Variation in semiochemical-mediated prey-predator interaction:Ips pini (Scolytidae) andThanasimus dubius (Cleridae)

The bark beetleIps pini (Say) displays variation in its response to and production of enantiomeric blends of its pheromone ipsdienol. One of the principal predators ofIps pini isThanasimus dubius (F.), which uses ipsdienol as a kairomone for prey location. During 1988 and 1989, in Wisconsin and Michigan, the response of both species to a range of enantiomeric blends of ipsdienol was investigated. Blends tested had the following ratios of the (S)-(+) to (R)-(–) enantiomers: 3%97%, 25%75%, 50%50%, 75%25%, and 97%3%. Either75% (+) 25% (–) or 50% (+) 50% (–) ipsdienol captured the mostIps pini in both years at both sites. The 25% (+)75% (–) blend also caught moreIps pini than the control during both years at both sites. All blends tested were attractive toThanasimus dubius in both years at both locations. Blend preferences of both species were variable and labile at both sites. Response patterns of both species in Wisconsin were different from those in Michigan each year. Furthermore, response patterns of both species to the ipsdienol blends changed from 1988 to 1989 at both locations. A genetic component to this variation would permit predator-prey coevolution, as well as the development of resistance byIps pini to management strategies based on mass-trapping with single blends.     

Variation in semiochemical-mediated prey-predator interaction:Ips pini (Scolytidae) andThanasimus dubius (Cleridae)

The bark beetleIps pini (Say) displays variation in its response to and production of enantiomeric blends of its pheromone ipsdienol. One of the principal predators ofIps pini isThanasimus dubius (F.), which uses ipsdienol as a kairomone for prey location. During 1988 and 1989, in Wisconsin and Michigan, the response of both species to a range of enantiomeric blends of ipsdienol was investigated. Blends tested had the following ratios of the (S)-(+) to (R)-(–) enantiomers: 3%97%, 25%75%, 50%50%, 75% 25%, and 97% 3%. Either 75% (+)25% (–) or 50% (+)50% (–) ipsdienol captured the mostIps pini in both years at both sites. The 25% (+)75% (–) blend also caught moreIps pini than the control during both years at both sites. All blends tested were attractive toThanasimus dubius in both years at both locations. Blend preferences of both species were variable and labile at both sites. Response patterns of both species in Wisconsin were different from those in Michigan each year. Furthermore, response patterns of both species to the ipsdienol blends changed from 1988 to 1989 at both locations. A genetic component to this variation would permit predator-prey coevolution, as well as the development of resistance byIps pini to management strategies based on mass-trapping with single blends.Editor's Note: This paper originally appearedin Journal of Chemical Ecology, Volume 17, Number 3 (March 1991), pp. 515–524. Due to several significant printer's errors, the paper is reprinted here in entirety in its corrected form.     

Inter- and intrapopulation variation of the pheromone,ipsdienol produced by male pine engravers,Ips pini (Say) (Coleoptera: Scolytidae)

We determined the chirality of ipsdienol in individual male pine engravers,Ips pini (Say), from New York, California, and two localities in British Columbia (BC). Both quantity and chirality of ipsdienol varied significantly between and within populations ofI. pini. Beetles from California and southeastern BC produced primarily (R)-(–)-ipsdienol with mean ratios of (S)-(+) : (R)-(–) of 9 : 91 and 11 : 89, respectively, while beetles from New York produced primarily (S)-(+)-ipsdienol with a mean (S)-(+) : (R)-(–) ratio of 57 : 43. A population from southwestern BC was unlike any other known western population, producing primarily (S)-(+)-ipsdienol with a mean (S)-(+) : (R)-(–) ratio of 66 : 34. In contrast to the unimodal chirality profiles for ipsdienol production in populations from California and southeastern BC, the profiles of the populations from southwestern BC and New York were bimodal, with a common mode at approximately 44 : 56 (S)-(+) : (R)-(–). Bimodality in the profiles of ipsdienol chirality in two populations ofI. pini and remarkably high levels of intrapopulation variation in pheromone chirality in all four populations suggest that evolutionary change in pheromone channels of communication could occur, possibly in response to artificial selection pressures such as mass trapping.     

