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Molecular Phylogeny of Birdwing Butterflies of the Tribe Troidini (Lepidoptera: Papilionidae) - Using All Species of the Genus Ornithoptera

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Molecular Phylogeny of Birdwing Butterflies of the Tribe Troidini (Lepidoptera: Papilionidae) - Using All Species of the Genus Ornithoptera Biogeography 2. 103-111. Aug. 28, 2000 Molecular Phylogeny of Birdwing Butterflies of the Tribe Troidini (Lepidoptera: Papilionidae) - Using All Species of the Genus Ornithoptera - l 2 Sadaharu Morinaka , Nobuhiro Minaka , Masayuki SekiguchP, Erniwati4, Siti Nuramaliati Prijono\ Ida Ketut GinarsaS, Tadashi Miyata6 and Toshitaka Hidaka7 I Aichi Study Center, The University of the Air, Yagotohonmachi, 10 1-2, Showa-ku, Nagoya 466-0825, Japan 2 National Institute of Agro-Environmental Sciences, Kannon-dai 3-1-1, Tsukuba, Ibaraki 305-8604, Japan J National Institute of Neuroscience, Ogawahigasimachi 4-1-1, Kodaira-shi, Tokyo 187-8502, Japan 4 Biological Division, Research and Development for Biology, Indonesian Institute of Sciences, Gedung Widyasatwaloka J1. Raya Cibinong, Cibinong Bogor, 16002 Indonesia 5 Pk. Jelati, Yeh Sumbul, Mendoyo Bali, 82261 Indonesia h Department of Biological Mechanisms and Functions, Graduate School of Bioagricultural Sciences, Nagoya University, Huro-cho, Chikusa-ku, Nagoya 464-8601, Japan 7 The University of Shiga Prefecture, Hassaka 2500, Hikone, Shiga, 522-8533 Japan Corresponding address: S. Morinaka, Totsukahasami-cho 11-20, Kawaguchi-shi, Saitama-ken, 333-0805 Japan (e-mail: [email protected]) Abstract. The molecular phylogeny of birdwing butterflies based on the NOS gene of mitochondrial DNA (mtDNA) was studied by the maximum-likelihood method. maximum-parsimony method and neighbor- joining method. All species of the genus Omithoptera, some species of the genus Troides and other species representing most genera of the tribe Troidini were used. Results suggest the genus Omilhoptera forms a clade with Troides, supporting the previous results (Morin aka el al., 1999). In the case of the genus Omilhoptera, each of the subgenera Omithoptera and Schoenbergia was found to be monophyletic, however, phylogenetic relationships of O. alexandrae and O. vicloriae (the subgenus Aetheoptera) could not be clarified. In the subgenus Schoenbergia, two clades (0. paradisea + O. meridionalis, and O. goliath + O. chimaera + O. rothschildi) were found, however, the phylogenetic relationships of O. tithonus and other species remain unclarified, so as in the case of the genus Trogonoptera in the tribe Troidini. Key words: Aetheoptera, birdwing butterflies, mitochondrial DNA, molecular phylogeny, NOS, Omithoptera, Ripponia, Schoenbergia, Trogolloptera, Troides, Troidini. Introduction region), Central to South America and Indo- Australian regions. This tribe includes bird wing The tribe Troidini includes about 10 genera, butterflies, which are important for mankind because which inhabit Madagascar Is. (the Afrotropical their fate, except some species adapting to the -103- Molecular Phylogeny of Birdwing Butterflies of the Tribe Troidini countryside, is linked to the existence of tropical treated as hybrids of O. victoriae X O. priamus, and rainforests. They are considered as a symbol O. rothschildi