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ecLma bens. Iopid. Soc. IaPan 46 (4): 175-184, December 1995
The life histories and bielogy of Epicopeiidae of Taiwan
Shen-Horn YEN", Jia-Horn Mu, Jia-Lurng JEAN
Department of Entomology, National Chung-Hsing University, 250, Kuokuang Road., Taichung 402, Taiwan, R.O.C.
Abstract The Iife histories and biology ef the twu species of Epicopeiidae of Taiwan are clescribed and illustratecl fer the first time. Additionally, a comment on phylogenetic position of this family is also provided.
Key words Diurnal moths, immature stages, hostplants, phylogeny.
Introduction
The family Epicopeiidae embraces several medium- to large-sized diurnal moths across the Oriental and the Palaearctic regions. They are usually regarded as mimic to a certain species of Papilionidae, Danainae or bearing aposematic coloration. In Taiwan, two species of Epicopeiidae, EIPicopeia mencia Moore, 1874, and E. hainesii matsumuvai Okano, 1973, have hitherto been recognized. wnile the early stages of the same species from Japan and China have been described or illustrated, the life history of this family from Taiwan has remained unknown to date. In early May 1992, one of us, Yen, was able to collect larvae and eggs of E, mencia from Ulmus Parvijblia Jacq. (Ulmaceae) at Huei-Sen Forests, Nantou Co. and Shanping, Kaohsiung Co,, Taiwan, respectively.
They were subsequently brought to the laboratory of Department of.Entomology. Rearing was conducted at room temperatures, Eggs were put into a small transparent plastic bag ; hatched larvae were reared in a 15 × 8 × 20 cm plastic case with an abundant supply of leaves of the hostplant; pupae were left in the case to emerge; hibernate pupae were mantained at 100C. In the early June 1993, the larvae and eggs of the other species, E hainesii matsumurai, were collected by us at Kukuan, Taichung Co., Hawan- shan, Wushe, and Wanda, Nantou Co. and Chinshuei, Hualien Co. They were also reared and observed by the same way. The purpose of this paper is to describe the elements of the life histories and present some notes on the biology of these fascinating members in the moth fauna of Taiwan.
Morphology and bielogy of immature stages and adult
Elpicqpeia mencia Moore, 1874 (Figs 1--7, 18-28)
Egg. Ovum diameter 1.1-1.2 mm, yerlow, spherical with a slightly flattened base and 16- 17 vertical ribs, aggregative on the under surface of termina] host leaves.
Larva. Coloration : Hatching larva length 2.2-2.4 mrn, ground color light yellow with dorsal and subdorsal stripes white ; head capsule yellow but soon turning to black after I hour ; body covered with white waxy secretion after few hours, Instars 2-6 similar in ground color and pattern of waxy secretion ; mature larva (instar 6) length 30.0-50.0 mm, ground color yellowish white with dark brown patches plus several waxing pores on these patches, skin with 3-4 wrinkles transverse ; waxy cover white, granular, uniform on each segment. Head : Capsule glabrous dark brown, chaetotaxy only with AF2 and
* Present address : Research Institute of Life Science, National Sun Yat-Sen University, Kaohsiung, Taiwan, R.O.C.
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Figs1-7. Last instar of Eipicopeia mencia Moore: 1. Chaetotaxy. 2. Head chaetotaxy (frontal view). 3. Head chaetotaxy (lateral view). 4. Hypopharyngeal complex (ventral view). 5. Mandible (left rear). 6. Mandible (right front). 7. Crochets.
