TThhee LLeepipdiopdteorpoltoegircaollSoocgieitycal Society ooff JJaapapnan utLma 1)'anslopktSoc.Jqpan 54(1):47-67,January 2003 CopulationposturesinLepidoptera(Insecta) AkiraMyATAi),H.S.YoNG2),Yatsukaho IKEDA') Hideo HAsEGAwAi) and i)Department oHnfectious Diseases ,Oita Medical Universit yH,asama, Oita Pref. 8,79-5593Japan 2)lnstitut SainsBiologi,Univcrsiti,Malaytt,50603 Kuala Lumpur, Malaysia Abstract In copulation, most Lepidoptera unite with their abdomens end-to-end, their heads pointing in opposite direction asn,d their forewing smaking slopes to both sides Lik earooi We call i t'`roof type" copulation. However, the slope ofthe roofvaries according to difierence sin length of the body and size of the wings. In a typical case "female's wings cover male's wings (FCM)" or conversely "male's wings cQver female' s(MCF)" .In these cases, the female always occupies the upper side with her head pointing upwurd during copulatien on incline dslope or vertical surfacc. We eall this "female dominant (FD)" .`・Male dominant CMD)" is possible but x,ery exceptional. Females always keep the position FD with very rare exception. In the case of wing arrangement, FCM is apparently dominant, but often MCF occurs under natural conditioii. We classify copulation in Lepidoptera ,especially nocturnal moths, into fiv epostures as fbllows: Subtype 1-Typical roof type (RT) ;Subtype 2-1-V-shaped postur e(VSP) ;Subtype 2-2- Face-to-fa cpeostur e(FFP) ;Subtype 3-1-Horizontal wing posture (HWP) ;and Subtype 3-2- Butterfiy posture (BP). All subtypes may be derive dfl]o ma primitiye RT; Subtype 1 is the true RT in the strict sense. In Subtype 2, the heads of both sexes point upward when resting on incline dsiope or verticat surface. VSP is V-shaped copulation, in which the male approaches the female laterally h,ence their bodies make a V shape. In FFP, the male late ron moves to his partner's ventral side and assumes face-to-fa pcoesture. In HWP, as the wing size becomes large ra,nd the abdomen size becomes slender, the slope of RT is reduced gradually f,inalt byecoming almost horizonta l.If during copulation, the pair' wsings are held horizontall wye, call it HWP. And in BP, the pair' s wings are held upright on their backs. Key Copulation, LepidQptera, butterfly. words posture, classification, nocturnal moth, Observation sofcopulation in nocturnal Lepidoptera are very rare compared to the day-flying species, because of diMcult iyn findin gthem at night, Fortunately we observed seme cases under natural conditions and also on the white screen illuminate dby ligh tat night fbr collecting moths. In ,the present paper, we describ ethe copulation postur ein each pair observed in detai la,nd in addition we examine and discus sthe many photegraphs ofmating pair sthat appeared in various llteratures. tt Method 1. When a pair of moths was found in copulation, we took picture sand observed them in detai l.Our main observation fiel dwas Jizobaru, especially in the garden of the Kokonoe Institu toef Natural History (KINH) ,situated at 3229-317 Jizobaru ,Machida, Kokonoe- machi, Kusu-gun, Oita-pref i(abou 8t30 meters above sea level ;33009'05"N, 131011'oo"E). 2. As our observations are not enough to discu stshe copulation posture ofall moth families, we reviewed many photographs from the literatu reA.mong them, we fbund some copula- NII-Electronic Library Service TThhee LLeepipdiopdteorpoltoegircaollSoocgieitycal Society ooff JJaapapnan 48 Akira MiyAiA, H. S. YoNG, Yatsukaho IKEDA and Hideo HAsEGAwA tion picture sof moths from three journa l(sJdpa nHeteroceiis tJsb'urnal ,Ytzgato ,and insectarium) and two illustrat beodoks ashizaw a19,56; Hidaka [Ed, ]19,97). However, we cou]d not find good picture sof copulation except butterfli fersom other journ asluchs as (]ekkan=Mushi, Tinea,Kontyn, T7ansactibnsofthe Lepidbpterologr'Scbaclietyoj'Jbpan, .lbumal of Researeh on the Lepidbptera, and .Jbumal of the Lepidbpteits tSsb'etety ,and several books on Lepidoptera written in English or Japanese. 