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MALE SCENT IN LEPIDOPTERAN COMMUNICATION: THE ROLE OF MALE PHEROMONE IN MATING BEHAVIOUR OF PSEUDALETIA UNIPUNCTA (HAW.) (LEPIDOPTERA: NOCTUIDAE)

Published online by Cambridge University Press:  31 May 2012

Sheila M. Fitzpatrick  and
Jeremy N. McNeil
Sheila M. Fitzpatrick
Affiliation:
Département de biologie, Université Laval, Ste-Foy, Québec, Canada G1K 7P4
Jeremy N. McNeil
Affiliation:
Département de biologie, Université Laval, Ste-Foy, Québec, Canada G1K 7P4
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Abstract

Pseudaletia unipuncta (Haw.) males, experimentally deprived of hairpencils (male-specific scent organs), were less acceptable to females than were intact males. The degree to which females discriminated against males lacking hairpencils depended on the experimental protocol. When females were placed with males before scotophase onset, they tended to evade repeatedly and/or terminate genital contact with hairpencil-less males before eventually copulating, although this tendency was not statistically significant. When males were introduced after females had initiated calling, females refused hairpencil-less males significantly more often than intact ones. The incidence of copulation by hairpencil-less males was also lower than that of intact males, significantly so in one of the two populations tested. Neither copulation duration nor the dimensions of the cuticular portion of the spermatophore were correlated with the presence of hairpencils. Yet in all cases, females that mated with hairpencil-less males were more fecund and laid more fertile eggs than those mated to intact males. Hairpencil eversion could not be seen during courtship, nor did females show any obvious change in calling behaviour in response to scent from a freshly excised hairpencil. However, gas chromatographic analysis of hairpencils showed a decrease in titre of two scent components, acetic acid and benzaldehyde, just before the moment of genital contact between male and female, indicating that the pheromone was released at this time. These results suggest that P. unipuncta male pheromone facilitates female acceptance of males. The hypothesis that male pheromone carries information about the quality of a courting male is discussed in relation to recent work on arctiids, nymphalids, pierids and pyralids. An appendix of lepidopteran species possessing male scent structures is included.

Résumé

Les femelles de Pseudaletia unipuncta (Haw.) ont préféré des mâles intacts à des mâles dont les pinceaux androconiaux (organes odoriférants spécifiques aux mâles) avaient été extirpés. Le niveau de discrimination exprimé par les femelles envers les mâles sans pinceaux dépendait du protocole expérimental. Lorsque des femelles ont été placées en présence de mâles avant le début de la scotophase, celles-ci ont eu tendance à éviter et/ou terminer les contacts génitaux avant d'en arriver éventuellement à la copulation. Cependant, cette tendance ne fut pas significative. Mais lorsque les mâles furent introduits après que les femelles aient initié l'appel, ces dernières ont refusé les mâles dépourvus de pinceaux significativement plus souvent que les mâles intacts. Le nombre de copulations entreprises par des mâles sans pinceaux fut également inférieur à celui de mâles intacts, et ce de façon significative dans une des deux populations testées. Ni la durée de la copulation, ni les dimensions de la portion cuticulaire des spermatophores furent corrélées, avec la présence de pinceaux. Et pourtant dans tous les cas, les femelles accouplées à des mâles sans pinceaux furent plus fécondes et déposèrent plus d'oeufs fertiles que celles accouplées à des mâles intacts. Le déploiement des pinceaux au moment de l'approche n'a pas pu être observé et les femelles n'ont pas semblé modifier leur comportement d'appel en réponse à l'odeur de pinceaux fraîchement prélevés. Cependant, la chromatographie en phase gazeuse a révélé une diminution du titre de deux des composantes de la phéromone des mâles, l'acide acétique et le benzaldéhyde, juste avant le contact génital mâle-femelle, indiquant que la phéromone est relâchée à ce moment. Ces résultats suggèrent que la phéromone des mâles de P. unipuncta facilite l'acceptation de ceux-ci par les femelles. L'hypothèse selon laquelle la phéromone des mâles constitue, pour les femelles, une source d'information relative à la qualité des mâles, est discutée en relation avec certains ouvrages récents sur les Arctiidae, Nymphalidae, Pieridae et Pyralidae. Une liste des espèces de Lépidoptères dont les mâles possèdent des structures androconiales est inclue en annexe.