Opposing Effects of Host Monoterpenes on Responses by Two Sympatric Species of Bark Beetles to Their Aggregation Pheromones

The effects of host tree monoterpenes on primary and secondary attraction of the bark beetles, Ips pini and Ips grandicollis, and their associated phloeophagous insects were investigated. Monoterpenes alone were not attractive to I. pini. However, monoterpenes mediated the attraction of I. pini to its aggregation pheromones. With the exception of 3–carene, the effect of monoterpenes on I. pini's response to its pheromone, ipsdienol plus lanierone, was inhibitory. In contrast, (–)--pinene both attracted I. grandicollis and enhanced the attraction of I. grandicollis to its pheromone, ipsenol. No monoterpene inhibited the response of I. grandicollis to its pheromone. The inhibitory effect of host monoterpenes on I. pini response to its aggregation pheromone differs from previous work, in which monoterpenes either synergized responses or had no effect. In addition to possible geographic differences, the concentrations used in our study simulated trees that had begun to respond to attack, whereas previous studies deployed lower concentrations, which simulated constitutive phloem from unattacked trees. These results support the view that trees that undergo induced responses to bark beetles sometimes inhibit attraction of additional beetles, despite the beetles' production of aggregation pheromones. Neither species displayed cross-attraction to the pheromone of the other. The red turpentine beetle, Dendroctonus valens, showed weak and consistent attraction to (+)--pinene and in some cases to (–)--pinene. Attraction to (–)--pinene was usually enhanced by Ips spp. pheromones. The absence of strong attraction to (+)--pinene and partial attraction to (–)--pinene suggest that the effects of different stereoisomers of -pinene on D. valens vary throughout its geographical range. Hylastes porculus was also attracted to some monoterpenes, particularly (–)--pinene. An additional 10 species of phloeophagous insects were caught in response to monoterpenes and/or pheromones, including the pine root weevils, Hylobius pales and Pachylobius picivorus, and the longhorned beetle, Monochamus carolinensis.     

IPS pini: The basis for interpopulational differences in pheromone biology

Ips pini from Idaho (ID) produce (–)-ipsdienol and respond to it in the laboratory.I. pini from New York (NY) produce (+) and (–)-ipsdienol in a 6535 ratio and respond much more strongly to the (+) than to the (–) enantiomer. Response byI. pini-ID to (–)-ipsdienol in the laboratory was inhibited by (+)-ipsdienol, while the field response byI. pini-NY was optimal to a 50:50 mixture of (+) and (–)-ipsdienol. This differential production and response to enantiomers of ipsdienol is sufficient to account for populational specificity observed in earlier experiments and confirmed by our current work. Reciprocal hybrids of the NY and IDI. pini populations did not differ in their patterns of attraction and response; those of both types of F₁ were similar to NY beetles. Thus, it appears that (+)-ipsdienol is produced by the hybrids and is necessary for their maximum response, but this supposition was not tested. The genes controlling the pheromone biology ofI. pini are not sex linked. Since previous whole-antenna and single-cell electrophysiological data show that the receptor systems of the two populations are virtually identical, response behavior seems to be governed by the central nervous system rather than by the characteristics of the olfactory receptors.Coleoptera: Scolytidae.Supported by grants from the Rockefeller Institute, the National Science Foundation, and the Research Foundation of Providence, Rhode Island.     

Response of mountain pine beetle,Dendroctonus ponderosae Hopkins,and pine engraver,Ips pint (SAY), to ipsdienol in southwestern British Columbia

In bioassays conducted with walking beetles in the laboratory (S)–(+)–, (R)-(–)-, and (±)-ipsdienol were attractive alone, but reduced the attraction of both sexes of the mountain pine beetle,Dendroctonus ponderosae Hopkins, to extracts of female frass. Field trapping studies indicated that attraction ofD. ponderosae to a stimulus composed of myrcene +trans-verbenol +exo-brevicomin was significantly reduced with the addition of (±)- and sometimes (S)-(±)-ipsdienol. Thus, (S)-(+)-ipsdienol produced by males of this species may act as an antiaggregation pheromone. (S)-(+)-Ips-dienol is thought to function as a repellent allomone against the pine engraver,Ips pini (Say), in regions whereI. pini utilizes (R)-(–)-ipsdienol as an aggregation pheromone. However, in southwestern British ColumbiaI. pini was attracted to the (±)-ipsdienol used in field bioassays ofD. ponderosae, a finding consistent with the production of both enantiomers byI. pini in this region. When presented with the ternary semiochemical bait forD. ponderosae, (±)-ipsdienol was not attractive toI. pini. Thus, the activity of (S)-(+)-ipsdienol as a repellent allomone againstI. pini seems to be replaced in southwestern British Columbia by the inhibitory effects of myrcene,trans-verbenol,exo-brevicomin, or some combination thereof.Research supported in part by Natural Sciences and Engineering Research Council, Canada, Operating Grant A3881, Strategic Grant G1039, a Postgraduate Scholarship to D.W.A. Hunt, and the Science Council of British Columbia Grant 1 (RC-10)     