3. In the presen tpaper, the fo11owin gterms are used for explanation ofcopulation. In most cases of copulation, male and female unite with their abdomens end-to-end and their heads pointin gin opposite direction s.Their forewing smake slopes on both sides, resembling a smal1 roof We cal1 it "roof typ e(RT in the presen tpaper)" as shown in Figs 1, IO-15 ,and 26-27. In Geometridae ,the slepe oftheroofis gentle a,lmost horizonta l.In these cases, we call it "`horizontal forewing postur e(HWiP) "(Fig 1s8, 5],54 ,55, 58-60, and 63) . The RT is apparently the basic postur ein copulation, but many other variations derived from it were observed. 4. In the case ofRT, when the pair rests on a venical or inclin esdurface, usually the female point sher head upward, and the male's head downward. We call it "female dominant (FD)" ,and in the case of male's head upward, "male dominant (MD)" .FD is apparently very common compared with MD. In some copulation cases, `lhe end of the male's fbrewing scovers the female' swings (MCF)" b,ut mostly "the female' swings cover the male's (FCM)" .These are used fbr descriptio onf the pair' swing arrangement, 5. In a posture derive dfrom RT, the male points his head upward, paralle lto the female, and copulates with her lateral ly.The pair fbrms a V-shape . ifthey retain the posjtion until the end ofcopulation as shown in Figs 33, 35 and 38, we call it "V-shape postur e(VSP)" .IC after coupling, the male moyes to the female' sventral side, they mate face-to-fa acse shown in Figs 30-32, 37 and 61, We call it "face-to-face posture (FFP)". 6. In most butterfl iaensd some moths, the wings ofboth sexes during copulatjon are closed in an upright positio nabove the body as shown in Figs16,17,S2 and 56. We cal1 it "butterfly posture (BP) ".HWP and BP probably reflect close affinity; we observed some- times beth postures in one species or one genus, but one posture only in many species, 7. 0ther copulation postures ,derive dfrom conspicuous lengt hof wing and abdomen or ditferenc ein body and wing sizes due to sexual dimorphism, will be detaile dunder descrip- tion of the observed species. Obseryation We listbelow from literature; examples ofcopulation our observations and where published illustrated but observations were with only photographs without any note, we append a short supplementary explanation, [I lFamily HEPIALIDAE = 9 e V Jij"*IP 1. Endoclyta excrescens U VZ V Jif RT, FD, MCF: Ueda (1985 p)hotographed a pair by hand-pairing. 2. Gazo,yctr amacdenta t IJ V >i >< 9 - V va V RT, FD: Pair photographed on a stem ofthe fatnil Gyramineae at 7 p.m,, August 26, 1983 ,in Nagano Pref (Nishi 1o9,86b) . This is RT lik eEL excrescens; in the photograph male and female rested in horizonta lpositien ,but FD when Nishio fir sftbund it .Wings of both sexes were closed, but abdomens were very long and did not cover each other, NII-Electronic Library Service TThhee LLeepipdiopdteorpoltoegircaollSoocgieitycal Society ooff JJaapapnan CopulationPosturesinLepidoptera 49 One hour 30 minutes late rb,oth individual tsurned their heads face to face. [I IF]amily INCURVARIJDAE =?')ij" V Jij"lilF 3. Nemophora staudingereZla VXf<=tb'tJifff RT, MCF: A pair on horizonta lfia tsurface was photographed by