108 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY

LITERATURE CITED EBERHARD W. C. 1096. Female control. Sexual selection by cryptic female choice. Princeton University Press, Princeton, U.SA CORDERO, C. 1993. The courtship behavior of Callophrys xami SIMMONS , L. W. & M. T. SIVA-JOTHY. 1998. Sperm competition in (Lycaenidae). Res. Lepid. 32:99-106. J. insects: mechanisms and the potential for selection, pp, --. 199R. Ecologia del Comportamiento Sexual de los Machos 341-434. In Birkhead, T. R. & A, P. M011er (eds.), Sperm com­ de la Mariposa Callophrys xami (Lycaenidae), con Algunas petition and sexual selection. Academic Press, London. Consideraciones Acerca de la Evoluci6n del Semen de los In­ sectos. Doctoral Thesis, UACPyP/CCH, UNAM, Mexico. CARLOS CORDERO ' , CEAM ISH, Universidad Aut6noma del Es­ CORDERO, c. , R. MACiAS & C. JIMENEZ. 2000. The number of cop­ tado de Morelos, Av. Universidad 1001, Col. Chamilpa, Cuemavaca ulations of territorial males of the butterfly Callophrys xami 622]0, Morelos , Mexico. (Lycaenidae). J. Res, Lepid. 35:7R-R9. Received for publication 7 January 2001; revised and accepted 28 & CORDERO, C. J. SOBERON. 1990. Non-resource based territorial­ November 2001 ity in males of the butterfly Xamia xami (Lepidoptera: Ly­ caenirlae) J. Insect Behav. 3:719-732. DRUMMOND III, B. A. 1984. Multiple mating and sperm competi­ 'Current address. Departamento de Ecologia Evolutiva, Instituto tion in the Lepidoptera, pp. 291- 370. In Smith, R. L. (eeL ), de Ecologia, UNAM, Apdo. Post. 70-275, Ciudad Universitari(1, Sperm competition and the evolution of animal mating systems. Coyo(1c(1n 04570, M.ixico, DF, Mexico. Academic Press, New York.

.Tournai of the Lepidopterists' Society .56(2).2002. 108- 111

PAPILIO DEMOLEUS (PAPILIONIDAE) IN BORNEO AND BALI Additional key words: Malaysia, Indonesia, Malay Archipelago, invasion, deforestation.

Papilio derrwleus L. is widely distributed in the trop­ 1991) identified P d. libanius from Sandakan, Sabah, ical and subtropical regions of Asian continent, Aus­ and he mentioned (Ishii 1991) that he saw only a few tralia, and the islands of Taiwan, Hainan, New Guinea, individuals of this species in 1981, while he found and Lesser Sunda Islands (Sumba, Flores and Alor), many individuals of the same species in 1983, and sug­ hut it had been lacking in Sumatra, Java, Borneo, thc gested that the Taiwanese subspecies may have in­ Philippines, and the Moluccas until its recent invasion vaded from the Philippines during the early 1980's. of these islands (Corbet & Pendlebury 1978, 1992), Sumatra received ssp. malayanus Wallace from the TABLE 1. Year of first record and presumed origin of the Papilio Malay Peninsula and the Philippines ssp, libanius demo/eus populations recently established in the Southeast Asian Is­ Fruhstorfer from Taiwan in the 1960-70's (Jumalon lands.

