ponderosa pine bolts containing maleI. paraconfusus. I. pini from California and New York equally inhibit the response ofI. paraconfusus to maleI. paraconfusus. Ipsenol, one component of the attractant pheromone ofIps paraconfusus, inhibits attacks byIps pini on ponderosa pine logs baited with maleI. pini. The concentration of ipsenol used appears to be critical for effective suppression of attacks.This investigation was supported in part by a grant from the U.S. Forest Service.     

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.     

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.     

Ancestral semiochemical attraction persists for adjoining populations of siblingIps bark beetles (Coleoptera: Scolytidae)

Three species ofIps pine bark beetles in Hopping's group IX (S.L. Wood'sgrandicollis group),Ips confusus, I. lecontei, andI. paraconfusus, are parapatrically distributed in the American Southwest. They share post-Pleistocene altitudinal ecotones with their host pines. Adjacent to these areas of host overlap, we tested the hypothesis that aggregation pheromones produced by male beetles and/or host volatiles are sufficient to elicit the speciesspecific colonization behaviors typical of these threeIps species in nature. A more distantly related species,I. pini (Hopping's group IV, S.L. Wood'spini group) was used for outgroup comparison. Under the influence of pheromone, males ofI. confusus andI. paraconfusus do not discern among unin-fested log bolts of host and nonhost pine prior to bark contact. Males responding to pheromones emanating from infested bolts are similarly undiscriminating. Females ofI. confusus andI. lecontei olfactorily discern the combination of conspecific males in host pine from other possible beetlepine combinations; females ofI. paraconfusus do not. FemaleI. pini discerned conspecific pheromone from that ofI. lecontei. The bark beetle predator,Enoclerus lecontei, is attracted by, but does not discriminate among, the male-produced volatiles of these Group IXIps species. These results support a hypothesis that divergence in pheromonal responses by these group IXIps species has evolved following their speciation, having been manifested first in the female sex. Evolutionarily, the derived pheromonal messages have preceded their behavioral discrimination by these beetles. Additional speciesspecific cues may operate between the sexes in the field that may preclude heterospecific pairings.     

Attractive and inhibitory pheromones produced in the bark beetle,Dendroctonus brevicomis,during host colonization: Regulation of inter- and intraspecific competition

Quantities of attractive (exo-brevicomin and frontalin) and inhibitory (trans-verbenol, verbenone, and ipsdienol) pheromones were monitored in both sexes ofDendroctonus brevicomis during their colonization of a ponderosa pine. Verbenone was found in males in the greatest amounts at the time of landing, and it declined more rapidly than the other pheromones in either sex. The amounts of frontalin andexo-brevicomin in males and females, respectively, increased after initial boring within the host but began to decline after mating. The quantity oftrans-verbenol in both sexes (females had significantly more) declined more gradually thanexo-brevicomin, frontalin, and verbenone. Ipsdienol was found only in males during the initial stages of attack when encountering the resin. It is suggested that along with a general decline in all pheromonal components, a sufficient change in the ratio of the attractive pheromones to an inhibitory pheromone,trans-verbenol, may play a role in termination of aggregation.trans-Verbenol may also function along with verbenone and ipsdienol in limiting the density of attack and thus intraspecific competition. These inhibitory pheromones also appear to cause several competing species of bark beetle to avoid landing in areas infested withD. brevicomis, even when their own pheromone is present.Coleoptera:Scolytidae. These studies were supported in part by the Rockefeller Foundation, USDA Forest Service, Regional research project W-110, 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.     

Conversion of verbenols to verbenone by yeasts isolated fromDendroctonus ponderosae (Coleoptera: Scolytidae)

A variety of symbionts associated with bark beetles are capable of producing compounds that are used as pheromones by their hosts. We report that two yeasts associated withDendroctonus ponderosae Hopkins,Hansenula capsulata Wickerham, andPichiapinus (Hoist) Phaff, are capable of convertingcis- andtrans-verbenol efficiently into verbenone.trans-Verbenol, which is produced by femaleD. ponderosae, acts as an aggregation pheromone for this scolytid, while verbenone, which other studies have indicated that microbe-reducedD. ponderosae are incapable of producing, acts as an antiaggregation pheromone.D. ponderosae appears to rely primarily on microbial symbionts for terminating aggregation and mass attack on individual host trees.Research supported in part by the Natural Sciences and Engineering Research Council, Canada, Operating Grant A3881, and Strategic Grant G1039.     