Type
Research Article
Information
The Memoirs of the Entomological Society of Canada , Volume 120 , Supplement S146 , 1988 , pp. 131 - 151
Copyright
Copyright © Entomological Society of Canada 1988

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References

Agee, H.R. 1969. Mating behavior of bollworm moths. Ann. ent. Soc. Am. 62: 1120–1122.Google Scholar
Aplin, R.T., and Birch, M.C.. 1970. Identification of odorous compounds from male Lepidoptera. Experientia 26: 1193–1194.Google Scholar
Atkinson, P.R. 1982. Structure of the putative pheromone glands of Eldana saccharina Walker (Lepidoptera: Pyralidae). J. ent. Soc. S. Afr. 45: 93–104.Google Scholar
Baker, T.C., and Cardé, R.T.. 1979. Courtship behavior of the oriental fruit moth (Grapholitha molesta): experimental analysis and consideration of the role of sexual selection in the evolution of courtship pheromones in the Lepidoptera. Ann. ent. Soc. Am. 72: 173–188.Google Scholar
Baker, T.C., Nishida, R., and Roelofs, W.L.. 1981. Close range attraction of female oriental fruit moths to herbal scent of hairpencils. Science 214: 1359–1361.Google Scholar
Barrer, P.M., and Hill, R.J.. 1978. The acceptance response of Ephestia cautella (Walker) (Lepidoptera: Phycitidae) females, obtained in the absence of courting males. Experientia 34: 343–344.Google Scholar
Barth, R. 1958. Sobre o glandular do aparelho copulador do macho de Prodenia ornithogalli Gn. (Lepidoptera: Noctuidae). Mem. Inst. Oswaldo Cruz 51: 187–202; cited in Grant 1971a.Google Scholar
Barth, R. 1960. Orgaños odóriferos dos Lepidópteros. Parque Nac. Itatiaia, Rio de Janeiro, Bull. 7. 159 pp.; cited in Grant 1978.Google Scholar
Benz, G., and Schmid, K.. 1968. Stimulation der Eiablage unbegatter weibchen des Mondspinners Actias selene durch ein Mannchen-Pheromon. Experientia 24: 1279–1281.Google Scholar
Bergstrom, G., and Lundgren, L.. 1973. Androconial secretion of three species of butterflies of the genus Pieris (Lepidoptera, Pyralidae). ZOON Suppl. 1: 67–75.Google Scholar
Bijpost, S.C.A., Thomas, G., and Kruijt, J.P.. 1985. Olfactory interactions between sexually active males in Adoxophyes orana (F.v.R.) (Lepidoptera: Tortricidae). Behav. 95: 121–137.Google Scholar
Birch, M.C. 1970a. Pre-courtship use of abdominal brushes by the nocturnal moth, Phlogophora meticulosa (L.) (Lepidoptera: Noctuidae). Anim. Behav. 18: 310–316.Google Scholar
Birch, M.C. 1970b. Persuasive scents in moth life. Nat. Hist. 79: 34–39.Google Scholar
Birch, M.C. 1972. Male abdominal brush-organs in British noctuid moths and their value as a taxonomic character. Part II. Entomology 105: 233–244.Google Scholar
Birch, M.C. 1974. Aphrodisiac pheromones in insects, pp. 115–134in Birch, M.C. (Ed.), Pheromones. North-Holland, Amsterdam.Google Scholar
Birch, M.C. 1979. Eversible structures, pp. 9–18in Heath, J., and Emmet, A.M. (Eds.), Moths and Butterflies of Great Britain and Ireland. Curwen, London; cited in Birch and Hefetz 1987.Google Scholar
Birch, M.C., and Hefetz, A.. 1987. Eversible organs in male moths and their role in courtship behavior. Bull. ent. Soc. Am. 33: 222–229.Google Scholar