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Fl present ; labrum with 4 pairs of major setae dorsally ; 6 stemmata ; rnandible almost rectangular, with 8 teeth ; hypopharyngeal complex as illustrated, with blunt spinneret. Thorax: Surface texture with some minute secondary setae; thoracic spiracles Iarge; Tl with doral plate dark brown ; chaetotaxy with SDI and SD2 apparently as single seta, with an L-group seta near spiracle ; T2 with Dl as a single seta and D2 plus several associated short setae over surface; T3 with L-group emitted on weakly sclerotized pinaculum, Ll and L2 anterior to L3. Abdomen: Al-2 similar to T2 except spiyacle present and SV group as approximate individual setae ; AIO with strongly sclerotized amber plate having several pairs of setae ; prolegs normal, all with many setae ; cro- "tetraordinal". chets uniserial and
Pupa. Length 20.0-28.0mm, width 8.0-10.emm; dark brown, stout, slightly setal; prothoracic legs equal to proboscis in length ; wing tips nearly reaching posterior margin of A4 ; dorsum with one or two setae on each abdominal segment ; spiracles visible, cremaster reduced.
Biology of immature stages. Larvae of all instars are strongly aggregative. The hatching larva does not consume egg shell, and the first instar larva behaves ag- gregatively with curving or arising the body when disturbed. Mature larva moves downwards to the ground to build a soft cocoon covered with white wax. Prepupal stage lasts a week and pupal stage takes approximately 25-45 (unhibernating) to 200 (hibernating) days. The complete early stage needs 60-90 days (unhibernating) to 240- 270 days (hibernating).
Adult flight period. Adult usually appears from April to October with two or three generatlons.
Adult. Adult coloration is dimorphic and aposematic with red tegula and transverse patterns on abdomen, one forrn is similar to a Troidini species byasa alcinezas, the other looks like I]kechliqPla an'stolochiae. The ratio between two forms is nearly equal. This
diurnal insect usually flies high near the hostplant and visits flowers of Rutaceae and Euphorbiaceae ; when captured, false death is always observed.
Hostplants. Ulnzus Parvtfolla Jacq. (Ulmaceae) is the sole confirrned hostplant of E mencia in China (Chu & Wang, 1981 ; Liu, 1983), Japan (Tsushima I.) (Miyakawa, 1993), and Taiwan (Chang et ag.. 1992). We supplied Ulmtts tcyematsui Hayata and Zbthova servata (Thunb.) Makino while the larvae did net accept the leaves. We suppose that physical condition of leaves is a limiting factor confining E. mencla on UL Parvijbha.
Habitat and distribution. The habitat is usualiy primary evergreen oak forest consist- ing dominantly of Querczts and Machilus. This insect is sparsely distributed to the low (500m) to medium altitudes (2,OOOm) of the whole island of Taiwan and strongly associated with its hostplant.
llpicQpeia hainesii matsumurai Okano, 1973 (Figs 8-17, 29-36)
Egg. Ovum diameter 1.0-1.1 mm, slightly smaller than E. mencia, light yellow, spherical with a slightly flattened base and 18 vertical ribs, aggregative on the under surface of terminal host leaves,
Larva. Coloratien: Hatching larva length 2.0-2.2rnm, ground color creamy yellow without dosal or subdorsal stripes white ; head capsule light gray but soon turning to black ; body covered with thin layer of white waxy secretion. Instars 2-6 similar in ground celor but dissimilar in pattern of waxy secretion, 2nd instar with thin layer of wax, 3rd instar having wax tufts more developed on Tl-3 and A8-10, wax secretion on
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Figs8-14. Last instar ofEipicopeia hainesii matsum"7uiOkano: 8. Chaetotaxy. 9. Head chaetotaxy (frontal view). 10. Head chaetotaxy (lateral view). 11, Hypopharyn- geal complex (ventral view). 12. Mandible (left rear). 13. Mandible (right front). 14. Crochets.
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Figs 15-17. Pupa of EipicQPeiahainesii matsumu7ui Okano : 15. Ventralview. 16. Dorsal view. 17. Lateral view.
Figs IS-20. Pupa oi EipicqPeia mencia Moore : 18. Ventral view. 19. Dorsal view. 20. Lateral view.
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Figs21-28. Life history of Eipicopeia mencia Moore. 21. Hatching larvain egg. 22. Ist instar. 23. Istand2ndinstars. 24. Matureinstar. 25. Pupaewithwaxycocoons. 26. Adult male. 27. Adult female, 28. Adult female (white-banded form).