Seasonal variability in response ofIps pini (Coleoptera: Scolytidae) to ipsdienol in New York

In May,Ips pini in New York did not respond in the field to 50–98.5% (R)-(–)-ipsdienol (synthetic). In September, beetles responded strongly to 50–60% (R)-(–)-ipsdienol (synthetic). In May and June, New York beetles showed marked preference for their own males over Arizona males, which produce an average of 94.1% (R)-(–)-ipsdienol. This suggested that ipsdienol stereochemistry alone does not ensure activity and that an additional compound is necessary for attraction in May. In the second year of field tests, attraction to synthetic ipsdienol and male beetles was tested in the spring, summer, and fall. There was response only to males in the spring and mid-summer and to both males and synthetic ipsdienol in the late summer and fall, causing a significant treatment x sampling period (date) interaction. Laboratory-reared beetles were not significantly more attracted to ipsdienol than to a blank airstream in laboratory assays, while male volatiles were significantly more attractive than ipsdienol and the blank. These data demonstrate that there is one or more unknown semiochemicals necessary for pheromonal response and that the behavioral activity of synthetic ipsdienol varies seasonally.Deceased.     

Enantiospecific pheromone production and response profiles for populations of pine engraver,Ips pini (Say) (Coleoptera: Scolytidae), in British Columbia

Analyses of the enantiomeric composition of ipsdienol produced by individual male pine engravers,Ips pini (Say), from six populations in British Columbia, support the hypothesis that New York and Idaho races of this species hybridize in southeastern British Columbia. Production profiles, expressed as frequency distributions of (+):(-) ipsdienol ratios [= ratio of (S)-(+)-ipsdienol to (R)-(-)-ipsdienol], were bimodal for four western British Columbia populations. The (+):(-) ratios ranged from 63:37 to 71:29. consistent with those previously found for the New York race. The profile for a southeastern population from Radium, British Columbia, was intermediate between those for the four western British Columbia populations and that from one population in Kimberley, British Columbia, just south of Radium. Males in the Kimberley population produce predominantly (R)-(-)-ipsdienol, typical of California nad Idaho males. Response profiles of different individuals ofI. pini, determined by captures of beetles in multiple-funnel traps baited with ipsdienol of 11 different (+):(-) ratios, were not consistent with production profiles. Populations in Williams Lake and Princeton, in western British Columbia, and Radium, in southeastern British Columbia, had response profiles with maximal attraction to ipsdienol over a broad range of (+):(-) ratios, falling off as enantiomeric purity was approached at either end of the spectrum. This type of response profile is consistent with that for the New York race, which has been shown to respond optimally to (+):(-) ratios ranging from 40:60 to 70:30. The response profile of the Kimberley population gradually declined from maximal attraction to ipsdienol with a (+):(-) ratio of 2:98 to the lowest response at a (+):(-) ratio of 98:2. The attraction ofI. pini to chemical stimuli in California is interrupted by ipsdienol with a (+):(-) ratio >5:95, a pheromone of a host competitor, the California five-spined ips,Ips paraconfusus Lanier. We hypothesie that the Idaho race, which does not compete withI. paraconfusus due to geographical separation, is characterized by a Kimberley-type enantiomeric response profile, intermediate between those of the New York and California races.     

Repeatability of female response to ipsdienol enantiomeric mixtures by pine engraver,Ips pini (Coleoptera: Scolytidae)

Repeatability, a measure of the proportion of variance in a character that occurs among rather than within individuals, is assessed for the phenotypic trait of female response preference for enantiomeric blends of ipsdienol inIps pini at two pheromone concentrations—1 µg and 5 g of ipsdienol. Average female response shows greater repeatability at the higher pheromone dosage when females are tested in two successive sets of five assays than when assayed in three successive sets or at the lower dosage. Repeatability within each set of five assays is highest for the first set and decreases thereafter. Thus the response phenotype of females for enantiomeric blends of ipsdienol in this experiment is context dependent; female choice of an enantiomeric blend differed between dosages and among sets of assays.     

