Invertebrate Systematics Invertebrate Systematics Society
Systematics, phylogeny and biogeography

When and where did troidine butterflies (Lepidoptera : Papilionidae) evolve? Phylogenetic and biogeographic evidence suggests an origin in remnant Gondwana in the Late Cretaceous

Michael F. Braby A B D , John W. H. Trueman A and Rod Eastwood B C

A School of Botany and Zoology, The Australian National University, Canberra, ACT 0200, Australia.

B Museum of Comparative Zoology, Harvard University, 26 Oxford St, Cambridge, MA 02138, USA.

C Australian School of Environmental Studies, Griffith University, Nathan, Queensland 4111, Australia.

D Corresponding author. Email:

Invertebrate Systematics 19(2) 113-143
Submitted: 9 August 2004  Accepted: 9 March 2005   Published: 28 June 2005


The age, geographic origin and time of major radiation of the butterflies (Hesperioidea + Papilionoidea + Hedyloidea) are largely unknown. The general modern view is that butterflies arose during the Late Jurassic/Cretaceous in the southern hemisphere (southern Pangea/Gondwana before continental breakup), but this is not universally accepted, and is a best guess based largely on circumstantial evidence. The extreme paucity of fossils and lack of modern, higher-level phylogenies of extant monophyletic groups have been major impediments towards determining reliable estimates of either their age or geographic origin. Here we present a phylogenetic and historical biogeographic analysis of a higher butterfly taxon, the swallowtail tribe Troidini. We analysed molecular data for three protein-encoding genes, mitochondrial ND5 and COI–COII, and nuclear EF–1α, both separately and in combination using maximum parsimony (with and without character weighting and transition/transversion weighting), maximum likelihood and Bayesian methods. Our sample included representatives of all 10 genera of Troidini and distant ingroup taxa (Baroniinae, Parnassiinae, Graphiini, Papilionini), with Pieridae as outgroup. Analysis of the combined dataset (4326 bp; 1012 parsimony informative characters) recovered the Troidini as a well supported monophyletic group and the monophyly of its two subtribes, Battina and Troidina. The most parsimonious biogeographic hypothesis suggests a southern origin of the tribe in remnant Gondwana (Madagascar–Greater India–Australia–Antarctica–South America) sometime after the rifting and final separation of Africa in the Late Cretaceous (<90 Mya). Although an ancient vicariance pattern is proposed, at least four relatively recent dispersal/extinction events are needed to reconcile anomalies in distribution, most of which can be explained by geological and climatic events in South-east Asia and Australia during the late Tertiary. Application of a molecular clock based on a rate smoothing programme to estimate various divergence times based on vicariance events, revealed two peculiarities in our biogeographic vicariance model that do not strictly accord with current understanding of the temporal breakup of Gondwana: (1) the troidine fauna of Greater India did not become isolated from Gondwana (Antarctica) until the end of the Cretaceous (c. 65 Mya), well after Madagascar separated from Greater India (84 Mya); and (2) the faunas of Greater India, Australia and South America diverged simultaneously, also at the K/T boundary. A recent published estimate of the time (31 Mya) of divergence between Cressida Swainson (Australia) and Euryades Felder & Felder (South America) is shown to be in error.


Ackery P. R. , Smith C. R. , and Vane-Wright R. I. (1995). ‘Carcasson’s African Butterflies. An Annotated Catalogue of the Papilionoidea and Hesperioidea of the Afrotropical Region.’ (CSIRO Publishing: Melbourne, Australia.)

Ackery P. R. , de Jong R. , and Vane-Wright R. I. (1999). The Butterflies: Hedyloidea, Hesperioidea and Papilionoidea. In ‘Lepidoptera, Moths and Butterflies. Volume 1. Evolution, Systematics and Biogeography’. (Ed. N. P. Kristensen.) pp. 263–300. (de Gruyter: Berlin, Germany.)