Boppré, M. 1984. Chemically mediated interactions between butterflies, pp. 259–275in Vane-Wright, R.I., and Ackery, P.R. (Eds.), The Biology of Butterflies, Symposium of the Royal Entomological Society of London, 11. Academic Press, London.Google Scholar
Boppré, M. 1986. Insects pharmacophagously using defensive plant chemicals (pyrrolizidine alkaloids). Naturw. 73: 17–26.Google Scholar
Boppré, M., Petty, R.L., Schneider, D., and Meinwald, J.. 1978. Behaviorally mediated contacts between scent organs: another prerequisite for pheromone production in Danaus chrysippus males (Lepidoptera). J. Comp. Physiol. A. 126: 97–103.Google Scholar
Brower, L.P., and Jones, M.A.. 1965. Precourtship interaction of wing and abdominal sex glands in male Danaus butterflies. Proc. R. ent. Soc. Lond. A 40: 147–151.Google Scholar
Brower, L.R., van Zandt Brower, J., and Cranston, F.R. 1965. Courtship behavior of the queen butterfly, Danaus gilippus berenice (Cramer). Zool. 50: 1–39.Google Scholar
Brown, K.S. 1984. Adult-obtained pyrrolizidine alkaloids defend ithomiine butterflies against a spider predator. Nature 309: 707–709.Google Scholar
Callahan, P.S., and Chapin, J.B.. 1960. Morphology of the reproductive systems and mating in 2 representative members of the family Noctuidae, Pseudaletia unipuncta and Peridroma margaritosa, with comparison to Heliothis zea. Ann. ent. Soc. Am. 53: 763–782.Google Scholar
Clearwater, J.R. 1971. The role of the male produced pheromone in the reproduction behaviour of the southern army worm, Pseudaletia separata (Walk.). M.Sc. thesis, Massey University, New Zealand.Google Scholar
Clearwater, J.R. 1972. Chemistry and function of a pheromone produced by the male of the southern army worm, Pseudaletia separata. J. Insect Physiol. 18: 781–789.Google Scholar
Clearwater, J.R. 1975a. Pheromone metabolism in male Pseudaletia separata (Walk.) and Mamestra configurata (Walk.) (Lepidoptera: Noctuidae). Comp. Biochem. Physiol. 50B: 77–82.Google Scholar
Clearwater, J.R. 1975b. Structure development and evolution of the male pheromone system in some Noctuidae (Lepidoptera). J. Morph. 146: 129–176.Google Scholar
Connor, W.E., Eisner, T., Vander Meer, R.K., Guerrero, A., and Meinwald, J.. 1981. Precopulatory sexual interaction in an arctiid moth (Utetheisa ornatrix): role of a pheromone derived from dietary alkaloids. Behav. Ecol. Sociobiol. 9: 227–235.Google Scholar
Corbet, S.A., and Lai-Fook, J.. 1977. The hairpencils of the flour moth, Ephestia kuehniella. J. Zool. Lond. 181: 377–394.Google Scholar
Culvenor, C.C.J., and Edgar, J.A.. 1972. Dihydropyrrolizine secretions associated with coremata of Utetheisa moths (family Arctiidae). Experientia 28: 627–628.Google Scholar
Dahm, K.H., Meyer, D., Finn, W.E., Reinhold, V., and Röller, H.. 1971. The olfactory and auditory mediated sex attraction in Achroia grisella (Fabr.). Naturw. 58: 265–266.Google Scholar
Dickins, G.R. 1936. The scent glands of certain Phycitidae (Lepidoptera). Trans. R. ent. Soc. Lond. 85: 331–362; cited in Weatherston and Percy 1969.Google Scholar
