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Journal of the Lepidopterists' Society

Literature Cited

Cordero, C. 1993. The courtship behavior of Callophrys xami (Lycaenidae). J. Res. Lepid. 32:99-106.

---------. 1998. Ecologia del Comportamiento Sexual de los Machos

de la Mariposa Callophrys xami (Lycaenidae), con Algunas Consideraciones Acerca de la Evolution del Semen de los In-sectos. Doctoral Thesis, UACPyP/CCH, UNAM, Mexico.

Cordero, C, R. Macias & G. Jimenez. 2000. The number of copulations of territorial males of the butterfly Callophrys xami (Lycaenidae). J. Res. Lepid. 35:78-89.

Cordero, C. & J. Soberon. 1990. Non-resource based territoriality in males of the butterfly Xamia xami (Lepidoptera: Lycaenidae). J. Insect Behav. 3:719-732.

Drummond III, B. A. 1984. Multiple mating and sperm competition in the Lepidoptera, pp. 291-370. In Smith, R. L. (ed.), Sperm competition and the evolution of animal mating systems. Academic Press, New York.

Eberhard W. G. 1996. Female control. Sexual selection by cryptic female choice. Princeton University Press, Princeton, U.S.A.

Simmons, L. W. & M. T. Siva-Jothy. 1998. Sperm competition in insects: mechanisms and the potential for selection, pp. 341-434. In Birkhead, T. R. & A. P. M0ller (eds.), Sperm competition and sexual selection. Academic Press, London.

Carlos Cordero], CEAMISH, Universidad Autonoma del Es-tado de Morelos, Av. Universidad 1001, Col. Chamilpa, Cuernavaca 62210, Morelos, Mexico.

Received for publication 7 January 2001; revised and accepted 28 November 2001

Current address. Departamento de Ecologia Evolutiva, Institute de Ecologia, UNAM, Apdo. Post. 70-275, Ciudad Universitaria, Coyoacan 04510, Mexico, DF, Mexico.

Journal 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 demoleus L. is widely distributed in the tropical and subtropical regions of Asian continent, Australia, and the islands of Taiwan, Hainan, New Guinea, and Lesser Sunda Islands (Sumba, Flores and Alor), but it had been lacking in Sumatra, Java, Borneo, the Philippines, and the Moluccas until its recent invasion of these islands (Corbet & Pendlebury 1978, 1992). Sumatra received ssp. malayanus Wallace from the Malay Peninsula and the Philippines ssp. libanius Fruhstorfer from Taiwan in the 1960-70's (Jumalon 1968, Hiura 1973, Miyata 1973, Tsukada & Nishiyama 1980). These two subspecies can be easily discriminated based on the wing markings; Fruhstorfer (1908), in his description of ssp. libanius, stated that the Taiwanese specimens appear darker than the nominate subspecies specimens from "Tonkin." The dark appearance of ssp. libanius is mainly due to the fact that the yellow spots in spaces la and lb of forewing upper surface are always narrow, whereas these spots are always very broad in ssp. malayanus, though as was not mentioned in the Wallaces (1865) description of the subspecies (Figs. 1-4).

The Malay subspecies was confirmed to have established its population in Java, supposedly having invaded from Sumatra during late 1980s (Kato 1989, Matsumoto & Noerdjito 1996). The species has also been found from Borneo. Otsuka (1988) illustrated a male and a female specimen of P. demoleus from Keningau, Sabah, without mentioning that the records were new in Borneo. Although Otsuka (1988) did not identify the subspecies, the illustrated specimens exhibited typical characters of ssp. libanius. Ishii (1987,

1991) identified P. d. libanius from Sandakan, Sabah, and he mentioned (Ishii 1991) that he saw only a few individuals of this species in 1981, while he found many individuals of the same species in 1983, and suggested that the Taiwanese subspecies may have invaded from the Philippines during the early 1980?s.

Table 1. Year of first record and presumed origin of the Papilio demoleus populations recently established in the Southeast Asian Islands.

* Except old sporadic records which are unrelated to the present population (see Jumalon 1968, Moonen 1991 for further details).

** Tsukada & Nishiyama (1980) first stated occurrence of the species in Sumatra without indicating earliest collection data.

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m w

Figs. 1-4. Papilio dernoleus collected in Borneo and Bali: 1, a male with characters of ssp. malayanus (Bukit Soeharto, East Kalimantan, 16 Nov. 1998); 2, a male with characters of ssp. lihanius (Bukit Soeharto, East Kalimantan, 23 Nov. 1998); 3, a female with intermediate characters between ssp. malayanus and ssp. lihanius (Tawau, Sabah, 11 June 1996); 4, male P. d. malayanus (Nusa Dua, Bali, 17 February 1991).