Matsuka (Hidaka, 1997) :Female with both leg sstretched and head pointed slightly upward; maie's head downward. pointed rather [IIIF]arnilyPSYCHIDAE l'7 ffliSF 4. Acanthopsyche nigTaplaga J}k tiv q .7 J(f Fig. 46, sketched from the photograph by Kobayashi (1973 )Th:e male's abdomen insert eddeeply into the female' spupa case to copulate. After emerging, the wingless female waited for a male's visit, with her head pointing downward, therefore her abdomen end was in the deepes tpart of the case. The male's abdomen has elasticity to reach the female' ssexual organ. Saigusa (197 4h)ad reviewed the copulation in this family .He describe dfbur steps of evolution in copulation of the family .In primitive species, the female has wings and can fl yout from her case for mating. [IV]Family YPONOMEUTIDAE XJifllSl・ 5. )[PonomeuM todyonetZtz `szi'n .$ JdiJIJX>if We fbund a pair in the early morning efJuly 14, 2oo1, on the sliding shutter ofKINH (Fi g1.; the righ ts{de is female). [V] Family SESIIDAE JJ <de )i7"Jij"X)I/ 6. Pennisetia juen ijixen -itzXY-do S/," We found a pair of this species on a horizonta lleaf of a roadside plant in Kokonoe near Mt Kurodake on the afternoon of September 1, 1980 (Fig 2.). The height ofthe leaf was about 60-70 cm above the ground. The upper individu ainl the pictur emay be the male. Two copulation picture sof IVbkona pexnix (t ?( 7 F >i LJi jSi7") taken by Nojiri appeared in I>zsectari u(m198 51,988) , These pictures depicte dthe same pair as they were taken by the same person and showed similar arrangement of the leave saround the pair, The pictur ethat appeared in 1985 is an example ofFD, and that in 1988 MD. The fbrmer picture is probably correct, as the latte ris apparently upside down. The pair might be affected by the presence of the photographer, with the female wanting te meve hence dragging the male, which didn' twant to move and resisted her tugging. In these pictures, the true copulation postur ewas cempletely disturbe dby the observer, We must take care to approach a pair of day-fiyi nmgoths and butterfli beesc,ause the presenc eof an observer could cause the male or femal eof the pair to attempt to escape, hence rendering their copulation postur eabnormal, Arita and Ikeda (2000 )publishe da monograph ofJapanese Sesiidae . They recorded the copulation tirne of 2-3 species, with some observations of their mating behavior such as calling by the female . The copulation of Skisi ardynchioidles (t hSiVJk do j/79) and 7)ichocero taconstticta (AtfF t ?e XJij Sf,") on the surface of an inclined slope is apparently FD in a straight 1{ne . In the photographs of a single individpa lt,he head is also always pointed upward, therefor "eFD" is normal in thi sfamily. [VI ]Family CHOR.EUTIDAEixv i E F i )lf"liSl/ 7. Choreutisfabninea7x>ifStlf7xviEFi Accerding to the photograph of a pair taken by Ohbayashi (199 7on) March 7, 1995, in Chichijima, Ogasawara Jslands ,the male and female rested with their wings upward, and NII-Electronic Library Service TThhee LLeepipdiopdteorpoltoegircaollSoocgieitycal Society ooff JJaapapnan so Akira MIyATA, H. S. YoNG, Yatsukali oIKEDA and Hideo HAsEGAwA the individual at the lower positio nmight be the female, Hidaka (199 7a)lso used the same photograph , Fig, 50 is our rough sketch, [VII ]Family OECOPHORIDAE -?7V7xS 7")erliSF 8. Diun2ea eurpreij2]raS-m"*'?7Wxi7"Jij' RT, FD: A pair on a tree stem photographed by Miyasaka (]99 2i)n Sanadamachi, Nagano-pre£ on April 30, 1983 , Fig, 47 is sketched from the picture, , [VIII] Family ZYGAENIDAE vMili'lilL 9. Erasmia putehella V' V v; Si S' RT, FCM: Ohkubo (1991) horizontalleaf photographed