Edgar, J.A. 1982. Pyrrolizidine alkaloids sequestered by Solomon Island Danaine butterflies. The feeding preferences of the Danainae and Ithomiinae. J. Zool. Lond. 196: 385–399.Google Scholar
Edgar, J.A., Culvenor, C.C.J., and Smith, L.W.. 1971. Dihydropyrrolizine derivatives in the ‘hair-pencil’ secretions of danaid butterflies. Experientia 27: 761–762.Google Scholar
Edgar, J.A., Culvenor, C.C.J., and Pliske, T.E.. 1976. Isolation of a lactone, structurally related to the esterifying acids of pyrrolizidine alkaloids, from the costal fringes of male Ithomiinae. J. Chem. Ecol. 2: 263–270.Google Scholar
Ellis, P.E., and Brimacombe, L.C.. 1980. The mating behaviour of the Egyptian cotton leafworm moth, Spodoptera littoralis (Boisd.). Anim. Behav. 28: 1239–1248.Google Scholar
Eltringham, H. 1913. On the scent apparatus in the male of Amauris niavius Linn. Trans, ent. Soc. Lond. 1913: 399–406; cited in Brower et al. 1965.Google Scholar
Eltringham, H. 1915. Further observations on the structure of the scent organs in certain male Danaine butterflies. Trans. ent. Soc. Lond. 1915: 152–176; cited in Brower et al. 1965.Google Scholar
Eltringham, H. 1925. On the abdominal brushes of certain male moths. Trans. R. ent. Soc. Lond. 1–5; cited in Weatherston and Percy 1969.Google Scholar
Farine, J.-R. 1982. Les glandes exocrines male and femelle de Pseudaletia unipuncta Haw. (Lepidoptera: Noctuidae) et leur rôle dans le comportement précopulatoire des deux sexes. Agronomie 2: 459–468.Google Scholar
Finn, W.E., and Payne, T.L.. 1977. Attraction of greater wax moth females to male-produced pheromones. Southwest. Ent. 2: 62–64.Google Scholar
Fitzpatrick, S.M., McNeil, J.N., and Miller, D.. 1989. Age-specific titer and antennal perception of acetic acid, a component of male Pseudaletia unipuncta (Haw.) hairpencil secretion. J. Chem. Ecol., in press.Google Scholar
Fitzpatrick, S.M., Miller, D., Weatherston, I., and McNeil, J.N.. 1985. Determining pheromone content of hair-pencils from individual virgin males of Pseudaletia unipuncta (Haw.) (Lepidoptera: Noctuidae). J. Chem. Ecol. 11: 207–215.Google Scholar
George, J.A. 1965. Sex pheromone of the Oriental fruit moth Grapholitha molesta (Busck) (Lepidoptera: Tortricidae). Can. Ent. 97: 1002–1007.Google Scholar
Gilbert, D.G. 1981. Ejaculate esterase 6 and initial sperm use by female Drosophila melanogaster. J. Insect Physiol. 27: 641–650.Google Scholar
Gilbert, L.E. 1976. Postmating female odor in Heliconius butterflies; a male-contributed antiaphrodisiac? Science 193: 419–420.Google Scholar
Gothilf, S., and Shorey, H. H.. 1976. Sex pheromones in the Lepidoptera: examination of the role of male scent brushes in courtship behavior of Trichoplusia ni. Environ. Ent. 5: 115–119.Google Scholar
Grant, G.G. 1970. Evidence for a male sex pheromone in the noctuid, Trichoplusia ni. Nature 227: 1345–1346.Google Scholar
Grant, G.G. 1971 a. Scent apparatus of the male cabbage looper, Trichoplusia ni. Ann. ent. Soc. Ann. 64: 347–352.Google Scholar
Grant, G.G. 1971 b. Electroantennogram responses to the scent brush secretions of several male moths. Ann. ent. Soc. Am. 64: 1428–1431.Google Scholar
