<I>Dendroctonus Jeffreyi</I>
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An Attractant Isolated from Female Jeffrey Pine Beetles, Dendroctonus jeffreyi1 J. A. A. RENWICK AND G. B. PITMAN Boyce Thompson Institute, Tower Rd., Ithaca, NY ]4853 ABSTRACT Environ. Entomol. 8: 40-41 (1979) The aggregation pheromone of Dendroctonus jeffreyi was investigated by combined gas chromatography and mass spectrometry of volatiles from hindguts of attacking beetles. The major compounds in females were identified as I-heptanol and 2-heptanol, which are pro- duced by oxidation of heptane present in the host resin. In field tests, I-heptanol in combi- nation with heptane was attractive, but racemic 2-heptanol was not. The addition of 2-hep- tanol to I-heptanol appeared to reduce the number of beetles responding. The results indicate a probable difference between the chemical communication systems of D. jeffreyi and D. ponderosae, both of which have apparently adapted to utilize the met- abolic products of potentially toxic hydrocarbons in their hosts. Downloaded from https://academic.oup.com/ee/article/8/1/40/2396242 by guest on 01 October 2021 The Jeffrey pine beetle, DendroclOnus jeffreyi Hopk, near Donner Lake, California. The beetles were imme- is a pest of Jeffrey pines throughout their range, which diately frozen at - 70°C and stored in a freezer for sub- extends from southern Oregon through California and sequent dissection and analysis. Sex determination was western Nevada to northern Baja California. This beetle initially based on the positions of the beetles in the gal- closely resembles the mountain pine beetle, D. ponde- leries and later confirmed during dissection. rosae Hopk., both in morphology and behavior, and the Gas chromatographic (GC) and mass spectrometric 2 species were synonymized, along with D. monticolae (MS) analyses were performed as previously described Hopk., by S. L. Wood (1963). Although observations (Hughes et a!. 1976). Preliminary GC patterns were ob- on the larval anatomy supported this synonymy (Thomas tained using headspace samples from hindguts, whereas 1965), physiological differences were reported in the re- ether extracts were prepared for the GC-MS analyses. action of D. jeffreyi and D. ponderosae to pine resin Field Bioassays vapors and their components, thus leading Smith (1965) Two areas were chosen in the Central Sierra for field to continue to distinguish between them. Controlled mating experiments along with observations on devel- testing of synthetic chemicals. The 1st site was at the North Star ski area near Lake Tahoe and the 2nd was opment rate, karyology and morphology were subse- quently used by Lanier and D. L. Wood (1968) to pos- close to the north shore of Donner Lake. In both areas, periodic Jeffrey pine beetle activity had been observed itively establish the species integrity of D. jeffreyi. The aggregation pheromone involved in the mass at- in preceding years. The trapping devices were sticky panels consisting of tack of D. ponderosae on white pine in Idaho was iden- large plastic-coated envelopes mounted on wooden stakes tified as trans- verbenol (Pitman and Vite 1969), which (Pitman et aI. 1975). The traps were arranged at the cor- the beetles produce by oxidation of a-pinene present in ners of squares of ca. IO-m sides. Test chemicals were the host resin (Hughes 1973a,b). The response of Jef- released from vials fitted with polyethylene stoppers frey pine beetles to trans- verbenol was tested in an area where a beetle population was known to be active, but which were perforated at the sides. Responding beetles were removed from the traps every few days for a period no D. jeffreyi were caught on the traps (Pitman, unpub- of 6 wk and their sex determined by dissection. lished results). The chemicals used were obtained from Aldrich The volatile constituents of resin from white pine and Jeffrey pine are quite different. In white pine, a-pinene Chemical Company. The purity of the 2-heptanol was 96%, and both the heptane and I-heptanol were 99% is the most abundant hydrocarbon, whereas in Jeffrey pure. pine, heptane constitutes ca. 90% of the volatile fraction of the resin (Mirov 1961). These differences in chemis- Results try of the host trees, along with the lack of response of D. jeffreyi to trans-verbenol, would strongly suggest Chemical Analysis that the 2 beetle species use different systems of chemi- Gas chromatograms of hindgut volatiles from attack- cal communication, and prompted the study reported ing female beetles revealed the presence of 2 major con- here to elucidate the chemical basis for aggregation of stituents which were relatively minor components in the Jeffrey pine beetle. samples from males. The mass spectra of these com- pounds indicated that they were 2-heptanol and I-hep- Materials and Methods tanol, and authentic samples of the alcohols had identi- Analysis of Hindgut Volatiles cal GC retention times and mass spectra. Attacking D. jeffreyi were removed from pitch tubes Field Bioassays and initial galleries in naturally infested Jeffrey pines Each of the identified heptanols was tested alone and in combination. In all cases, heptane was included along I Received for publication July 17. 