a pair on a 10. Elyewma westwoodZi Il7><7"))179pt rf According to a photograph (Mik i19,98) ,the female rested with her head up and both her fbrewing sfbrming a very gentl eslope; the male with his wings erect on his back like butterf lgyot into a slit between the female' fsorewings a,nd his body is concealed complete- ly from head to abdomen. Bocause this species has large forewin gwsith rather short abdomen, if the female holds up her wings, the copulation is classified as BP. However, she did net do so, and the male had to compromise to accomodate his partner .According to DrKoshio's comment attached fbr this picture ,this posture is standard to the species; the male fiie isn the early morning, and copulation takes plaee until noon time. 11. Zygaen aniphona /<= e )-vM7 RT, FD: Nishio (198 9p)hotegraphed a pair in the middle of July in Ueda City, Nagano pref The pair was on a sta!k. Nakajima (198 7s)howed two photographs of copulation, one on ItZibe triotsundota and the other IIZZbe ittesnuts. Both are apparently RT and MCF. We believ ethat both photographs were turned sideways, because of layout .The copulation might be FD. [IX ]Family HYBLAEIDAE -t z-} zV 'E F }Jij"7ilF 12. H),bla feoartis s;i)tKm a>fiz -l zV i F} Nishio photographed a pair on a horizonta lsurface at 11 : 30 p,m, ,Apri1 5, 1995 ,in Ueda City ,Nagano Pref OSJish 2i0e0,0a) . According to the short explanation attached, the male opening the hair tuft on his hind leg ,approached behind a female ,and attempted to copulate flappin ghis wings. The copulation posture is not true TR; their wings were somewhat oblique to the dorsa ldirectio snimilar to Figs 51 or 54. The pestur emust be classified as HWI', The female' shead is pointed upward, and the male's downward. [X]FamilyP\RALIDAE f1 tiN 13, Seirpophaga invertulas RT, FD, MCF: Fig. 57 was photographed by H. S, Yong in the daytime in Sekin- Selangor,Malaysia, July,1971, chan, on 14 . Ii:},men iraee,urvaiZy Ynttf/1d ff RT, MD, FCM: We fbund a pair of this moth on the white sereen for collecting moths by ligh tat KINH in the morning of August 11, 2oo1 (Fi g3.). 15 . Datanoidle fsascia t1urst a Yt t},'Y?pt i ti RT, FD, FCM: We observed the copulation of this species on two occasions, First, we fbund them on the white screen at KINH at 7 : 20 a.m., July 21, 2oo1 (Fi g4s-5) . The upper large rone was female ,with her 1arg eabdomen sticking out slantingly; the male with all his legs contracted was floatin gin the air at the end of her abdomen. RT, FD, MCF: in the second case, we found a pair on a screen door at KINH at 7:oo a,m,, July 26, 2oo1 (Fig 6s-7) , The pair stayed still in the same pose for 7-8 hours after we fbund them; after that we caught them because we had to leave .In thi scase, his NNIII-IE-leEcltreoncitcronic LMbirabrryary Service TThhee LLeepipdiopdteorpoltoegircaollSoocgieitycal Society ooff JJaapapnan CopulationPosturesinLepidoptera 51 foreleg asppeared to be touching the doer. 16. AmppaTu induator1)i`?fviVi'gptiff RT, FD, MCF: We fbund a pair on the white screen at KINH at 6 : oo a.m., July 6, 2001 (Fig.13), 17, Orthaga onerata )}t s7')t7 F ?