Grant, G.G. 1976. Courtship behavior of a phycitid moth, Vitula edmandsae. Ann. ent. Soc. Am. 69: 445–449.Google Scholar
Grant, G.G. 1978. Morphology of the presumed male pheromone glands on the forewings of tortricid and phycitid moths. Ann. ent. Soc. Am. 71: 423–431.Google Scholar
Grant, G.G., and Brady, U.E.. 1975. Courtship behavior of phycitid moths. 1: comparison of Plodia interpunctella and Cadra cautella and the role of male scent glands. Can. J. Zool. 53: 813–826.Google Scholar
Grant, G.G., Brady, U.E., and Brand, J.M.. 1972. Male armyworm scent brush secretion: identification and electroantennogram study of major components. Ann. ent. Soc. Am. 65: 1224–1227.Google Scholar
Grant, G.G., and Eaton, J.L.. 1973. Scent brushes of the male tobacco hornworm, Manduca sexta (Lepidoptera: Sphingidae). Ann. ent. Soc. Am. 66: 901–904.Google Scholar
Grula, J.W., McChesney, J.D., and Taylor, O.R. Jr., 1980. Aphrodisiac pheromones of the sulfur butterflies Colias eurytheme and C. philodice (Lepidoptera, Pieridae). J. Chem. Ecol. 6: 241–256.Google Scholar
Haberman, S.J. 1973. The analysis of residuals in cross-classified tables. Biometrics 29: 205–220;Google Scholar
cited in Everitt, B.S., 1977, The Analysis of Contingency Tables, Chapman and Hall, London.Google Scholar
Haynes, K.F., and Birch, M.C.. 1985. The role of other pheromones, allomones and kairomones in the behavioral responses of insects, pp. 225–255in Kerkut, G.A., and Gilbert, L.I. (Eds.), Comprehensive Insect Physiology, Biochemistry and Pharmacology, Vol. 9. Pergamon Press, London.Google Scholar
Heinrich, C. 1956. American moths of the subfamily Phycitinae. U.S. Nat. Mus. Bull. 207. 249 pp.; cited in Grant 1978.Google Scholar
Hendricks, D.E., and Shaver, T.N.. 1975. Tobacco budworm: male pheromone suppressed emission of sex pheromone by the female. Environ. Ent. 4: 555–558.Google Scholar
Hendrikse, A., van der Laan, C.E., and Kerkof, L.. 1984. The role of male abdominal brushes in the sexual behaviour of small ermine moths (Yponomeuta Latr., Lepidoptera). Med. Fac. Landbouww. Rijksuniv. Gent. 49: 719–726; cited in G.G. Grant personal communication.Google Scholar
Hirai, K. 1980. Male scent emitted by armyworms, Pseudaletia unipuncta and P. separata (Lepidoptera: Noctuidae). Appl. Ent. Zool. 15: 310–315.Google Scholar
Hirai, K. 1981. Morphology of the brush organs of the limabean pod borer, Etiella zinckenella Treitschke (Lepidoptera: Pyralidae). Appl. Ent. Zool. 16: 362–366.Google Scholar
Hirai, K. 1982. Directional flow of male scent released by Pseudaletia separata Walker (Lepidoptera: Noctuidae) and its repellent effect on adults and larvae of four noctuid and one phycitine moth. J. Chem. Ecol. 8: 1263–1270.Google Scholar
Hirai, K., Shorey, H.H., and Gaston, L.K.. 1978. Competition among courting male moths: male-to-male inhibitory pheromone. Science 202: 644–645.Google Scholar
Honda, K. 1980. Odor of a papilionid butterfly. Odoriferous substances emitted by Atrophaneura alcinous alcinous (Lepidoptera: Papilionidae). J. Chem. Ecol. 6: 867–873.Google Scholar