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Figs29-36. Life hlstory of EipicQPeia fitainesii matsblmuvaii Okane. 29. Egg shells. 30.34. 2nd instar. 31. 3rd instar, 32.5th instar. 33. 6th instar (mature larva>. Pupa with cocoon removed. 35. Aclult male. 36. Celor variation of adult.
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Tl-Al, A4 and A7-10 filamentous in the late period of 3rd instar, 4-6th instars with wax tufts dorsally and laterally, a dotted black stripe present dorsally ; mature larva (instar 6) length 30.0-42.0 mm, smaller than E, mencia, waxing pores not visible as that of EL mencin, skin with 3-4 wrinkles transverse. Head : Capsule glabrous, black, chaetotaxy with A2 nearer to Al than to A3; labrum with 4 pairs of major setae dorsally; 6 stemmata, I and II smaller than the others, V situated ventrally, III, IV and VI on a black area ; mandible almost rectangular, with margin entire ; hypopharyngeal complex as illustrated, with blunt spinneret. Thorax : Surface texture with few minute secondary setae;thoracic spiracles smaller and less sclerotized than those of E. mencia;Tl without identical dorsal plate; chaetotaxy with all primary setae emitted on weak pinaculum; Dl,D2,XDI,XD2,SDI and SD2 apparently single, with an L-group seta below spiracle ; T2 with Dl as a single seta ; T3 with Ll far from L2. Abdomen : Al- 2 similar in chaetotaxy ; A3-7 with Ll separate from L2 and L3 plus 2-3 short setae ; AIO with slightly s ¢ lerotized amber plate having several pairs of setae ; prolegs normal, all with many setae ; crochets uniserial and triordinal.
Pupa. Length 15.0-18.0 mm, width 6.0-6.2 mm ; dark brown, stout, slightly setal ; simi- lar to E. mencia but with body much more slender, having prothoracic setae absent, spiracles reduced and cremaster developed,
Biology of immature stages, The first instar larvae are slightly aggregative, while the other instars are scattered on the hostplants. The hatching larva does not consume egg shell, and the first instar larva shows no defensive behavior when disturbed. Mature larva moves downwards to the ground to build a soft cocoon covered with white wax. Prepupal stage lasts a week and pupal stage takes approximately 25-45 (unhibernating) to 200 (hibernating) days, The complete early stage needs 60 days (unhibernating) to 210 days (hibernating).
Adult flight period. Adult usually appears from April to October with two generations,
Adult. Adult coloration is polymorphic with continuous variation of white/black ratio on wing pattern. According to our observation the complete black form appears with low frequency. This diurnal insect usually fiies high near the hostplant and visits flowers of Euphorbiaceae, when captured, false death is always observed, an undeterminated chemical secretion is released from the thorax.
Hostplants. The early stages of the Japanese E. h. hainesii were first reported by Nagano (1907) but he misidentified the hostplant. Sugi (1964) was the first to report Cornus controve7sa Hemsl. (=Sboidu controveTsa) and C, macmp1rylla Wall, (=Swicin macmpitylla) as the hostplants of E. h. hainesii, stating that the hostplant of hainesii had been erroneousry cited as Linde7u glauca Sieb. (= IV2olitsea sen'ca (Bl.) Koidz.), a member of Lauraceae. Sugi (1987) furthermore stated that the error might be caused in confu- sion between the local vernacular names of plants, The incorrect hostplant was also cited by Chu & Wang (1983) for the Chinese subspecies. Issiki (1969) recorded this insect additionally from C. hoztsa (= Benthamidia 1'mponica var. JJmponica). In Taiwan, we have confirmed Cornzas controversa, C. macropdylla and C. kousa as the hostplants of this species in natural condition.