( -t )lj" RT, FD, MCF: We fbund a pair en a wall at KINH at 6 : oo a.m., July 13 ,2002, 18, E?idotntha olivaceads VXfxq= F ff V pt d ti Sawazaki (198 4sh)owed a photograph of a female copulating with a dead male. It was considered to be RT and FD. [XI]FamilyPTEROPHORIDAE b V7")ij'1;)P 19, Oc]tyrode aconcmsa tt))2vl J{i b Vi" According to a photograph of a pair of this species taken by Shinkai (Hidak a1,997), they formed an H-shape because the pair' swings stretched jus tbesid etheir bodies. [XII ]Family THYATIRIDAE F ff V i"Jiflill・ 20. Demopsestt spunctiger aJtK S! ri ?)i F ff V !" RT, FD, FCM: We found a pair on the white screen at KINH at about 10 : oo p.m., April3,2oo2(Fig3.9). [XM] Family GEOMETRIDAE )/ v 7 tilil} 21, Archieants parthenia sip79Sf- J?7 Nakajima and Jinbo (1989 f)bund a male of thi sspecies fiying in the sky against the wind in Mt Irikasayama N,agano Prefi at 4: 30 p.m. ,May 5, 1988. Then it stopped on a small tree of Quencu msongoiica var. grastserwa taand copulated with a female waiting there, Aocording to the photograph ,the pair was on a horizonta lside twig, and the female' sside was somewhat higher than the male's side, Hewever, the photograph is considered not to be in a natural position, but rather the twig was jncline dfbr layout, 22, A lsophilo iaderboasma - )S JE >f 7 u. Si Jv 9 RT, FD: Tanaka et aL (199 4p)ublished a photograph of a pair in which the female stayed in the lower position with her head pointed downward. Later one of the authers (Mizun o1,994) pointed out that the photograph was printe dupside down. 23. AJaxa seriaria rk Y S/ te 7 FD: The femal eopened her wings horizontail byu,t the male closed his wings dorsally like butterfly his left (Ishizaw1a95,6). a and crept under partner' s wings 24. Entephri acaesiata -V'V't ) -7' } Si rt ra RT., FD, MCF: A pair was fbund insid ea window gla sosf a cottage in Ina City, Nagano Prefi ,at 4 : 40 a.m., August 25, 1987 (Nish i1o98,8). 25. Heterothera taigana sounkeana Y9 >f7pJt tft ) 'Yte 7 BP, FCM: A pair was fbund in Ina City ,Nagano Pref, at 7 : 30 p.m. ,August 14, 1992 (Nishi o19,92) .The pair appeared to be hanging from the underside of a twig, with the female' swings hel dupright like butterfly. A similar photograph of Opa,zlphte rbarunnea (-t l -7 r-:}m ) -S/ ftiP) published by Nakajima (1992 s)ho,ws it to be FD. In this winter appearing species, the female's wings are very small but the male's wings are norrnal and held upright like buttediy .At leas tsome moths of the family Geometridae normally rest with wings upright over their backlike butterfly. a 26. Abraxas tptv Sf"i)zSf"SivO niphombia FD, BP: We fbund a pair resting on the white screen at KINH at 5 : oo am., August 25, 2001, as shown in Fig. 17. In this case, the pair held their wings upright like butterflies. NII-Electronic Library Service TThhee LLeepipdiopdteorpoltoegircaollSoocgieitycal Society ooff JJaapapnan 52 Akira MiyATA, H. S, YoNG, Yatsukaho IKEDA and Hideo HAsEGAwA Figs 1-15. Copulation postures ofLepidoptera. 1. I))ononieuta tokyoneaus 2. Pennisetia jZyeni]tu en3i. .R:ymenia recurvalts: 4-5. Datanoiclesfosciatus(samepai r)6,-7, D. .fasciat u(ssam epair). 8-11, Athetis athisignata (8-10 :moving of pair ;11: different angle of10). 12 .A.athtsigna ta(oth eparir) . I3. Arippara indicato n 14-15 .Rivula (same serieeads pair). NII-Electronic Library Service TThhee LLeepipdiopdteorpoltoegircaollSoocgieitycal Society oofJfap anJapan CopulationPosturesinLepidoptera 53 I'' . t -, .- Figs l6-29. Copulation postures of Lepidoptera. 16. Everes argiades I7. Abraxas nP phont h ia , 18 .A, latdesciat a.19-20 .Eilema doplanapavescen s(same pairin diiferent angle). 2 1 .E doplan apavescens 22-25 .Lithosi aquadea (al ldiifere nptairs ).26- 27. T71anatarct iianaequads (2 ditferen tpairs) . 28-29. Amata fortun e(isam epair. but moving). NII-Electronic Library Service TThhee LLeepipdiopdteorpoltoegircaollSoocgieitycal Society ooff JJaapapnan 54 Akira MiyATA, H. S. YoNq Yatsukaho IKEDA and Hideo HAsEGAwA 27. Abraxas latdescia tta F X si "7 N' i7 :z 5' i"i ft r7 FD, HWP: We fbund a pair resting on a leaf of a fer nnear KJ'NH (Fi g1.8). 