Insect predator-prey coevolution via enantiomeric specificity in a kairomone-pheromone system

Insect predators can be guided to their prey by a kairomonal response to the prey pheromone. We found this phenomenon to be highly specific in the bark beetle predatorThanasimus dubius. Olfactory responses and behavioral tests revealed that the predator is guided to its major preyDendroctonusfrontalis by the primary enantiomer of the pheromone of the prey, (1S, 5R)-(–)-frontalin. These and other findings suggest the co-evolution of a kairomone system of the predator and the pheromone system of its prey.Texas Agricultural Experiment Station paper No.TA17770. The work was funded in part by McIntire-Stennis project 1525 (TAES) and the USDA program entitled The Expanded Southern Pine Beetle Research and Applications Program through TAES-CSRS grant 680-15-10. The findings, opinions, and recommendations reported herein are those of the authors and not necessarily those of the United States Department of Agriculture.     

Electrophysiological and Olfactometer Responses of Two Histerid Predators to Three Pine Bark Beetle Pheromones

We measured electrophysiological responses in the antennae of two predaceous hister beetles, Platysoma parallelum and Plegaderus transversus, exposed to racemic mixtures of primary aggregation pheromones of scolytid bark beetle prey, ipsenol, ipsdienol, and frontalin. No significant differences were found for either histerid species between male and female antennal responses to any of the three pheromones. Measurement of antennal threshold responses indicated that Pla. parallelum has increasing antennal sensitivity to ipsdienol, ipsenol, and frontalin. In contrast, Ple. transversus exhibited similar detection thresholds to all three pheromones. Pla. parallelum antennae exhibited different response amplitudes to the three pheromones at quantities above the detection threshold, while Ple. transversus had similar responses to each. Behavioral responses to the same three pheromones were evaluated for both histerid species using pedestrian olfactometer bioassays. Both species were attracted to frontalin and ipsenol, but not ipsdienol. Pla. parallelum was significantly more attracted to frontalin than ipsenol, while Ple. transversus showed no significant preference for either compound. Our results suggest that histerids that prey upon pine bark beetles may have different host or host habitat preferences, which could reduce interspecific competition.     

Specialization of olfactory cells to insect-and host-produced volatiles in the bark beetleIps pini (say)

Electrophysiological recordings inIps pini were made from single olfactory cells stimulated by six concentrations of eleven compounds—either pheromones or host constituents. The receptor cells were grouped according to their differential responses to these biologically significant compounds. Cells specialized to either ipsdienol or ipsenol (pheromones for severalIps species) appeared to be relatively uniform in their sensitivity to all compounds tested. Another group of cells specialized tocis- andtrans-verbenol were more variable in their sensitivities to the substances tested. The activation of separate groups of cells by the different pheromones and host compounds shows (by exclusion) that behaviorally expressed syner-gistic as well as inhibitory action mainly is due to an interaction in the central nervous system rather than an interaction of the compounds on the receptors. All ipsdienol cells responded to both enantiomers of ipsdienol. Although some differences between responses to the enantiomers by individual cells were observed, these did not form a consistent pattern, and no net differences could be found between the eastern and western populations ofI. pint.Coleoptera: Scolytidae.     

Chemically Mediated Predator-free Space: Herbivores Can Synergize Intraspecific Communication Without Increasing Risk of Predation

Natural enemies of herbivores often locate cryptic insects by responding to volatiles associated with the prey's feeding and mating. For example, predators of bark beetles (Coleoptera: Scolytidae) exploit the aggregation pheromones that their prey use to attract mates and secure hosts. Bark beetles are cryptic insects that feed and develop in the subcortical tissues of trees and spend all but a portion of their life history within this habitat. The pine engraver, Ips pini, produces the pheromone ipsdienol throughout its transcontinental range. Predators of I. pini exploit this chemical as a kairomonal cue. Eastern and Midwestern I. pini populations also produce lanierone, which synergizes their attraction to ipsdienol. We evaluated the effects of varying amounts of lanierone, in combination with a constant amount of racemic ipsdienol, on the relative attraction of I. pini and its major predators in Wisconsin. Higher numbers of I. pini were captured with increasing release rates of lanierone. In contrast, the numbers of the major predators, such as Thanasimus dubius, Enoclerus nigrifrons, Platysoma cylindrica, and P. parallelum, did not differ among different lanierone release rates. The response of I. pini but not their predators to lanierone at ecologically realistic release rates may be part of a coevolving interaction between predators and prey and offers new strategies for semiochemically based pest management by selectively removing pests and leaving predators.     

