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Reproductive Behaviour of the Lesser Waxmoth, Achroia Grisella (Pyralidae: Galleriinae): Signalling, Pair Formation, Male Interactions, and Mate Guarding

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Reproductive Behaviour of the Lesser Waxmoth, Achroia Grisella (Pyralidae: Galleriinae): Signalling, Pair Formation, Male Interactions, and Mate Guarding REPRODUCTIVE BEHAVIOUR OF THE LESSER WAXMOTH, ACHROIA GRISELLA (PYRALIDAE: GALLERIINAE): SIGNALLING, PAIR FORMATION, MALE INTERACTIONS, AND MATE GUARDING by MICHAEL D. GREENFIELD') and JAMES A. COFFELT2) (Insect Attractants, Behavior, and Basic Biology Research Laboratory, ARS, USDA, Gainesville, Florida, U.S.A.) (With4 Figures) (Ace.10-XII-1982) Introduction The mating systems of most species of moths conform to a general pat- tern in which pair formation can be modelled as female signalling and male searching (SILBERGLIED, 1977). Females typically remain close to their emergence site after eclosion, emitting long-distance chemical signals for particular periods of the day or night, often exhibiting a characteristic "calling posture" during emission. Males search per- sistently for such females (HAGSTRUM & DAVIS, 1980) via an extremely acute sensitivity to conspecific pheromone (KAISSLING & PRIESNER, 1970; SOWER et al., 1971 ). However, several species are known that do not con- form to the stereotyped model (GREENFIELD, 1981). We studied the reproductive behaviour of one of these "non-conventional species," the 1) Present address; Department of Biology, University of California, Los Angeles, CA90024 U.S.A. 2) We wish to thank J. BENNER,R. W. MANKIN,W. T. MCCLELLAN,K. N. PRESTWICH,F. ROBINSON,J. SHARP,K. W. VICK,J. C. WEBB,and G. P. WHITMERfor able technical assistance without which this work would not have been possible. The in- itial phases of the work benefitted greatly from discussions with T. BURK,D. BIEMAN,and T. J. WALKER.P. R. ATKINSONgenerously allowed us to read several of his unpublished manuscripts. The study was supported by a postdoctoral fellowship to M. D. G., jointly administered by the U.S. Department of Agriculture and the University of Florida. Critical readings by D. BIEMAN,C. BOGGS,T. BURK,M. DALY,R. MANKIN,M. S. MAYER, J.R. MCLAUGHLIN, SIVINSKI,J. T. J. WALKER,translation of the summary by S. TAYLORand E. VILLALOBOS,and typing by E. TURNERgreatly aided the manuscript's final preparation. Mention of a commercial or proprietary product does not constitute an endorsement by the USDA or the State of Florida. 288 lesser waxmoth, Achroia grisella (Fabricius) (Pyralidae: Galleriinae), to ex- plore the extent of its deviation from the usual system. A. grisella is a cosmopolitan inhabitant of honeybee colonies, the larvae infesting detritus and brood combs (MILUM, 1935). KUNIKE (1930) briefly described its mating behaviour, reporting the strong odour released by the males, their stationary wing-fanning activity, and the attractiveness of males to females over long distances. Later, FINN (1967) showed that the odour was released from a pair of glands located ventrobasally on the forewings. DAHM et al. (1971) chemically identified the odour as a com- bination of undecanal and cis-11-octadecenal, demonstrated that these 2 compounds were necessary for female attraction, and suggested that sex- ual communication involved an additional acoustic signal produced by wing-fanning males. Therefore, pair formation in A. grisella superficially - appears to be a sex-role reversal of the typical female signaller male searcher system of moths. In this report, we describe observational and experimental analysis of various aspects of A. grisella reproductive behavior. Initially, the periodicity of male signalling in the laboratory is described, corroborated by field observations on emergence, signalling, and mating activity near a beehive. We then quantitatively analyze the pheromones emitted by signalling males for comparison with the quantities of pheromones releas- ed by females in other moth species, typically very low ( < 100 ng/h) values (GREENFIELD, 1981). The air movement produced by wing- fanning males and the energetic requirement of wing-fanning are also ex- amined to more completely characterize the nature of male signalling. Because of the apparent sex-role reversal in A. grisella, we discuss its pair- forming sequence in detail and report the results of experiments on female choice of signalling vs non-signalling males, the effects of female age on the response of females to males, and the occurrence of multiple mating in females. In insects in which males signal, e.g. various Diptera (BURK, 1981) and Orthoptera (OTTE, 1977), temporal signalling interactions (chorusing), aggression between signallers, non-signalling males, and aggregations of signallers are often present. We report observations demonstrating that these phenomena occur in male lesser wax moths in the Male Interac- tions section of this paper. Additionally, experiments are described in which we tested the orientation of males toward signalling vs non- signalling males, whether females ever mate with non-signalling males situated close to signalling males, and the orientation of females toward lone as paired (aggregated) males. Finally, in the Mate Guarding section, .
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