Jacobsen, M., Adler, V.E., Kishaba, A.N., and Priesner, E.. 1976. 2-phenylethanol, a presumed sexual stimulant produced by the male cabbage looper moth, Trichoplusia ni. Experientia 32: 964–966.Google Scholar
Komae, H., Nishi, A., Tanaka, T., Hayashi, N., Wesou, C., and Kuwahara, Y.. 1982. Major components in the hairpencil secretions of danaid butterflies from Far East Asia. Biochem. System. Ecol. 10: 181–183.Google Scholar
Krasnoff, S.B., and Vick, K.W.. 1984. Male wing-gland pheromone of Ephestia elutella. J. Chem. Ecol. 10: 667–679.Google Scholar
Leopold, R.A. 1976. The role of male accessory glands in insect reproduction. A. Rev. Ent. 21: 199–222.Google Scholar
Lundgren, L., and Bergstrom, G.. 1975. Wing scents and scent-released phases in the courtship behavior of Lycaeides argyognomon (Lepidoptera: Lycaenidae). J. Chem. Ecol. 1: 399–412.Google Scholar
Mallet, J. 1984. Sex roles in the ghost moth, Hepialus humuli (L.) and a review of mating in the Hepialidae (Lepidoptera). Zool. J. Linn. Soc. 79: 67–82.Google Scholar
Marshall, L.D. 1985. Protein and lipid composition of Colias philodice and C. eurytheme spermatophores and their changes over time (Pieridae). J. Res. Lepid. 24: 21–30.Google Scholar
McLaughlin, J.R. 1982. Behavioral effect of a sex pheromone extracted from forewings of male Plodia interpunctella. Environ. Ent. 11: 378–380.Google Scholar
Meinwald, J., Boriack, C.J., Schneider, D., Boppré, M., Wood, W.F., and Eisner, T.. 1974. Volatile ketones in the hairpencil secretion of danaid butterflies (Amauris and Danaus). Experientia 30: 721–722.Google Scholar
Meinwald, J., Meinwald, Y.C., Wheeler, J.W., Eisner, T., and Brower, L.P.. 1966. Major components in the exocrine secretion of a male butterfly (Lycorea). Science 151: 583–585.Google Scholar
Meinwald, J., Thompson, W.R., Eisner, T., and Owen, D.F.. 1971. Pheromones. 7: African monarch: major components of hairpencil secretion. Tetrahedron Lett. 1971: 3485–3488.Google Scholar
Müller, F. 1877. Úber Haarpinsel, Filzflecke und ahnliche Gebilde auf den Flügeln mannlicher Schmetterlinge. Jena. Z. Naturw. 5: 99–114; cited in Boppré 1984.Google Scholar
Myers, J.H. 1972. Pheromones and courtship behavior in butterflies. Am. Zool. 12: 545–551.Google Scholar
Myers, J., and Brower, L.P.. 1969. A behavioral analysis of the courtship pheromone receptors of the queen butterfly, Danaus gilippus berenice. J. Insect Physiol. 15: 2117–2130.Google Scholar
Palaniswamy, P., Seabrook, W.D., and Ross, R.J.. 1979. Precopulatory behavior of males and perception of a potential male pheromone in spruce budworm, Choristoneura fumiferana. Ann. ent. Soc. Am. 72: 544–551.Google Scholar
Peterson, W. (1907) 1908. Die Arberechtigung von Miana latruncula Hb. (Lepidoptera: Noctuidae). Revue russe d'entomologie 7: 206–210; cited in Birch 1972.Google Scholar
Phelan, P.L., and Baker, T.C.. 1986. Male-size-related courtship success and intersexual selection in the tobacco moth, Ephestia elutella. Experientia 42: 1291–1293.Google Scholar
Phelan, P.L., and Baker, T.C.. 1987. Evolution of male pheromones in moths: reproductive isolation through sexual selection? Science 235: 205–207.Google Scholar