Anderson J. M. Anderson H. M. 1984 The fossil content of the Upper Triassic Molteno Formation, South Africa. Palaeontologia Africana 25 39 59

Archer M. , Hand S. J. , and Godthelp H. (1991). ‘Riversleigh, The Story of Animals in Ancient Rainforests of Inland Australia.’ (Reed Books: Melbourne, Australia.)

Australian Heritage Commission (1986). ‘Tropical Rainforests of North Queensland. Their Conservation Significance. Special Australian Heritage Publication Series No. 3.’ (Australian Government Publishing Service: Canberra, Australia.)

Barker F. K. Lutzoni F. M. 2002 The utility of the incongruence length difference test. Systematic Biology 51 625 637

Barlow B. A. (1981). The Australian flora: its origin and evolution. In ‘Flora of Australia. Volume 1. Introduction’. (Ed. A. S. George.) pp. 25–75. (Australian Government Publishing Service: Canberra, Australia.)

Barlow B. A. (1983). Biogeography of Loranthaceae and Viscaceae. In ‘The Biology of Mistletoes’. (Eds M. Calder and P. Bernhardt.) pp. 19–46. (Academic Press: Sydney.)

Braby M. F. (2000). ‘Butterflies of Australia. Their Identification, Biology and Distribution.’ (CSIRO Publishing: Melbourne, Australia.)

Bremer K. 1994 Branch support and tree stability. Cladistics 10 295 304 DOI

Briggs J. C. 2003 The biogeography and tectonic history of India. Journal of Biogeography 30 381 388 DOI

Brower L. P. (1984). Chemical defense in butterfleis. In ‘The Biology of Butterflies. Symposium of the Royal Entomological Society of London. Number 11’. (Eds R. I. Vane-Wright and P. R. Ackery.) pp. 109–133. (Academic Press: London, UK.)

Brown F. M. 1976 Oligodonta florissantensis, gen. n., sp. nov. (Lepidoptera: Pieridae). Bulletin of the Allyn Museum 37 1 4

Brown K. S. J. (1987). Biogeography and evolution of Neotropical butterflies. In ‘Biogeography and Quaternary History in Tropical America’. (Eds T. C. Whitmore and G. T. Prance.) pp. 66–104. (Clarendon Press: Oxford, UK.)

Carpenter F. M. (1992). ‘Treatise on Invertebrate Paleontology. Part R. Arthropoda 4. Volume 4: Superclass Hexapoda.’ (The Geological Society of America, Inc. and The University of Kansas: Boulder/Lawrence, KA, USA.)

Castoe T. A. Doan T. M. Parkinson C. L. 2004 Data partitions and complex models in Bayesian analysis: the phylogeny of Gymnophthalmid lizards. Systematic Biology 53 448 469

Caterino M. S. Sperling F. A. H. 1999 Papilio phylogeny based on mitochondrial cytochrome oxidase I and II genes. Molecular Phylogenetics and Evolution 11 122 137 DOI

Caterino M. S. Cho S. Sperling F. A. H. 2000 The current state of insect molecular systematics: a thriving tower of babel. Annual Review of Entomology 45 1 54 DOI

Caterino M. S. Reed R. D. Kuo M. M. Sperling F. A. H. 2001 A partitioned likelihood analysis of swallowtail butterfly phylogeny (Lepidoptera: Papilionidae). Systematic Biology 50 106 127 DOI

Cho S. Mitchell A. Regier J. C. Mitter C. Poole R. W. Friedlander T. P. Zhao S. 1995 A highly conserved nuclear gene for low-level phylogenetics: Elongation Factor-1α recovers morphology-based tree for heliothine moths. Molecular Biology and Evolution 12 650 656

Clary D. O. Wolstenholme D. R. 1985 The mitochondrial DNA molecule of Drosophila yakuba: nucleotide sequence, gene organization, and genetic code. Journal of Molecular Evolution 22 252 271

Common I. F. B. (1990). ‘Moths of Australia.’ (Melbourne University Press: Melbourne, Australia.)

Cox C. B. , and Moore P. D. (2000). ‘Biogeography. An Ecological and Evolutionary Approach.’ 6th edn. (Blackwell Science: Oxford, UK.)