Influence of aggregation inhibitors (verbenone and ipsidenol) on landing and attack behavior ofDendroctonus brevicomis (Coleoptera: Scolytidae)

The influence of the aggregation inhibitors verbenone and ipsdienol on the response of western pine beetle,Dendroctonus brevicomis, to attractive host trees was investigated. Paired ponderosa pine trees (Pinus ponderosa) were baited with aggregation semiochemicals to stimulate mass attack. One tree in each pair received an inhibitor treatment consisting of five sets of two verbenone and two ipsdienol dispensers spaced 1 m apart vertically along the tree bole. Beetle landing was monitored with sticky traps on the tree bole, and attack density was assessed from bark samples removed four or seven days after baiting. The inhibitor treatment resulted in a significant reduction of both the numbers of beetles landing on trees and the density of attacking beetles compared to control trees (without inhibitors). The ratios of beetle landing density to attacking density were not different between inhibitor-treated and control trees, nor were the vertical distributions of beetles landing or attacking, suggesting that beetle behavior was primarily influenced at a longer range, prior to landing on the tree. Although the application of verbenone and ipsdienol did not preventD. brevicomis from attacking baited trees, our results suggest that when applied to unattacked (and unbaited) trees, their effectiveness at reducing the attack pressure might allow trees having a certain amount of resistance to survive attack by pioneer beetles.     

Semiochemicals from bark beetles: New results,remarks, and reflections

A brief survey is given about recent results in the identification of semiochemicals in bark beetles: Males ofIps sexdentatus (Boern.), stressed by the attack on resinous trees produce large amounts of 3(S)-1-methyl-5-(1-hydroxyl-1-methylethyl)-cyclohexa-1,3-diene. The compound appears to be derived from ³-carene and acts as a repellent. Males ofIps typographus (L.), stressed through the attack on unsuitable host material release 3-methyl-7-methylene-1,3(E), 8-nonatriene, which seems to act as a repellent. The odor bouquet of three species ofPityogenes is described. The occurrence of (+)-grandisol and other compounds related to weevil pheromones points to a close relation between Scolytidae and Curculionidae. Females ofDendroctonus simplex (Le Conte) use (–)-frontalin as the main pheromone. 6-Methyl-6-hepten-2-one, a minor component among the volatile compounds released by the females, is regarded as a possible precursor of frontalin. Similarly, (2R,5S)-2(1-hydroxyl-1-methylethyl)-5-methyltetrahydrofuran, pityol, a pheromone ofPityophthorus spp., is regarded to at least share a common biogenetic precursor with 6-methyl-5-hepten-2-ol, sulcatol. A new bicylic acetal, 2-ethyl-1,5-dimethyl-6,8-dioxabicyclo[3.2.1]octane, is described as an aggregation pheromone of the beech bark beetle,Taphrorychus bicolor (Herbst). Structural relationships between bark beetle pheromones and plant volatiles are discussed.     

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.     

Allelopathic potential ofErica vagans,calluna vulgaris,andDaboecia cantabrica

In laboratory tests the allelopathic potential ofErica vagans, Calluna vulgaris, andDaboecia cantabrica was determined. Aqueous extracts of flowers ofD. cantabrica and leaves ofC. vulgaris inhibit root and hypocotyl growth of red clover, the former causing 51% inhibition of germination. Intact aerial parts of the Ericaceae here studied drastically reduced the growth of red clover and 100% inhibition of germination was caused by flowers ofD. cantabrica. Inhibition of aqueous extracts remains after Chromatographic separation, and two well-defined inhibition zones may be observed. Hydrosoluble organic compounds (phenol-like compounds) could probably be responsible for the inhibitions detected.     

(S)-(+)-Ipsdienol: Interspecific inhibition ofIps latidens (leconte) byIps pini (Say) (Coleoptera: Scolytidae)

In south-central British Columbia, the attraction ofIps latidens (LeConte) to its pheromone, ipsenol, was inhibited by (S)–(+)-ipsdienol, a pheromone forI. pini (Say). (R)-(–)–lpsdienol had no effect onI. latidens. (S)–(+)-lpsdienol probably plays a role in interspecific communication between the two species, facilitating reductions in interspecific competition for breeding material and/or interspecific mating interference.     