Phelan, P.L., Silk, P.J., Northcott, C.J., Tan, S.H., and Baker, T.C.. 1986. Chemical identification and behavioral characterization of male wing pheromone of Ephestia elutella (Pyralidae). J. Chem. Ecol. 12: 135–146.Google Scholar
Pierce, F.N. 1909. The genitalia of the Group Noctuidae of the Lepidoptera of the British Islands. Liverpool; cited in Birch 1972.Google Scholar
Pivnick, K.P., and McNeil, J.N.. 1985. Mate location and mating behavior of Thymelicus lineola (Lepidoptera: Hesperiidae). Ann. ent. Soc. Am. 78: 651–656.Google Scholar
Pliske, T.E. 1975a. Courtship behavior of the Monarch butterfly, Danaus plexippus (L.). Ann. ent. Soc. Am. 68: 143–151.Google Scholar
Pliske, T.E. 1975b. Courtship behavior and use of chemical communication by males of certain species of Ithomiine butterflies (Nymphalidae: Lepidoptera). Ann. ent. Soc. Am. 68: 935–942.Google Scholar
Pliske, T.E., and Eisner, T.. 1969. Sex pheromone of the queen butterfly: biology. Science 164: 1170–1172.Google Scholar
Redfern, R.E., Cutt, B.A., and Cantu, E.. 1970. Bioassay of the sex pheromone of the southern armyworm. J. econ. Ent. 63: 658.Google Scholar
Richards, O.W., and Thomson, W.S.. 1932. A contribution to the study of the genera Ephestia Gn. (including strymax, Dyar), and Plodia Gn. (Lepidoptera: Phycitidae), with notes on parasites of the larvae. Trans. R. ent. Soc. Lond. 80: 169; cited in Barrer and Hill 1978.Google Scholar
Rutowski, R.L. 1977. Chemical communication in the courtship of the small sulfur butterfly Eurema lisa (Lepidoptera: Pieridae). J. Comp. Physiol. A 115: 75–85.Google Scholar
Rutowski, R.L. 1980. Male scent-producing structures in Colias butterflies. Function, localization and adaptive features. J. Chem. Ecol. 6: 13–26.Google Scholar
Schneider, D., Boppré, M., Zweig, J., Horsley, S.B., Bell, T.W., Meinwald, J., Hansen, K., and Diehl, E.W.. 1982. Scent organ development in Creatonotos moths: regulation by pyrrolizidine alkaloids. Science 215: 1264–1265.Google Scholar
Seabrook, W.D., Hirai, K., Shorey, H.H., and Gaston, L.K.. 1979. Maturation and senescence of an insect chemosensory response. J. Chem. Ecol. 5: 587–594.Google Scholar
Sellier, R. 1972. Etude structurale en microscopie électronique à balayage et essai d'interprétation du mode de fonctionnement des poils androconiaux alaires chez les Hesperiidae. C.R. Acad. Sci. Ser. D 275: 2239–2242.Google Scholar
Shorey, H.H. 1964. Sex pheromones of noctuid moths. II: Mating behavior of Trichoplusia ni with special reference to the role of the sex pheromone. Ann. ent. Soc. Am. 57: 371–377.Google Scholar
Shorey, H.H., and Hale, R.L.. 1965. Mass rearing of nine noctuid species on a simple artificial medium. J. econ. Ent. 58: 522–524.Google Scholar
Sokal, R.R., and Rohlf, F.J.. 1981. Biometry. W.H. Freeman and Co., New York, U.S.A.Google Scholar
Stobbe, R.H. 1912. Die abdominalen Duftorgane der mannlichen Sphingiden und Noctuiden. Zool. Jb. 32: 493–532; cited in Birch 1972.Google Scholar
Swinton, A.H. 1908–1909. The family tree of moths and butterflies traced in their organs of sense. Societas ent. 23: 99–101, 114–116, 124–126, 131–132, 140–141, 148–150, 156–157, 162–165; cited in Birch 1972.Google Scholar