X O. priamus, respectively, and O. exemplifying biodiversity. richmondia and O. euphorion as subspecies of O. Many taxonomic and/or phylogenetic studies on priamus. Permission was obtained for species troidine butterflies have been published (Zeuner, protected by the international contract, "Conservation 1943; Ford, 1944; Munroe & Ehrlich, 1960; Munroe, on International Trade in Endangered Species of Wild 1961; Hancock, 1983; Miller, 1987; Haugum & Low, Fauna and Flora" (C.I.T.E.S.) or other regulations 1978-1985; D' Abrera, 1975; Igarashi, 1979, 1984; when necessary. Tsukada, 1980; Ohya, 1983; Tyler et al., 1994; Fujioka et al., 1997; Parsons, 1999: Morinaka et al., Preparation of mtDNA, amplification and 1999). Their classifications are partly different from sequencing of ND5 gene each other. Morphological characters are useful for Details of DNA extraction, amplification and investigating phylogeny, whereas DNA sequences sequencing techniques for the ND5 gene of mtDNA, encode more discrete characters, determination of are described in Morinaka et ai. (1999). Nucleotide which would yield a high degree of accuracy sequences of the primers are as follows: a, 5'- (Brower, 1996; Su et al., 1998; Caterino & Sperling, CCTGTTTCTGCTTTAGTTCA-3' (Su et al., 1996); 1999; Yagi et ai., 1999, Maekawa et al., 1999). We and B3, 5'-TAACCTCTATATATYTCTCTT-3'. have previously reported that the genera of Troidini excluding the genus Bartus are well united, forming Phylogenetic Analyses one clade; Tl'Oides and Ornithoptera also form one Two or more individuals from an intrataxon were clade (Morinaka et al., 1999). However, the used for examination, except for some species which phylogeny of Trogonoptera and the intrageneric one were difficult to obtain. In the analyses, 877 of the of Ornithoptera remain unclarified. In this study, 924 nucleotide sequences were used (Fig. 1). phylogenetic analyses are carried out using Editing and aligning of the sequence data were Trogonoptera and all species of Ornithoptera. completed utilizing DANASIS (Hitachi Software Engineering Co., Hitachi) and CLUSTAL W Materials and Methods (Thompson et ai., 1994). Phylogenetic trees were constructed by the maximum-likelihood (ML) Taxa used in the study method, maximum-parsimony (MP) method and Taxa used in this study are shown in Table I, neighbor-joining (NJ) method with the evolutionary including their sampling localities and accession distances by Kimura's two-parameter method using numbers in DDBJ/EMBLlGenBank. Species name of PAUP Version 4.0b4a (Swofford, 2000). The Troides, Ornithoptera and Tl'Ogonoptera are based on statistical confidence of each clade was examined Aoki et al. (1998). O. aiotti and O. akakeae are with the bootstrap tests (Efron, 1982; Felsenstein, a: 20 mer 83: 21 mer ... 924 bp .... His UbP I Phe 877 bp N05 Gene Fig. I. Schematic representation of the amplified and coding regions of the NDS gene used in the study. a: 5'-CCTGTTTCTGCTTTAGTTCA-3'; B3: 5'- TAACCTCTATATATYTCTCTT-3'. -104- S. Morinaka, N. Minaka, M. Sekiguchi, Erniwati, S. N. Prijono, l. K. Ginarsa, T. Miyata and T. Hidaka Table I. Specimens used in the study (classified arbitrarily). Genus (Subgenus) Species Locality DDBJI EMBU GenBank Accession Number Parides hahneli Brasil AB027579 Par. elraces Upper HualJaga Y., Peru AB027580 