28, Cystid sinnatonice F )-)t?J[zY"Siftr7 In Ishizawa' pshotograph (1956 )a ,pair of this specieg was FD and BP. However, Kuribayashi's photograph (198 5o)fthe same species shovLred a differe nptose, HWP. This means this species has two copulation postures B,P and HWP. The genus Abraxas also might have two postures similar te the genus C)istidia. 29. eoncinnata FltL!=:S'" )tir )' ,Rrotalois FD, MCF: A pair on the stem of Pntnus ansu at 9 : 40 p,m. ,March 29, 1979 ,in Ueda City ,Nagano Pref i(Miyasaka&Nish i19o7,9) , With strong stimulation, both individuals began to move, then the male flutteri nhgis wings and the fernal ewalking. TD sum up, the pair could not fly and the female dragged her partner strongly for one centimeter. Apparently this pair could not cooperate to move to one directio angainst the stimulation. 30. PhigaEa sinuosa,t a)ie7V FtixtiYv7 RT, FD: In the geometrid moths that appear during winter season, the female 'wisngs do not develop ,therefore after evening the female stays on a stem and her abdomen end turns upward to release her pheromone (Nakajim 1a9,91) . Nakajima (199 8p)ub]ishe dan excellent review with many photographs of this group. 31. Btsto nrobustus } ti '{i>'le1ji5i'Yv7 RT, FD: As shown in Fig, 43, sketched from the photograph by Miyasaka (1983a). The copulation posture in most of the larg egeometrid moths is similar to B. robustus, as shown in the dead pair of Bisto n.fragt7 Zrseported by Inoue (1960). [XIV ]Family CALLIDULIDAE 1 )ig V e>f>ij'7ilF 32. Pteroclect a.fekler iK Ji jV 'E )-7V' BP: Nishio (199 9s)tudied the mating behavior ofthis day-flyi nsgpecies, and showed a photograph ofapair on the ground, The postur eis the same as BP, with the female' s wings hiding her mate's. [XV] Family LASIOCAMPIDAE Ji jLiiNJij"$II・ 33, Poeedocampa popufi 9XJ7r S do wx RT, FD, MCF: Four pairs areund a streetlight in Wadamura, Nagano Pre£, en November 6, 1983 (Nish i1o98,6a), [XVI ]Family BRAHMAEIDAE d Jt:S >if$iP 34. Brahmaea veai7ibhii 1 r£iS>V RT, FD, FCM: a pair photographed by Ishizawa (195 6a)nd another by Hashimoto (200 0a)ppeared on the cover of lnseetan'u m(Apri l20,00) (Fi g4,9, sketched from the picture), [XVll]FamilySATURNIIDAE 'e-?"? LJif1$F 35, Attacus atlas 9J}Ji)"=-V-Y VSP: phetographed by Jinbo (1995 ).The female' shead is pointed upward, and the male was approaching her right side to copulate. The male's wings were held upright over hi sback like a butterfl bye,cause ofobjects such as leave sand twigs betwee nthe two moths. FFP: photographed by Minato (Hidak a1,997) , In this case, the female hang from her cocoon and the front one was female , Fig. 61 was photographed by H. S. Yong in the daytime at University of Malaya, Malaysia ,on August, 1983 on Ilsidiu mguay'ava, According to the descripti onby Azuma and Kiajyo (1986 )a, male of A, atlas responding to the pheromone visited a femal eand stayed at her side, then he bent his abdomen end to her . The copulation postur ewas at fir sVtSP but soon the male turned NNIII-IE-leEcltreoncitcronic LMbirabrryary Service TThhee LLeepipdiopdteorpoltoegircaollSoocgieitycal Society ooff JJaapapnan Copulation Postures in Lepidoptera 55 180 degrees ,then became one straight line .However, thi sobservation was quite excep- tional ;we thought tha tthis observation was carried out in a rearing cage and other males prebably interfbr ewidth the pair . We asked Prof Azuma about this, and he answered that FFP was the usual copulation postur ein this species, 36. Samia qynthia pt:yer iSi>'9aV')- FFP: We found a pair of this species hanging from the twig ofa street tree, Ilex rotuncia, at 7 : 45 a.m., August 16, 2000, in Fojimigaoka ,Oita City (Fig 3s0-32) . The pair was about l60 cm above the ground, The copulation continued until the end of our observatjon at 10:OO a.m.; we carne back at 5:OO p.m. to continue observation, but they had disappeared. 