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On the Mechanism of Copulation in Hepialid Moths, Endoclita Excrescens (Butler) and Hepialus Fusconebulosa (De Geer) (Lepidoptera: Hepialidae)

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On the Mechanism of Copulation in Hepialid Moths, Endoclita Excrescens (Butler) and Hepialus Fusconebulosa (De Geer) (Lepidoptera: Hepialidae) Bull. Kitakyushu Mus. Nat. Hist., 3: 27-39. December 31, 1981 On the mechanism of copulation in Hepialid moths, Endoclita excrescens (Butler) and Hepialus fusconebulosa (De Geer) (Lepidoptera: Hepialidae) Kyoichiro Ueda Kitakyushu Museum of Natural History, Nishihonmachi,Kitakyushu, 805 Japan. (Received on 20. Sept., 1981) Abstract The male and female genitalia of Endoclita excrescens (Butler) and Hepialus fusconebulosa (De Geer) are described and figured. The musculature of male genitalia of Endoclita excrescens (Butler) is described and figured. The mode of firm connection between male and female genitalia, which has not been recorded in Lepidoptera, is effected in the following sequences: (1) the mesosome was in serted into the dorsal groove of central process (E. excrescens) or grappled the dorsal sclerotized portion of central process (H. fusconebulosa), (2) the valvellae were applied to the lateral walls ofseminal gutter (E. excrescens) or the depression ofsubanal plate (H.fusconebulosa), which was opened by the inflation oftegumen, (3) thevalvae clasped the weakly sclerotized portions of female diaphragma (E. excrescens) or lateral portions of female ninth tergum and sternum (H. fusconebulosa). The characteristics of these copulation mechanism in Hepialidae and their phylogenetic significance are shortly discussed. Introduction The male and female genitalia ofLepidoptera are much variable in structure. They have been carefully studied in detail and used as important characters in taxonomy. On the contrary, the function of parts of the genitalia, especially in copula, has almost been remained unknown with the exception of De Jong (1978) and other few works. In the family Hepialidae, such studies are few and we hardly find Stekol'nikov's remark that "The nature of fixation of the sexes in copulation is quite different in H. humuli from what has been observed in the other Lepidoptera. The dorsal and ventral tegminal lobes open and are inserted in special depressions on the abdomen of the female, and the valves hold the abdomen of the female laterally. The weight during copulation is therefore principally borne by the tegminal lobes and the muscles responsible for move ment of the lobes are most strongly developed" (Stekol'nikov, 1967: 403-404.). From 1979 to 1980,1 could make several pairs of Endoclita excrescens (Butler) in copula by hand-pairing method (Fig. 1). The male and female in copula were 28 Kyoichiro Ueda fixed in boiling Carnoy fluid, then dissected and observed in detail. In addition to them, I had an opportunity to examine a pair of Hepialusfusconebulosa (De Geer) in copula, which was fixed in Bouin fluid, through the courtesy of Dr. Kauri Mikkola, University of Helsinki, Finland and I could make a comparison between the two species. The terms used in this paper for each structure are based on Ueda (1978, 1980). 1. Description of male and female external genitalia. Fig. 1. A pair in copula. Arrow indicates 1.1. Endoclita excrescens the fore and mid legs of male. Upper; (Butler)*" female, lower; male. Male external genitalia: Tegumcn very large with dorsal margin bearing a pair of swollen portions, which bears dense small serrations; twin process almost reduced; processus momenti broad and hook-shaped, strongly bent downwards at middle, and its posterior portion producing dorsally; valvclla long, producing ventrally, and tapering towards tip; the dorsal margin of valvclla with dense denticulations, and producing laterally; vinculum almost as deep as tegumcn, with no special process on the posterior margin; saccus large and rounded. Valva small, tapering towards apex and with rounded tip. Mesosomc fused with tabulatum near its base, strongly sclerotized ventrally and ending in a sharply pointed tip. Tabulatum strongly concaved at middle portion and with a large keel-like ridge running through its ventromedial portion (muscle attachment of m.7). Female external genitalia (Fig. 2): Eighth sternum well sclerotized, almost rectangular. Ninth tergum almost three limes as deep as ninth sternum, slender and semicircular, with minute hairs on dorsal margin. Ninth sternum producing a pair of slender and flat processes from lateral margins; ventromedial portion of the sternum forming central process which has a slender and deep groove on its dorsal surface. Diaphragma very broad, with a seminal gutter running middle of diaphragma. Weakly sclerotized subanal plate on lateral portion of dia- *1) The external genitalia of this species were figured on Ueda, 1978 and 1980 (male; Figs. 8 A-C, Ueda 1979, female; Figs. 11-12, Ueda 1980). Copulation mechanism in Hcpialid moths 29 Dorsal groove Seminal gutter Central process Lateral process Fig. 2. Female ninth sternum of E. excrescens (Butler). Right dorsolateral aspect. phragma, and almost elliptical. 