1968, Hiura 1973, Miyata 1973, Tsukada & Nishiyama Year of Presumed 1980), These two subspecies can be easily discrimi­ Island first record origin Literature nated based on the wing markings; Fruhstorfer (1908), Lllzon 1967* Taiwan Tumalon (1968) in his description of ssp, libanius, stated that the Tai­ Cebu 1968? Taiwan Hiura (197.3 ) wanese specimens appear darker than the nominate Leyte 1968? Taiwan Hiura (1973) subspecies specimens from "Tonkin," The dark appear­ Palawan 1969 'Iaiwan Hiura (1973) ance of ssp, libanius is mainly due to the fact that the Negros 1969? Taiwan Miyata (197:3 ) Mindanao 1969 Taiwan Miyata (1973) yellow spots in spaces la and Ib of forewing upper sur­ Mindoro 1971 Taiwan Hillra (1973) face are always narrow, whereas these spots are always Talallrl Unknown** Taiwan Tsukada & very broad in ssp, malayanus, though as was not men­ Nishiyama (1980) tioned in the Wallace's (1865) description of the sub­ Sangihe Unkno"l1** Taiwan Tsukada & species (Figs, 1-4). Nishiyama (191l0) Sula Unknown** Taiwan Tsukada & The Malay subspecies was confirmed to have estab­ Nishiyama (l91l0) lished its population in Java, supposedly having in­ Borneo 1983 Taiwan Ishii (1987) vaded from Sumatra during late 1980's (Kato 1989, 1996 Malay Peninsula This study Matsumoto & Noerdjito 1996). The species has also Sumatra U nkno"l1 ** Malay Peninsula Tsukada & been found from Borneo. Otsuka (1988) illustrated a Nishiyama (1980) Java 1988* Malay Peninsula Kato (1989) male and a female specimen of P demoleus from Bali 1991 Malay Peninsula This study Keningau, Sahah, without mentioning that the records * Except old sporadiC records which are unrelated to the present were new in Borneo. Although Otsuka (1988) did not population (see Jumalon 1968, Moonen 1991 for further details). identify the subspecies, the illustrated specimens ex­ ** Tsukada & Nishiyama (1980) nrst stated occurrence of the hibited typical characters of ssp, libanius, Ishii (1987, species in Sumatra "ithout indicating earliest collection data, VOLUME .56, NUMBER 2 109

F[Gs. 1-4. Papilio demoleLls collected in Borneo and Bali: 1, a male with characters of ssp. malayanl1s (Bukit Soeharto. East Kalimantan, 16 Nov. 1991:»; 2, a male with characters of ssp. libanil1s (Bukit Soeharto, East Kalimantan, 23 Nov. 1998); 3, a female with intermediate char­ acters behveen ssp. malayanus and ssp. libanius (Tawau, Sabah, 11 June 1996); 4, male P d. malayanl1s (Nusa Dua, Bali, 17 February L991 ). 110 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY

TAIWAN

PHILIPPINES

MINDANAO

SANGIHEt '1 ff...... • b b ~ SUlAC) ~

o FLORES ALORP '. .11

D~'~'~~ AUSTRALIA SUM~ • t?

FIC, 5. A map of Southeast Asia, showing islands recently invaded by Papilio demoleus. Arrows indicate presumable direction of the invasion.

Furthermore, P d. libanius has been also recorded March 1996; 1 C) Tawau, 11 June 1996; 1 cS Bllkit Soe­ from Talaud, Sangihe and Sula islands of the Moluccas harto, 6 October 1998; 4 cS Bukit Soeharto, 2.3 Novem­ (Tsukada & Nishiyama 1980, Corbet & Pendlebury ber 1998; 2 cS Bukit Soeharto, 8 December 1998; 1 cS 1992), again suggesting invasion via the Philippines Bukit Soeharto, 30 November 1998), eight looked like (Table 1, Fig . .5). ssp. malayanus (1 0 Luasong, 19 March 1996; 1 0 Lu­ Recently, I confirmed P d. malayanus had invaded asong, 2.5 May 1996; 1 0 Bukit Soeharto, 12 January into Borneo (including Indonesian Kalimantan) and 1999; 2 cS Bukit Soeharto, 6 October 1998; 2 ? Bukit Bali as described below. Soeharrto, 2 November 1998; 1 0 Bukit Soeharto, 9 P. demoleus in Borneo. I collected four males and November 1998), and nine exhibited various interme­ one female at Luasong (near Tawau) and one female at diate characters between the two (1 0 Lllasong, 1 June Tawau in Sabah, Malaysia, in 1996 (the specimens are 1996; I ? Lllasong, 28 May 1996; 10 Bukit Soeharto, 6 kept at Luasong Forestry Centre, Luasong, Sahah). October 1998; 1 0, 1 ¥ Bukit Soeharto, 6 November Hiroshi Makihara of Forestry and Forest Products Re­ 1998; 1 0 Bukit Soeharto, 9 November 1998; 1 0 Bukit search Institute (FFPRI), Japan, also collected 18 Soeharto, 16 Novemher 1998; 2 cS Bukit Soeharto, 2:3 males and three females at Bukit Soeharto (near November 1998), indicating that the population in Bor­ Samarinda), East Kalimantan, Indonesia, in 1998 and neo is now a mixture of the two subspecies (Figs. 1-:3). showed them to me for examination (the specimens P. demoleus in Bali. I found two males of P de­ are temporarily kept by Makihara and to be kept else­ mo/eus collected by Keizi Kiritani at Nusa Dua, Bali, where in Indonesia in the future). Ten of these 27 Indonesia, on 17 Fehruary, 1991, in the collection of specimens looked like ssp. libanius (l cS Luasong, 19 National Institute of Agro-Environmental Sciences VOLUME 56, N U MBE K 2 HI