Habitat and distribution, The habitat is usually primary evergreen oak ferest. In Taiwan, this insect is restricted to several localities of low (500 m) to medium altitudes (2,OOO m) of the north, central and east part of Taiwan and strongly associated with its hostplant
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Phylogenetic position of Epicopeiidae
The position of Epicopeiidae is not established, and it is usually included in Uranioidea only tentatively by Scoble (1992) and Inoue (1992). This family has recently been included in Drepanoidea by Minet (1991) on the basis of the presence of a complete prespiracular sclerite connecting the anterior corner of the first abdominal sternum with the lateral bar of the first abdominal tergum in the adult;the concealed or slightly exposed foreleg femur in the pupa ; the presence of setae on a flat and laterally delimited area of the larval mandible; and the occurence, in the larva, of at least one secondary seta associated with seta L3 on abdominal segments 1-8. However, accorcling to our preliminary observation of Sthistomiva fanemlds Butler and Rsychostmphia melanargia Butler, which have recently been transferred from Epiplemidae into Epicopeiidae by Minet (I986), immature morphology seems variable within this family. We consider that further biological investigation of other species will be needed to support Minet's concept of this family. Therefore, at the present we tend to adopt the treatment by Minet (1986) and suppose there are at least three major lineages, lipicopeia, IVbssa+krabraxas and Schistomiva+Psychostrophia and the phylogenetic pattern of this famiJy is deeply forked.
Acknowledgments
We are very grateful to Win-Yi Lin (NCHU), Yeong-Chium Sen (Taiwan Forestry Research Institute), Yasunori Kishida (Tokyo) and Yuan-Po Yang (NSYSU) for their helpful information and offering valuable materials. Thanks are due to Yi-Bin Fan (Taiwan Forestry Research Institute), Jean-Rong Her (Taiwan Endemic Species Research Institute), Juo-Yen Lai (NCHU) and Yeng-Jye Chen (NTU) for their various aids.
References
Chang, B.S., 1989. Epicopeiidae [Illblstmted Motlms of 71zizvan] 1: 174 175 (in Chinese). Chang, Y.C., Fan, Y.B. and Y.C. Sen, 1992. A name list of moths and their ioodplants at Shanping (Kaohsiung, Taiwan). Bull, 7keizvan thz Res. ins. New Series. 7 {1) : 39-71 (in Chinese with English summary). . Chu, H. F. ancl L. Y.Wang, 1981. Epicopeiidae. In Wang, P.Y., et al. (ed.>. ft:onag,mphia Heterocer- oium Sinicorum 1 : 107-108, pl. 27. Science Press, Beijing. {In Chinese). Draudt, M., 1931. Family Epicopeiidae. In Seitz, A. (ed.), MizcivlqPidoPtem of the WbTld 2 (Supp].) : 51. Inoue, H., 1978. Genus Elpicopeia Westwood from Japan, Korea and Taiwan (Lepidoptera: Epicopeiidae). Y},6 Ga 29 : 69-75 (in Japanese with English summary). ,l982. Epicopeiidae. in Inoue, H. et al. (ed.), Motlzs of 1of)an 1: 574-575, 2: pl. 109. Kodan- sha, Tokyo. (In Japanese). Issiki, S. (ed.), 1969. Etzrly Stages of 1mpanese Moths in color 2. vi, 237pp., 68pls. Hoikusha, Osaka. (In Japanese). Janet. A., 1933. Family Epicopeiidae. in Seitz, A. {ed.), Mdecrolopidoptem of the I･7brgd 10: 57-59. Liu, S. R., 1983. Uraniidae. in Tzai, B. Il. et al.
l991. , Tentative reconstruction of the ditrysianphy]ogeny (Lepidoptera:Glossata). Entomotogica scand. 22 : 69-95. Miyakawa, Y. 1993. Larvae of an IlipicQPeia taken in Tsushima, probably of mencia Moore (Epicopeiidae). ]1ipan Hbterocerists' J (178) : 36 (in Japanese with English summary).