Electroantennogram responses of two populations ofIps pini (Coleoptera: Scolytidae) to insect-produced and host tree compounds

Electroantennograms in two populations ofIps pini (Idaho and New York) were very similar, even though previous work documented behavioral specificity and revealed that the populations produce different ratios of the enantiomers of the aggregating pheromone ipsdienol. The insect- and host-produced compounds tested evoked characteristic response curves in both populations. EAG amplitude was highest to ipsdienol and the aggregation inhibitor ipsenol. Among the other compounds tested, responses were generally higher to the beetle-produced odorants (linalool, verbenone,trans-verbenol) than to host terpenes (1-octanol, -pinene). The antennal club and its sensory receptors are described and illustrated by scanning electron micrographs.From a thesis presented in partial fulfillment of the degree Master of Science. Department of Environmental and Forest Biology, SUNY College of Environmental Science and Forestry, Syracuse, New York 13210     

Genetic Control of the Enantiomeric Composition of Ipsdienol in the Pine Engraver, Ips pini

The genetic nature of pheromone variation within species has rarely been studied, and never for male-produced long-range pheromones. Males from western North American populations of Ips pini produce predominantly (R)-(−)-ipsdienol, whereas those from eastern North American populations produce higher proportions of (S)-(+)-ipsdienol. From a population in the hybrid zone, we divergently selected lines for the opposing pheromonal types and then created F1, F2, and backcross lines. We formed additional F1, F2, and backcross lines, first by using populations with low (+)-ipsdienol enantiomeric ratios near to and distant from the hybrid zone, and then by using populations with high (+)-ipsdienol enantiomeric ratios near to and distant from the hybrid zone. Three types of analysis were employed: (1) line means analysis; (2) Mendelian analysis of assigned high and low (+)-ipsdienol enantiomeric ratio phenotypes when applicable; and (3) Castle–Wright estimation of the number of effective factors. Dominance at one autosomal locus explains much of the variation in ipsdienol blend between the divergently selected lines. Thus, as in all previously studied female long-range pheromone systems, a major genetic element is implicated. The populations with low (+)-ipsdienol enantiomeric ratios near and distant to the hybrid zone differ negligibly for this trait. We confirmed previous studies showing slightly higher ratios of (+)-ipsdienol at the hybrid zone than in a distant eastern population and reveal a genetic basis for this difference.     

Lanierone: A new pheromone component fromIps pini (Coleoptera: Scolytidae) in New York

A new pheromone component, lanierone, (2-hydroxy-4,4,6-trimethyl-2,5-cyclohexadien-1-one) was isolated and identified from a Porapak Q collection of volatiles from maleIps pini from New York through GC fractionation, bioassay, and spectrometry. In both the laboratory and the field, synthetic lanierone, in a 1:100 ratio with synthetic ipsdienol, is as attractive as natural pheromone sources. Synthetic ipsdienol alone is not attractive in the laboratory and only weakly attractive in the field. Varying the ratio of lanierone to ipsdienol in the field from 10^–41 to 11 in 10-fold increments resulted in an increased number of beetles trapped at the three lower ratios, but also in an increase in the proportion of males trapped. In the field, all combinations of lanierone to ipsdienol attracted proportionately fewer males than did pheromone-producing male beetles. GC and GC-MS analyses of Porapak Q-trapped volatiles revealed that lanierone is produced in an amount equal to about 0.2% of that of ipsdienol and is produced exclusively by males. The small amount of lanierone produced, together with a GC retention time similar to that of ipsdienol on a nonpolar column, probably confounded its detection in earlier studies.Deceased.     

Pheromonal attraction and allomonal interruption ofIps pini in California by the two enantiomers of ipsdienol

Air containing volatile compounds from around maleIps pini boring in ponderosa pine logs from California was condensed, fractionated by GC, and assayed in the laboratory and field. The only fraction that showed consistent activity in laboratory assays contained a single compound identified as ipsdienol (2-methyl-6-methylene-2,7-octadien-4-ol). Synthetic racemic ipsdienol showed no activity in either the laboratory or field. However, (–)-ipsdienol, the naturally occurring enantiomer, was attractive toI. pini in the laboratory and field, whereas (+)-ipsdienol interrupted the response ofI. pini to a natural source of attraction in field tests. (–)-Ipsdienol is a major component of the attractant pheromone of this species, since its level of activity in laboratory assays was quantitatively comparable to that of the condensed volatiles, and it was as attractive as maleI. pini boring in ponderosa pine in the field. (+)-Ipsdienol is a component of the pheromone of the competing species,I. paraconfusus.(C.B.R.I. Contribution No. 1043.)     