Byasa alcinous Tokyo, Japan AB027583 B. polyeuctes Formosa AB044646 Atrophaneura luchti Jawa Is., Indonesia AB027584 A. nox Bali Is., Indonesia AB027586 Losaria coon Jawa Is., Indonesia AB027587 Pachliopta aristolochiae Bali Is., Indonesia AB027590 Troides (Ripponia) hypolitus Sulawesi Is., Indonesia AB027592 T. (Troides) amphrysus Jawa Is., Indonesia AB027593 T. (T. ) helena Bali Is., Indonesia AB027594 T. (T. ) haliphron Sulawesi Is ., Indonesia AB044649 T. (T. ) prattorum Buru Is., Indonesia AB044650 T. (T. ) staudingeri Lakor Is., Indonesia AB04465 I Ornithoptera (Ornithoptera) priamus Seram Is., Indonesia AB027596 O. (0. ) priamus Malaita Is., Solomon Isis AB027597 O. (0. ) priamus Timika, Irian Jaya AB044656 O. (0. ) priamus I Is., Solomon IsIs AB044657 O. (0. ) croesus Halmahera Is., Indonesia AB027595 O. (0. ) croesus Bacan Is., Indonesia AB044654 O. (0. ) aesacus Obi Is., Indonesia AB044655 O. ( Aetheoptera) alexandrae Papua New Guinea AB044652 O. (A. ) victoriae Malaita Is., Solomon Isis AB027598 O. (A.) victoriae Gela Is., Solomon Isis AB044658 O. ( Schoenbergia) paradisea Timika, Irian Jaya AB027603 O. (S. ) paradisea Arfak Mts, Irian Jaya AB044659 O. (S. ) meridionalis Timika, Irian Jaya AB044653 O. (S. ) goliath Seram Is., Indonesia AB027599 O. (S. ) goliath Arfak Mts, Irian Jaya AB027600 O. (S. ) rothschildi Arfak Mts, Irian Jaya AB027601 O. (S. ) chimaera Wayland Mts, Irian Jaya AB044660 O. (S. ) tithonus Arfak Mts, Irian Jaya AB027602 Trogonoptera brookiana Padang, Sumatera Is. AB044647 Trog. brookiana Cameron Highland, Malaysia AB044648 Papilio protenor Tokyo, Japan AB027604 Pieris (Pieridae, as outgroup) rapae Mie, Japan AB044594 -105 - Molecular Phylogeny of Birdwing Butterflies of the Tribe Troidini P. ro"'poe Ahhrcvialions: r :;llIlding.:r; AI I : A.-fak MiS, lriun Jay" Iio/iphr. (I 8m.;: Bi1(.·tm Is v' T. prof/arum Cam; Cameron Highland Gd : Gclu. Solol11on Isis T f,elello GUll: GumJ<1lcanul. Solomon Isis r gmphl)'slIs Hal: Hnlmnhclu Is T hypolilus Mal: MaJaila, Solomon [sis ,--------- O . o/sxol1drcre Aelileoptera SCI: S!.:ram Is Sum: SUIOi.llcra r;;:--- 0 pad Tim Tim : Tirnika, Ilian Juyu '---- 0 men 'ono/(s golio/h i\rr Sclroe"hergiaJ ,,,. 0.. 0.. o llOSer 0. chimaera ____":.- .:.._-:..--_-_-_-_-_- 0. lolhschildi Omifhopfero L O. filhonus L _"-.. pr/omu!> Tim ' ()d. JflOmU5 Sar . POOIT/US Gu. 0 ill tJ o ptiomus M:t! rJI lOp era o.. aesaCfiS I QoIJO. croesus Bac 12 0. croe5u5 Hoi o.. "it...! /i ,,001 O. vic/a fiae M .1 B !X,I /,JeI 1'-4 B oHnous L_ __ .. A luchli 1---------4% ====- A. nox L ____ '" Pa r. hahne/i Poe ori5/%chioe 91 t coon '-_______-"-' '',' Trag. bfLlOk/al1o :\"'" Trogonoptero Trag brookiono Cam I prolenor -- 001 subslitutions/site a r---------------------------------------------- P ropae T, 510udinge ri f. ho/iphron T prof/arum T he/ella '--------' T ompluysus T /lypo/iIV$ ,--------- () 01 ondtne Aetheoplera " O .... priamus Tim 67 O.pnomus Ser j II 0 priomus Gua O. priomus Mal OrllltllOptera O . oe50CUS O. croe5us Bac O. croesus Hal Omifhop/era O . porodlr!)' T.m '----- 0 me/idiono/is j 0., go/,a/lt Arl' Scl,oel/hergia .12 0 .. 9 Jlo Ser I., 0. '--------- 0 /ilhonus L-----------fiWll" 0.. IIICI(;>1 /oe Gel 0.. vlc/orloe Ml.1 ---------...I ::---E====-EZ-::i. B. po/yeltc/es [ B. "'CIIlOU5
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