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− − − 184 Shen Horn YEN ， JiaHorn Mu and Jia Lurng JEAN
Okano ， M ，，工958． New or little known moths from Formosa （1）．．4 朋 、 ROp ． Gaflugei Fac．如 α ’θ ひ卿 ． − （13＞2 ： 51 52 （in Japanese with English summary ）， ，1964．On the Formosan species of the family Epicopeiidae（Lepidoptera＞． Tohokblκ o 咒 o 舷 − Kenkyu l ： 25 26 （in Japanese with English summary ）． ，1973．Arevision of the Formosan species of the family Epicopeiidae （Lepidoptera ＞， Aries − Liberales 13 ： 81 84 （in Japanese with English summary ）． Scoble， M ． J．，1992． The Lepidoptera， Fo ， Funct吻 and Diversity． xi ，4 4 pp ． Oxford University Press， New York ． Sugi ， S ．，1964． ［Hostplarlts of Ei ）icof｝eia hainesii Ho11and ］． κ o η 砂露 32 ： 405 （in Japanese）． ’ Sugi， S．（ed ．），1987、 La7va召 げ larger Moths in ノ砂α η ．453 pp ．（120 pls）． Kodansha ， Tokyo 、（ln Japanese with English summary ）．
摘 要 ・ ・ 台 湾 産 ア ゲ ハ モ ドキ ガ 科 の 生 活 史 （顔 聖 紘 穆 家 宏 虐 家 龍 ）
ハ モ mencia ハ モ − 台 湾 に は2種 の ア ゲ ドキ （EpicoPeia Moore ， オ ナ ガ ア ゲ ドキ とEhainesiimatSu
murai ア ゲ ハ モ ド が る そ れ ら は や で 生 が か っ て い る の の Okano ， キ ） 分 布 す ． 日本 中 国 活 史 分 も ，
で は に べ ら れ て い な か っ た ， は 1992 か ら 1993 に か け て こ れ ら の で 台湾 充 分 調 私 達 年 年 ， 両 種 台 湾 の 生 活 史 を 調 べ る こ と が で き た の で ， こ こ に 報 告 し た ．
E ）icoPein mencia Moore オ ナ ガ ア ゲ ハ モ ド キ
食樹 は ニ レ 科 の ア キ ニ レ ． こ れ は 中国 お よ び 日本 （対 馬） で 知 られ る 食 樹 と 同 じ で あ る．卵 は食 樹 の 葉 の 裏面 に ま と め て 産 み 付 け ら れ ，幼 虫 は 終齢 （6 齢） ま で 白色 の 蝋状 物質 を ま と う．蛹化 は 白 い − 鑞物 質 で 覆 わ れ た 柔 ら か い 繭 内 で 行 な わ れ る ．成虫 は 4 月 か ら 10 月 ま で 見 ら れ 年 2 3 化 ．台湾 − ， 土 の 500 2 000m ま で の カ シ に る が の に さ て で る ． 全 標 高 ， 常緑 帯 分布 す ， 食樹 分 布 限定 れ 局所 的 あ
EpicoPela hainesii matsum π rai Okano ア ゲ ハ モ ド キ
の た ミ ズ マ ノ マ シ 私達 確認 し 食樹 は キ科 の ミズ キ ， ク ミズ キ ，ヤ ボ ウ で ， 日本 で の 記 録 と同 じで あ る ．卵 は ，前種 同様 ，食樹 の 葉裏 に ま と め て 産 み 付 け ら れ ，前種 よ りや や 小 さ い ．幼 虫 は終 齢 （6 ま で の ま と が 3 の よ い つ 齢） 白色 蝋 状物 質 を う ， 齢以 降 蝋物 質 の 分 泌 は前種 り も多 く， く か の 体 節 で は い 毛 は よ ス マ ー で る ． は 4 か 10 で 2 ． 細 束状 ，蛹 前種 り も ト あ 成 虫 月 ら 月 ま 見 られ ， 年 化 台 − 湾 の 北部 ，中部 ，東部 の 標 高 500 2，000 m ま で の い く っ か の 産地 に 限 っ て 分布 す る ．
［文責 ： 吉 本 浩］
（Accepted June 1，1995）
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