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Journal of the Lepidopterists' Society

TAIWAN

SUMATRA

JAVA

SUMBA"

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Fig. 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 from Talaud, Sangihe and Sula Islands of the Moluccas (Tsukada & Nishiyama 1980, Corbet & Pendlebury 1992), again suggesting invasion via the Philippines (Table l,Fig.5).

Recently, I confirmed P. d. malayanus had invaded into Borneo (including Indonesian Kalimantan) and Bali as described below.

P. demoleus in Borneo. I collected four males and one female at Luasong (near Tawau) and one female at Tawau in Sabah, Malaysia, in 1996 (the specimens are kept at Luasong Forestry Centre, Luasong, Sabah). Hiroshi Makihara of Forestry and Forest Products Research Institute (FFPRI), Japan, also collected 18 males and three females at Bukit Soeharto (near Samarinda), East Kalimantan, Indonesia, in 1998 and showed them to me for examination (the specimens are temporarily kept by Makihara and to be kept elsewhere in Indonesia in the future). Ten of these 27 specimens looked like ssp. libanius (1 6 Luasong, 19

March 1996; 1 C) Tawau, 11 June 1996; 1 6 Bukit Soeharto, 6 October 1998; 4 6 Bukit Soeharto, 23 November 1998; 2 6 Bukit Soeharto, 8 December 1998; 1 6 Bukit Soeharto, 30 November 1998), eight looked like ssp. malayanus (1 6 Luasong, 19 March 1996; 1 6 Luasong, 25 May 1996; 1 6 Bukit Soeharto, 12 Januaiy 1999; 2 6 Bukit Soeharto, 6 October 1998; 2 9 Bukit Soeharrto, 2 November 1998; 1 6 Bukit Soeharto, 9 November 1998), and nine exhibited various intermediate characters between the two (1 6 Luasong, 1 June 1996; 1 9 Luasong, 28 May 1996; 1 6 Bukit Soeharto, 6 October 1998; 1 c5, 1 9 Bukit Soeharto, 6 November 1998; 1 6 Bukit Soeharto, 9 November 1998; 1 6 Bukit Soeharto, 16 November 1998; 2 6 Bukit Soeharto, 23 November 1998), indicating that the population in Borneo is now a mixture of the two subspecies (Figs. 1-3). P. demoleus in Bali. I found two males of P. demoleus collected by Keizi Kiritani at Nusa Dua, Bali, Indonesia, on 17 February, 1991, in the collection of National Institute of Agro-Environmental Sciences

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(NLAES), Japan. These specimens had typical characters of ssp. malayanus (Fig. 4). To my knowledge, this is the first record of the species from Bali. Although there has been no more information of this species in Bali, it seems likely that the species is established there.

Factors favoring spread of P. demoleus. Papilio demoleus frequents in and around villages and urban areas. It is originally a species of monsoon regions and prefers open habitats to thick shadowy rain forests as used to dominate in Sumatra, Java, Bali, Borneo and the islands of the Philippines. Hiura (1973) suggested that this habitat preference is the reason why this species had never occurred in the rain forest regions in Sunda Land and the Philippines before, while recent large scale deforestation in the Philippines prepared suitable habitats for the species. I share the same view with him and believe that the same logic applies to the recent establishment of the P. demoleus populations in Sumatra and Borneo. In densely human populated Java and Bali, forests have been exploited earlier and suitable habitats for P. demoleus have long been prevalent. However, Java and Bali had been far derived from the nearest P. demoleus population, until effects of extensive logging and consequent human activities, e.g., fire, shifting cultivation, illegal settlements, cattle breeding, etc. in the forests (or ex-forests) became conspicuous in Sumatra in the 1960-70s. After this stage, Sumatra played a role of stepping stone for P. d. malayanus to invade into Java, and then Java played a similar role between Sumatra and Bali. The occurrence of P. demoleus in Bali in 1991 indicates that the invasion into Bali took place almost immediately after its establishment in Java. The Malay subspecies in Borneo could have invaded either from Sumatra or Java, or directly from Malay Peninsula (Fig. 5).