(N LAES ), Japan. These specimens had typical charac­ Benth. in Papua New Guinea (Parsons 1999) (no fooel ters of ssp. rnalayanus (Fig. 4). To my knowledge, this plant record available for the Lesser Sunda popula­ is the first record of the species from Bali. Although tions). Although Psoralea plants occur in the Sunda Is­ there has been no more information of this species in lands (Bentham & Mueller 1967), there has been no Bali, it seems likely that the species is established evidence to indicate increase of these plants favored there. by recent environmental changes. The Citrus feeding Factors favoring spread of P. demoleus. Papilio Asian subspecies, on the other hand, are now expand­ demoleus frequents in and around Villages and urban ing its range in the Malay Archipelago, being favored areas. It is originally a species of monsoon regions and by increase of habitats with artificially planted Citrus prefers open habitats to thick shadowy rain forests as hosts. used to dominate in Sumatra, Java, Bali, Borneo and the islands of the Philippines. Hiura (1973) suggested 1 thank H . Makihara of FFPRI and NIAES for giving opportu­ nity for me to examine their specimens. Scott E. Miller and Carla M. that this habitat preference is the reason why this Pem. reviewed the manuscript and made helpful comments and sug­ species had never occurred in the rain forest regions in gestion s. Sunda Land and the Philippines before, whil e recent large scale deforestation in the Philippines prepared LITERATURE CITED suitable habitats for the species. I share the same view BENTHAM, G. & F. MUELLER. 1967. Flora Australiensis: a descrip­ tiOll of the plants of the Au stralian Territory. A. Asher & Co., with him and believe that the same logiC applies to the Amsterdam. 521 pp. recent establishment of the P derrwleus populations in COMMON , I. F'. B. & D. F. WATERHOUSF.. 1981. Butterflies of AllS­ Sumatra and Borneo. In densely human populated tndia. Angus & Robertson, Sydney. 682 pp. + 20 pis. CORB ET, A. S. & H. M. PE NDLEBURY. L978. The butterfli es of the Java and Bali, forests have been exploited earlier and Malay Peninsula. :3rd ed. (rev. by J. N. Eliot). Malayan Nature suitable habitats for P dernoleus have long been preva­ Society, Kuala Lumpur. 578 pp. + 3.5 pIs. lent. However, Java and Bali had been far derived - - . 1992. The butterflies of the Malay Peninsula. 4th ed. (rev. by J. 1\. Eliot). Malayan Nature SOCietY, Kuala Lumpur. 595 pp. from the nearest P dernoleus population, until effects + 69 pIs. of extensive logging and consequent human activities, FR UHSTO I\FI<:K , H. Hl08. Lepidopterologiscbcs pele-Mele. Ent. e.g., fire, shifting cultivation, illegal settlements, cattle Zeitschr. 