Outdoor attractancy of males ofMatsucoccus josephi (Homoptera: Matsucoccidae) andElatophilus hebraicus (Hemiptera: Anthocoridae) to synthetic female sex pheromone ofMatsucoccus josephi

The active component of the sex pheromone ofMatsucoccus josephi is (2E,6E,8E)-5,7-dimethyl-2,6,8-decatrien-4-one; the chemical is also a powerful kairomone of adult males and females of the bugElatophilus hebraicus the principal predator ofM. josephi. The presence of theZ isomer (2E,6Z,8E)-5,7-dimethyl-2,6,8-decatrien-4-one does not interfere with the attractancy of the activeE component forM. josephi males or the bug. Our results show a clear dose-response between trap catch ofM. josephi males andE. hebraicus. Conversely, increasing amounts of theZ isomer in the mixture did not affect the attraction of the scale insect males or the bug. The catch ofM. josephi males did not differ significantly among traps of different color, and was significantly higher with traps attached to the tree trunk than those suspended between trees. Comparison of the catch ofM. josephi among the three forests and between pine species suggests that the level of infestation ofPinus halepepsis andPinus brutia ssp.brutia is similar, despite the fact that the latter pine is resistant to the scale insect. Both sexes ofE. hebraicus were trapped in much lower numbers at the more infested sites. This may be related to interference with the activity ofE. hebraicus due to deterioration and drying of parts of the tree crowns and heavy colonization by generalist predators in injured trees.     

Effective attraction radius

The catches of bark beetles (Coleoptera: Scolytidae) were compared between attractive traps releasing semiochemicals and passive traps (cylindrical sticky screens hung, at 10 heights of 0.7–11.5 m, on poles). A central attractive-trap pole was surrounded by three passive-trap poles spaced 50 or 100 m away at the apices of an equilateral triangle. The catches ofTomicus piniperda and other scolytid species on the attractive-trap pole baited with host monoterpenes, or the catches ofIps typographus attracted to synthetic pheromone, were compared to passive trap catches in a Scots pine forest or in a Norway spruce clear-cut, respectively. Information about flight height distributions of the above scolytid species, andHylurgops palliatus, Cryphalus abietis, Pityogenes chalcographus, P. quadridens, P. bidentatus, andTrypodendron domesticum were obtained on the passive and attractive trap poles. A new method is presented for determining the densities of flying insects based on the passive trap's dimensions and catch, duration of test, and speed of insect. Also, a novel concept, the effective attraction radius (EAR), is presented for comparing attractants of species, which is independent of insect density, locality, or duration of test. The EAR is obtained by the ratio of attractive and passive trap catches and the dimensions of the passive trap, and thus should correlate positively with the strength of the attractant and the distance of attraction. EARs are determined from catch data ofT. piniperda andI. typographus as well as from the data of previous investigations on the same or other bark beetles.     

Cuticular hydrocarbons of eight species of north american cone beetles,Conophthorus hopkins

A study to determine the degree of similarity and/or diversity among eight of the 15 described species ofConophthorus is reported. Cuticular hydrocarbons were evaluated forC. conicolens, C. ponderosae, C. cembroides, C. edulis, C. radiatae, C. coniperda, C. resinosae, andC. banksianae. Seventy-eight individual and isomeric mixtures of hydrocarbons were identified by gas chromatography-mass spectrometry, includingn-alkanes, alkenes, alkadienes, 2- or 4-methylalkanes, 3-methylalkanes, and single-component and isomeric mixtures of internally branched mono-, di-, and trimethylalkanes. Differences in alkenes and mono-, di-, and trimethylalkanes can be used easily to separate the eight species.Conophthorus conicolens andC. ponderosae contain the most complex blends. Hydrocarbon patterns in three geographically separated populations ofC. ponderosae, each from a different host, are qualitatively identical with the exception of a homologous series of 3,7-dimethylalkanes from adults collected fromPinus lambertiana cones. The latter could comprise a sibling species. Hydrocarbon mixtures of two eastern species,C. resinosae andC. banksianae, are qualitatively identical, supporting the suspicion thatC. banksianae may not be a valid species. Closely relatedC. cembroides andC. edulis have similar combinations of hydrocarbons except for a unique and abundant alkene (C₂₇₁) inC. edulis and two dimethyhexacosanes inC. cembriodes.Coleoptera: Scolytidae.     