1.2. Hepialus fusconebulosa (De Geer)*2> Male external genitalia: Tegumen broad; twin process small, with rounded tip; processus momenti strongly bent downwards at middle and dilating towards distal margin; valvella bifurcate, the dorsal process of valvella short and triangu lar, the ventral processof valvella slender, projecting ventrally and its tip reached near ventral margin of vinculum; vinculum broad, with a pair of slender processes on lateral posterior margins; saccus large, ventromedial portion of saccus with acute irregular apodemal processes on dorsal wall (muscle attachment of m.78). Valva rather long, three times as long as wide, almost subequal in width throughout the length, tapering slightly on distal margin and projecting above beyond dorsal margin of tegumen; the ventral margin of valva slightly produced posteriorly at middle and its tip truncate. Mesosome large and well sclerotized. Tabulatum almost rectangle, with small serrations on internal surface. Female external genitalia (Figs. 3-4): Anterior portion of eighth sternum weakly sclerotized. Posterior margin of ninth tergum weakly concaved at middle; *2) Male external genitalia of this species was figured on Ueda, 1978 (Figs. 7 A-C). n Fig. 3. Hepialus fusconebulosa (De Geer). Terminal Fig. 4. Hepialus fusconebulosa (De Geer). Female genitalia. Caudal segments of female abdomen. Lateral aspect (left). aspect. Copulation mechanism in Hepialid moths 31 dorsomedial portion of ninth tergum membranous. Ninth sternum three times as wide as high; ventromedial portion weakly swollen with slight depressions on both sides; dorsal margin of ninth sternum with a pair of short and blunt processes, which obliquely protrudes upwards. Subanal plate very slender and weakly sclerotized. 2. Musculature of male external genitalia of Endoclita excrescens (Butler) (Figs. 5-8) The musculature system is mainly based on Birket-Smith (1974). m.12 (tegumen-twin process): untraceable, the absence of this muscle seems to m.i9-1 m.i9-2 m.i9- Fig. 5. Endoclita excrescens (Butler). Male genitalia showing its musculature. Frontal aspect. 32 Kyoichiro Ueda Fig. 6. Endoclita excrescens (Butler). Male genitalia showing its musculature. Lateral aspect (left). be correlated to the reduction of twin process. m.3 (tegumen-processus momenti): absent. m.4 (vinculum-processus momenti): a long and strong muscle, arising dorsally in lateral portion of vinculum-just laterally of the tabulatum in normal position- and inserted on the anterior margin of processus momenti. m.5 (vinculum-valva): thin and slender muscle, taking it origin on the middle of lateral portion of vinculum and inserted on dorsal portion of the base of valva. m.6 (tabulatum-valva): very thin muscle, arising on latero-frontal margin of tabulatum and inserted dorsally on the inner wall of the base of valva. m.7 (vinculum-ductus ejaculatorius): broad and thin muscle, arising on ventro- Copulation mechanism in Hepialid moths 33 Valva Tabulatum Vinculum Fig. 7. Endoclita excrescens (Butler): Male genitalia; right valva and vinculum with their muscles. Lateral aspect. m.i9 - 2 m.i9-3 Fig. 8. Diagram showing ductus ejaculatorius and its muscles of E. excrescens (Butler). frontal margin of vinculum, bent at an acute angle on anterior margin of tabu latum, and inserted on ventral portion of ductus ejaculatorius. m.8 (vinculum-tabulatum): very large and strong muscle, arising on ventral portion of vinculum and inserted on tabulatum. m.i9 (tegumen-tegumen, tegumen-ductus ejaculatorius, processus momenti-ductus ejaculatorius): extensively large muscle, divided into three parts: m.i9-l very large transverse muscle, which interconnects the subdorsal walls of tegumen. 34 Kyoichiro Ueda m.i9-2: rather small and short muscle, arising on the middle portion of tegumen and inserted on the dorsal wall of ductus ejaculatorius. m.i9-3: very large muscle, arising on the inner wall of processus momenti and inserted on lateral wall of ductus ejaculatorius, and ventral portion of this muscle passing below ductus ejaculatorius and interconnects the two processus momenti. m.10 (processus momenti-valvella): large and triangular muscle, arising on the bending portion of processus momenti and inserted on the membrane near the middle of valvella. m.11 (dorsal wall of sugenital crypt-valvella): absent. 3. The mechanism of copulation 3.1. Endoclita excrescens (Butler) (Figs. 9-10) First of all, the male was stimulated by rubbing the ventrocaudal portion of abdomen longitudinally, or gently pushing dorsal portion of tegumen. The male moth, which was fully stimulated, usually began to move his genitalia in the following sequences:*3) 1) the whole tegumen is turned upwards on the processus momenti (con traction of
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