37, Ctiligu ljaaponi c7aXV->f VSP: Three pairs were obs'erIvned on the white screen at KINH from midnight, October 3, 2oo1 to the next morning, the firs ctase, we found a pair at 10:45 a.m., October 3 (Fi g3.3), but the pair moved as shown in Fig. 34, at 2:44 a.m., October 4. The second pair was fbund at 3 : 10 a.m. October 4 (Fi g3.5), The third pair rested on the cover ofa black ligh tat 5:49 a.m., October 4. The postur ewas somewhat similar to FFP rather than VSP. We observed until 8 : OO a,m,, October4,but none moved. We observed fbur other pairs on October 8, 13, and 14, 2oo1; al 1ofthem had VSP and remained at the place we fbund them until the end of observation, Excepting Fig. 34, all other pairs assumed VSP even on the fla tground. in the present cases, the male couldn't move to the opposite side because of the very broad space on the white screen or the ground. Therefbr ewe believ ethat jf the femal ewere waiting at a twig or on her own FFP mightbe cocoon, observed. 38. Saturniajonasiitflevv=L FFP: photographed by Ishizawa (]95 6a)nd Obata (198 9a)s shown in Fig.48 (sketc fhreodm picture). 39. Rhodinia jugax I> J< 5' E' >if According to a photograph taken by Ishizawa (1956 a) ,female jus etmerged from her cocoon was copulating with a male that was also hanging her cocoon. The female' shead pointed upward, but the male's head pointed downward. This might be derive dfrom FFP. According to a persona !communication by Mr Mitsuo Nakajima, he observed FFP in the Actias artemts )trd l -2rJ71- in Oita Prefecture. [XVIII ]Family SPHINGIDAE XJCpt Jti'ffllF 40. ]Elyloicus caligineus 7vXfpt RT, FD, MCF: Ishizawa's photograph (195 6s)hows a pair resting on a stem, 41. Rsilogrammamenephron RT, FD, MCF: photographed by H, S. Yong in the daytime in Kepong, Selangor, MalaysiainApril,1990 inFig.62. as shown 42, Marumba sperchiusPt,"Xlrl FD: Miyata (20oo o)bserved apair as fbllow sA:t6:30 a.m., July 19, 1999 ,anewly emerged female was fbund on the hedge of Aheha grandlrJf iilno vFaLlj'imigao kO4ita City. Early next morning at 5: 30 a.m., she was in copulation with a male near her initial location (Fig 4.1). When the observer approached the pair to photograph them, the male with it sbody thrust fbrward watched him apprehensively (Fi g4.2) ,but the female djdn't react to the intrusio n.Until the evening (7 : oo p.m.) ,the pair was stil lin copu]ation, but about 8:30 p.m, the male flew off leavin gthe femal ealone in the thicket Miyata NII-Electronic Library Service TThhee LLeepipdiopdteorpoltoegircaollSoocgieitycal Society ooff JJaapapnan 56 Akira MTyATA, H. S, YoNG. Yatsukaho IKEiD Aand Hideo HAsEGAwA Figs 30-42 . Copulation posture sofLepidoptera. 30-32 ,Samia qynthia pt:Jie r(isamepai irn ditfere nantgle). 33-34 ,Caiigul aJ'aponic (asame pair ,but late rchanged position) . 35. C.iaponic a(otherpa ir36). .Emproctt spseudoconspeny a.37. Lymantria disparjapon- iea. 38. L, dispa r.iaponic a.39. Demosestis punctiger a.40. Marumba gaschkeivit- shii eclp)hron, 41-42. Ad Zsperchiu s(same pair, and male moved to watch observer). discusse tdhe presenc eof a second male, which was presen tat a site about 50 cm from the pair during the copulation period, In the famil ySphingidae ,the observed pair sare alway sin tandem as shown in Fig. 45, but the example of M sperchiu scopulating in the thlcket suggests strongly that this is NII-Electronic Library Service