22: 140-14] H IURA, I. ] 973. Butterflies flying across the sea. SOjl! Shobo, Tokyo. breeding, etc. in the forests (or ex-forests ) became 200 pp. conspicuous in Sumatra in the 1960-70's. After this 1SHlT , M. 1987 Diapallse potential in tropical Papilionids (Lepi­ stage, Sumatra played a role of stepping stone fo r P d. doptera: Papi!ionidae). App!. Ent. Zool. 22:11 4-15. - - -. J 991. Tropics for butterflies, pp. 59- 84. In Hidaka, T. & malayanus to invade into Java, and then Java played a M. Ishii (eds.) , Animals and plants in Borneo-studying their similar role between Sumatra and Bali. The occur­ li ves in tropical forests. Tokyo Kagaku Dojin , Tokyo. 248 pp. rence of P dernoleus in Bali in 1991 indicates that the JUMA LO N, J. N. 1968. A comment on the new papilionid fro m the Philippines. Tyo to Ca 19:105-109. invasion into Bali took place almost immediately after KATO, S. 1989. Notes 011 Papilio demoleus Linnaeus collected in its establishment in Java. The Malay subspecies in Bor­ Java, Indonesia (Lepidoptera, Papilionidae). Tyo to Ga neo could have invaded either from Sumatra or Java, 40: 189- 191 MATSUMOTO . K. & A. W. :-JOE RDjlTO. ] 996. Establishment of Fa ­ or directly from Malay Peninsula (Fig. 5). r,ilio demoleus L. in Java. J. Lepid. Soc. 50:139-140. Then, a question arises; why the Australian sub­ M IYATA, A. 1973. Notes on Princeps demo/eus libanius Fruhstorfe r species (i. e., ssp. sthenelus Macleay of Australia and in the Philippines. Tyo to Ca 24::17-41. MO ON EN, J. J M. 1991. Papilio demo/eus L. in Java (Lep.: Papil­ Sumba, ssp. sthenelinus Rothschild of Flores and Alor ion idae). Tyo to Ga 42:93-94. and ssp. novoguineensis Rothschild of New Guinea) OTSU KA, K. 1988. Butterflies of Borneo. Vol. 1 Tobishima Corp., would not spread? Food plant avail ability could be the To~'Y0 ' x + 61 pp. (in Japanese) + xix + 61 pp. (in E nglish ) + 80 pis. factor bvoring spread of the Asian subspecies, not the PARSONS , M 1999. Butterfli es of Papua New Guinea. Academic Australian subspecies. The Asian subspecies feed on Press, San Diego. 7:36 pp. + 136 pIs. oranges, Citrus hystrix DC., C. aurantifolia SWingle, TSU KADA, E. & Y. NISHIYAM A. 1980. Butterflies of South East Asian isl:lIlds. 1. Papilionidae. Plapac, Tokyo. 459 pp. C. amblyocarpa Ochse, etc. (Rutaceae; Corbet & WALLAC E. A. R. 1865. On the phenomena of variation and geo­ Pelldl ebury 1992, Matsumoto & Noerdjito 1996) graphical distribution as illustrated by the Papilionidae of the which are very commonly planted either as a crop or Malayan region. Transactions of the Linnean SOCiety 25: \- 71 + pis. I- VIlT. all ornament in towns, Villages and illegal settlements in the forest areas. The Australian subspecies feed on KAZUM A MATSliMOTO, Tam.a Forest Science Carden, Forestry lind wild leguminous plants of the genus Psoralea L.: i.e., P Forest Products Resea1'Ch Institute, Todori-chO , Hachiaji, Tokyo 193- 0843, Japan. tenox Lind!. , P patens Lind!., P cinerea LincH., P leu­ cantha F. Muel!. and P pustulata F. M uell. in Australia Received for puhlication 7 Febmary 2001 ; revised (Jnd accepted (Common & Waterhouse 1981 ) and P badocana 3 December 2001