Variation in cardenolide content of the lygaeid bugs,Oncopeltus fasciatus andLygaeus kalmii kalmii and of their milkweed hosts (Asclepias spp.) in central California

A colorimetric assay was used to quantify the amount of cardenolides in the lygaeid bugsOncopeltus fasciatus andLygaeus kalmii kalmii and their milkweed host plants (Asclepias spp.) in central California. The cardenolide content of individual insects, determined in microgram equivalents of digitoxin, varied from zero to over 300 g per insect. Sources of variation of cardenolide content in the insects include interspecific and intraspecific differences in the content of the host plant species and also differences in the content of plant organs on which insects were feeding. This last source of variability may explain temporal variation in the cardenolide content of the insects. Adults ofO. fasciatus, which migrate into California in the late spring and early summer, and adults ofL. k. kalmii, which emerge from winter hibernacula in the early spring, contained small to immeasureable amounts of cardenolides. The colonization pattern ofO. fasciatus on species ofAsclepias in north central California suggests that this species does not maximize its opportunities to sequester large quantities of cardenolides from potential hosts. The emetic potential of lygaeids in California to vertebrate predators is briefly discussed.     

Relationships betweenDendroctonus micans Kug. (Coleoptera: Scolytidae) survival and development and biochemical changes in Norway Spruce,Picea abies (L.) Karst., phloem caused by mechanical wounding

The bark of Norway spruce trees,Picea abies (L.) Karst., was wounded to produce areas of bark with differing biochemical characteristics. Adults and larvae ofDendroctonus micans Kug. were inserted into these trees at points around the wounds. Larval survival was higher, larval dry weights were higher, and adults were more likely to oviposit at sites around the wound that had higher phloem moisture contents. Larvae showed higher survival and dry weights in phloem with low terpene contents. Adults oviposited in phloem with lower stilbene glucoside contents. Phloem moisture differed at sites around the wound compared with that at a control site away from the wound, and the nitrogen content of entire bark cores was higher above wounds than at control sites away from the wounds. Phloem moisture correlated with the nitrogen content of entire bark. Phloem terpene contents correlated with each other, as did the stilbene contents. Associations between the host exploitation pattern ofD. micans and the biochemistry of the host phloem are discussed in relation to preselection by adults of sites suitable for larval development.     

Secondary attraction inGnathotrichus retusus and cross-attraction ofG. sulcatus (Coleoptera: Scolytidae)

Laboratory bioassays indicated that frass produced byGnathotrichus retusus males and male gut extracts were attractive to both sexes of beetles. The frass became attractive to females within 2 days of the commencement of boring activity, and attractive frass was produced for at least lOdays. Attraction of frass rose sharply after pairing of males with females. However, this increase in attraction could be due to increased boring activity and pheromone production by males following establishment of the mutualistic fungus. In field trapping experiments, male-infested logs were always highly attractive, but in one experiment, female-infested logs and logs infested by both sexes were also attractive. Thus, females could be involved in secondary attraction. High cross-attraction ofG. sulcatus toG. retusus infested logs in field experiments, as well as a moderate but significant response byG. retusus to sulcatol in the laboratory, suggests that they share sulcatol as a common pheromone.Research supported by Canadian Forestry Service Science Subvention Grant, National Research Council of Canada Operating Grant No. A3881, National Science Foundation (U.S.A.) Operating Grant No. PCM74-13643-A02, and the Council of Forest Industries of British Columbia.     

Enantiomer-based specificity in pheromone communication by two sympatricGnathoirichus species (Coleoptera: Scolytidae)

The aggregation pheromone ofGnathotrichus retusus was isolated and identified as (S)-(+)-sulcatol (6-methyl-5-hepten-2-ol). In laboratory and field experiments,G. retusus responded to (S)-(+)-sulcatol, but not to (±)-sulcatol, which was attractive to the sympatric species,G. sulcatus. G. sulcatus did not respond to optically pure (S)-(+)-sulcatol, but began to respond when 1% (R)-(–)-sulcatol was present in an enantiomeric mixture.Research supported by the National Science Foundation, U.S.A. (Grant BMS-74-13643), and the Natural Sciences and Engineering Research Council of Canada (Co-op grant A0243 and Operating Grant A3881 and A3785).