Szentesi, A., Tóth, M., and Dobrovolszky, A.. 1975. Evidence and preliminary investigations on a male aphrodisiac and a female sex pheromone in Mamestra brassicae (L.). Acta Phytopathol. Acad. Sci. Hung. 10: 425–429.Google Scholar
Tamaki, Y. 1985. Sex pheromones. pp. 145–191in Kerkut, G.A., and Gilbert, L.I. (Eds.), Comprehensive Insect Physiology, Biochemistry and Pharmacology, Vol. 9. Pergamon Press, London.Google Scholar
Teal, P.E.A., McLaughlin, J.R., and Tumlinson, J.H.. 1981. Analysis of the reproductive behavior of Heliothis virescens (F.) under laboratory conditions. Ann. ent. Soc. Am. 74: 324–330.Google Scholar
Thibout, E. 1978. Rôle des phéromones males et des corema dans le comportement sexuel précopulatoire d'Acrolepiopsis (Acrolepia) assectella Zell. (Hyponomeutoidea). C.R. Acad. Sci. Paris Ser. D 287: 1141–1144.Google Scholar
Thibout, E. 1979. Stimulation of reproductive activity of females of Acrolepiopsis assectella (Lepidoptera: Hyponomeutoidea) by the presence of eupyrene spermatozoa in the spermatheca. Ent. Exp. Appl. 26: 279–290.Google Scholar
Varley, G.C. 1962a. On the variations in scent distribution organs in the genus Leucania (Lepidoptera: Noctuidae). Proc. R. ent. Soc. Lond. C 26: 36; cited in Birch 1972.Google Scholar
Varley, G.C. 1962b. A plea for a new look at Lepidoptera with special reference to the scent distributing organs of male moths. Trans. Soc. Br. Ent. 15: 3–40; cited in Birch 1972.Google Scholar
Wagner, D.L. 1985. The biosystematics of Hepialus F.s. lato, with special emphasis on the californicus-hec-toides species group. Ph.D. dissertation, Univ. Calif., Berkeley.Google Scholar
Wago, H. 1978a. Studies on the mating behavior of the pale grass blue, Zizeeria maha argia (Lepidoptera: Lycaenidae). 3: olfactory cues used in sexual discrimination by males. Appl. Ent. Zool. 13: 283–289.Google Scholar
Wago, H. 1978b. Studies on the mating behavior of the pale grass blue, Zizeeria maha argia Lepidoptera: Lycaenidae). 4: experimental analysis of the role of male odor in male-male interactions. Zool. Mag. 87: 240–246.Google Scholar
Wakamura, S. 1977. Sound protection of the male moth Agrostis fucosa Butler (Lepidoptera: Noctuidae) during courtship behavior. Appl. Ent. Zool. 12: 202–203.Google Scholar
Weatherston, J., and Percy, J.E.. 1969. Studies of physiologically active arthropod secretions. 2: some observations on the scent pencils of male Vitula edmandsae (Lepidoptera: Phycitidae). Can. Ent. 101: 280–285.Google Scholar
Weatherston, J., and Percy, J.E.. 1977. Pheromones of male Lepidoptera. pp. 295–307in Adiyodi, K.G., and Adiyodi, R.G. (Eds.), Advances in Invertebrate Reproduction, Vol. 1. Peralam-Kenoth, India.Google Scholar
Willis, M.A., and Birch, M.C.. 1982. Lek formation and female calling in a population of the arctiid moth, Estigmene acrea. Science 218: 168–170.Google Scholar
Zagatti, P. 1981. Comportement sexuel de la pyrale de la canne à sucre Eldana saccharina (Wlk.) lié à deux phéromones émises par le male. Behav. 78: 81–98.Google Scholar