The role of lanierone in the chemical ecology ofIps pini (Coleoptera: Scolytidae) in California

Five doses of lanierone (2-hydroxy-4,4,6-trimethyl-2,5-cyclohexadien-1-one) were tested with one dose of enantiomerically pure [99.4% (4R)-(–)] ipsdienol (2-methyl-6-methylene-2,7-octadien-4-ol) for activity as an aggregation pheromone ofIps pini (Say) in California. The response ofI. pini to 1 mg/day ipsdienol + 20 g/day lanierone was significantly greater than the response to ipsdienol alone, but the response pattern did not demonstrate a clear dose-response relationship. The response to the highest dose of lanierone (2 mg/day) was significantly lower than the response to ipsdienol alone. Ipsdienol attracted significantly moreI. pini than a male-infested log. Lanierone did not alter the percentage of maleI. pini responding to ipsdienol alone. Neither sex ofI. pini orDendroctonus brevicomis LeConte from California produced detectable amounts of lanierone, but myrcene-aerated maleD. brevicomis produced 97.8%-(4S)-(+)-ipsdienol. The black-bellied clerid,Enoclerus lecontei (Wolcott) (Coleoptera: Cleridae) was attracted to lanierone when released with ipsdienol. Neither compound was attractive when released alone, proving synergism for the kairomone of this predator. Lanierone did not influence the response of the predatorsTemnochila chlorodia (Mannerheim) (Coleoptera: Trogositidae) andEnoclerus sphegeus (F.) (Coleoptera: Cleridae), which were attracted to all treatments containing ipsdienol.Tomicobia tibialis Ashmead (Hymenoptera: Pteromalidae) responded in significantly greater numbers to the male-infested log than it did to ipsdienol or ipsdienol + 20 g/day lanierone.     

Antennal olfactory responsiveness of three sympatricIps species [Ips avulsus (Eichhoff),Ips calligraphus (Germar),Ips grandicollis (Eichhoff)], to intra- and interspecific behavioral chemicals

Electroantennograms (EAGs) from male and femaleIps avulsus, I. calligraphus, andI. grandicollis to their pheromones and selected host odorants or kairomones verified the presence of antennal olfactory receptors in both sexes of each species capable of detecting ipsdienol, ipsenol,cis- andtrans-verbenol,endo-brevicomin -pinene, frontalin, and verbenone. Each species possesses receptors with lower thresholds and in greater abundance for the compounds they produce and to which they are behaviorally most responsive. Detection of bothIps andDendroctonus pheromones by the three cohabiting species provides a sensory basis for olfactory interactions among the species. Differences in both threshold and saturation levels for EAGs for the various behavioral chemicals could denote differences in specific behavioral roles for each compound.Texas Agricultural Experiment Station Paper Number TA-22463. This research was supported in part by McIntire-Stennis Project 1525, USDA-CR grants 85-CRCR-1-1856 and 86-CRCR-2019 and NATO Collaborative Research Grant 86-0710. The findings, opinions, and recommendations expressed herein are those of the authors and not necessarily those of the U.S. Department of Agriculture. All programs and information of the Texas Agricultural Experiment Station are available without regard to race, ethnic origin, religion, sex, or age.Mention of a trademark, proprietary product, or vendor does not constitute a guarantee or warranty of the product by the U.S. Department of Agriculture and does not imply its approval to the exclusion of other products or vendors that may also be suitable.     

β-Phellandrene: kairomone for pine engraver,Ips pini (Say) (Coleoptera: Scolytidae)

The responses ofIps pini (Scolytidae) to multiple-funnel traps baited with the pheromone, ipsdienol, and various monoterpenes were determined in stands of lodgepole pine in southern and central British Columbia.Ips pini was attracted to both ipsdienol and -phellandrene, demonstrating that -phellandrene is a kairomone for this species.Lasconotus complex (Colydiidae) and aCorticeus sp. (Tenebrionidae) were attracted to both ipsdienol and -phellandrene. TheCorticeus sp. exhibited a synergistic response to the combination of ipsdienol and -phellandrene; the responses of the other two species to the combination were additive. The predators,Thanasimus undatulus, Enoclerus sphegeus, andE. lecontei (Cleridae), were attracted to ipsdienol-baited traps, whileMonochamus clamator (Cerambycidae) andDendroctonus ponderosae (Scolytidae) were attracted to -phellandrene. Attraction of all eight species increased with increasing release rates of ipsdienol and/or -phellandrene.