1978. with the alcohols to provide a possible host factor. No 40 © 1979 Entomological Society of America 0046-225X/79/0 100-4002$00. 75/0 February 1979 RENWICK AND PITMAN: D. jeffreyi PHEROMONE 41 response was obtained to the heptane control or to the jeffreyi has developed the ability to utilize the I-heptano] combination of 2-heptanol with heptane, but traps baited as a pheromone. At the same time. some degree of sex with l-heptanol consistently caught beetles throughout specificity in the oxidation of heptane has been intro- the test period. Traps baited with 2-heptano] along with duced. so that production of the pheromone is primarily I-heptanol attracted fewer beetles than those with] -hep- restricted to the females. tanol alone (Table I). These results indicate probable differences between the chemical communication systems of D. jeffreyi and Discussion D. ponderosae. thus supporting the conclusions of Lan- The results clearly indicate that I-heptanol is a pher- ier and Wood (1968) that these beetles represent quite omone involved in the aggregation of D. jeffreyi on its distinct species. In both cases. it appears that adaptation host trees. Although the population of beetles in the test to particular hosts has involved metabolism of the poten- areas was extremely low, this alcohol consistently at- tially toxic hydrocarbons encountered and utilization of tracted both males and females. On the other hand, 2- the metabolic products as chemical messengers. heptanol was completely inactive as an attractant, and Downloaded from https://academic.oup.com/ee/article/8/1/40/2396242 by guest on 01 October 2021 may even have reduced the response to the active iso- REFERENCES CITED mer. The reason for this apparent inhibition is not clear, Hughes, P. R. 1973a. Effect of a-pinene exposure on trans- especially since the attacking female beetles produce al- verbenol synthesis in Dendroctonus ponderosae Hopk. most equal amounts of both geometrical isomers. How- Naturwissenschaften 60: 261-2. ever, the possibility exists that the beetles biosynthesize ]973b. Dendroctonus: Production of pheromones and related compounds in response to host monoterpenes. Z. Ang. only one enantiomer of the 2-heptanol, and the other EntomoL 73: 294-312. enantiomer could inhibit the response. This phenomenon Hughes, P. R., J.A.A. Renwick, and J. P. Vile. 1976. The has been observed recently in other bark beetles (Vite et identification and field bioassay of chemical attractants in al. 1976. 1978). The 2.heptanol used in our tests was a the roundheaded pine beetle. Environ. Entomol. 5: 1]65- racemic mixture. 8. I-Heptanol and 2-heptanol have been identified pre- Lanier, G. N., and D. L. Wood. 1968. Controlled mating, viously from hindguts of Dendroctonus vitei Wood at- karyology, morphology. and sex-ratio in the Dendroc- tacking Pinus ma.ximinoi Moore, the resin of which has tonus ponderosae complex. Ann. Entomol. Soc. Am. 61: 517-26. a relatively high heptane content. The heptanols are pro- Mirov, N. T. 1961. Composition of gum turpentines of pines. duced by both sexes of this species. but the key pher- Tech. Bull. USDA 1239: ]-158. omonal compound is frontalin. which is produced by Pitman, G. B., and J. P. Vile. ]969. Aggregation behavior of the females only. The heptanols in D. vitei may have Dendroctonus ponderosae (Coleoptera: Scolytidae) in re- some sex-regulating function in the response to frontalin sponse to chemica] messengers. Can. Entomol. ]0]: 143- (Renwick et al. 1975). It is likely that many bark beetles 9. are capable of oxidizing heptane to the alcohols. but D. Pitman, G. B., R. L. Hedden, and R. I. Gara. 1975. Syn- ergistic effects of ethyl alcohol on the aggregation of Den- droctonus pseudotsugae (Col.. Scolytidae) in response to pheromones. Z. Ang. Entomol. 78: 203-8. Table I.-Field response of Jeffrey pine beetles to hep- Renwick, J.A.A., P. R. Hughes, and J. P. Vite. 1975. The tane, l.heptanol and 2-heptanol, Central Sierra. June- aggregation pheromone system of a Dendroctonus bark Aug. 1976." beetle in Guatemala. J. Insect Physiol. 21: 1097-100. Smith, R. H. 1965. A physiological difference among beetles North Star Donner Lake of Dendroctonus ponderosae (= D. mOlllicolae) and D. ponderosae (= D. jeffreyi ). Ann. Entomol. Soc. Am. 58: Total no. Sex Total no. Sex 440-2. Test D. jeffreyi ratio D. jeffreyi ratio material caught o:'l' caught o:'l' Thomas, J. B. 1965. The immature stages of Scolytidae: the genus Dendroctonus Erichson. Can. Entomol. 97: 374- Heptane o o 400. Heptane + Vite, J. P., R. Hedden, and K. Mori. ]976./ps grandicollis: I-heptanol 56 ]:0.93 36 1:1.25 field response to the optically pure pheromone.