Then, a question arises; why the Australian subspecies (i.e., ssp. sthenelus Macleay of Australia and Sumba, ssp. sthenelinus Rothschild of Flores and Alor and ssp. novoguineensis Rothschild of New Guinea) would not spread? Food plant availability could be the factor favoring spread of the Asian subspecies, not the Australian subspecies. The Asian subspecies feed on oranges, Citrus hystrix DC, C. aurantifolia Swingle, C. amblyocarpa Ochse, etc. (Rutaceae; Corbet & Pendlebury 1992, Matsumoto & Noerdjito 1996) which are very commonly planted either as a crop or an ornament in towns, villages and illegal settlements in the forest areas. The Australian subspecies feed on wild leguminous plants of the genus Psoralea L.: i.e., P. tenax Lindl., P. patens Lindl, P. cinerea Lindl., P. leu-cantha F Muell. and P. pustulata F Muell. in Australia (Common & Waterhouse 1981) and P. badocana

Benth. in Papua New Guinea (Parsons 1999) (no food plant record available for the Lesser Sunda populations). Although Psoralea plants occur in the Sunda Islands (Bentham & Mueller 1967), there has been no evidence to indicate increase of these plants favored by recent environmental changes. The Citrus feeding Asian subspecies, on the other hand, are now expanding its range in the Malay Archipelago, being favored by increase of habitats with artificially planted Citrus hosts.

L thank H. Makihara of FFPRI and NIAES for giving opportunity for me to examine their specimens. Scott E. Miller and Carla M. Penz reviewed the manuscript and made helpful comments and suggestions.

Literature Cited

Bentham, G. & F. Mueller. 1967. Flora Australiensis: a description of the plants of the Australian Territory. A. Asher & Co., Amsterdam. 521 pp.

Common, l. F B. & D. F. Waterhouse. 1981. Butterflies of Australia. Angus & Robertson, Sydney. 682 pp. + 20 pis.

Corbet, A. S. & H. M. Pendlebury. L978. The butterflies of the Malay Peninsula. 3rd ed. (rev. by J. N. Eliot). Malayan Nature Society, Kuala Lumpur. 578 pp. + 35 pis.

----------. 1992. The butterflies of the Malay Peninsula. 4th ed. (rev.

by J. N. Eliot). Malayan Nature Society, Kuala Lumpur. 595 pp. + 69 pis.

Fruhstoreer, H. 1908. Lepidopterologisches Pele-Mele. Ent. Zeitschr. 22:140-141

Hiura, I. 1973. Butterflies flying across the sea. Soju Shobo, Tokyo. 200 pp.

Ishit, M. 1987 Diapause potential in tropical Papilionids (Lepi-doptera: Papilionidae). Appl. Ent. Zool. 22:114-15.

----------. 1991. Tropics for butterflies, pp. 59-84. In Hidaka, T. &

M. Ishii (eds.), Animals and plants in Borneo—studying their lives in tropical forests. Tokyo Kagaku Dojin, Tokyo. 248 pp.

Jumalon, J.N. 1968. A comment on the new papilionid from the Philippines. Tyo to Ga 19:105-109.

Kato, S. 1989. Notes on Papilio demoleus Linnaeus collected in Java, Indonesia (Lepidoptera, Papilionidae). Tyo to Ga "40:189-191

Matsumoto, K. & A. W. Noerdjito. 1996. Establishment of Papilio demoleus L. in Java. J. Lepid. Soc. 50:139-140.

MfYATA, A. 1973. Notes on Princeps demoleus libanius Fruhstorfer in the Philippines. Tyo to Ga 24:37-41.

Moonen, J. J M. 1991. Papilio demoleus L. in Java (Lep.; Papilionidae)" Tyo to Ga 42:93-94.

Otsuka, K. 1988. Butterflies of Borneo. Vol. 1 Tobishima Corp., Tokyo, x + 61 pp. (in Japanese) + xix + 61 pp. (in English) + 80 pis.

Parsons, M 1999. Butterflies of Papua New Guinea. Academic Press, San Diego. 736 pp. + 136 pis.

Tsukada, E. & Y. Nishjyama. 1980. Butterflies of South East Asian islands. I. Papilionidae. Plapac, Tokyo. 459 pp.

Wallace, A. R. 1865. On the phenomena of variation and geographical distribution as illustrated by the Papilionidae of the Malayan region. Transactions of the Linnean Society 25:1-71 + pis. I-VIIT.

Kazuma Matsumoto, Tama Forest Science Garden, Forestry and Forest Products Research Institute, Todori-cho, Hachidji, Tokyo 193-0843, Japan.

Received for publication 7 February 2001; revised and accepted 3 December 2001