Cracraft J. 2001 Avian evolution, Gondwana biogeography and the Cretaceous–Tertiary mass extinction event. Proceedings of the Royal Society of London. Series B. Biological Sciences 268 459 469

Crane P. R. Friis E. M. Pedersen K. R. 1995 The origin and early diversification of angiosperms. Nature 374 27 33 DOI

Cranston P. S. , and Naumann I. D. (1991). Biogeography. In ‘The Insects of Australia’. (Ed I. D. Naumann.) pp. 180–197. (Melbourne University Press: Melbourne, Australia.)

D’Abrera B. (1981). ‘Butterflies of the Neotropical Region. Part 1 Papilionidae & Pieridae.’ (Lansdowne Editions: Melbourne, Australia.)

Danforth B. N. Shuqing J. 1998 Elongation Factor-1α occurs as two copies in bees: implications for phylogenetic analysis of EF-1α sequences in insects. Molecular Biology and Evolution 15 225 235

de Jong R. (2001). Faunal exchange between Asia and Australia in the Tertiary as evidenced by recent butterflies. In ‘Faunal and Floral Migrations and Evolution in SE Asia-Australia’. (Eds I. Metcalfe, J. M. B. Smith, M. Morwood and I. Davidson.) pp. 133–146. (A.A. Balkema Publishers: Lisse, The Netherlands.)

de Jong R. 2003 Are there butterflies with Gondwanan ancestry in the Australian region? Invertebrate Systematics 17 143 156

de Jong R. Vane-Wright R. I. Ackery P. R. 1996 The higher classification of butterflies (Lepidoptera): problems and prospects. Entomologica Scandinavica 27 65 101

Dettmann M. E. (1994). Cretaceous vegetation: the microfossil record. In ‘History of the Australian Vegetation: Cretaceous to Recent’. (Ed R. S. Hill.) pp. 143–170. (Cambridge University Press: Cambridge, UK.)

Dixey F. A. 1894 On the phylogeny of the Pierinae, as illustrated by their wing-markings and geographical distribution. Transactions of the Entomological Society of London Part 2 249 334

Durden C. J. Rose H. 1978 Butterflies from the middle Eocene: the earliest occurrence of fossil Papilionoidea (Lepidoptera). Texas Memorial Museum, The Pearce-Sellards Series 29 1 25

Edwards E. D. , Gentili P. , Horak M. , Kristensen N. P. , and Nielsen E. S. (1999). The cossoid/sesoid assemblage. In ‘Lepidoptera, Moths and Butterflies. Volume 1. Evolution, Systematics and Biogeography’. (Ed. N. P. Kristensen.) pp. 423–462. (de Gruyter: Berlin, Germany.)

Eliot J. N. 1973 The higher classification of the Lycaenidae (Lepidoptera): a tentative arrangement. Bulletin of the British Museum (Natural History). Entomology 28 371 505

Emmel T. C. , Minno M. C. , and Drummond B. A. (1992). ‘Florissant Butterflies. A Guide to the Fossil and Present day Species of Central Colorado.’ (Stanford University Press: Stanford.)

Ericson P. G. P. Christidis L. Cooper A. Irestedt M. Jackson J. Johansson U. S. Norman J. A. 2002 A Gondwanan origin of passerine birds supported by DNA sequences of the endemic New Zealand wrens. Proceedings of the Royal Society of London. Series B. Biological Sciences 269 235 241

Faith D. P. 1991 Cladistic permutation tests for monophyly and nonmonophyly. Systematic Zoology 40 366 375

Farris J. S. Källersjö M. Kluge A. C. Bult C. 1994 Testing significance of incongruence. Cladistics 10 315 319

Feeny P. P. (1991). Chemical constraints on the evolution of swallowtail butterflies. In ‘Plant-animal Interactions: Evolutionary Ecology in Tropical and Temperate Regions’. (Eds P. W. Price, T. M. Lewinsohn, W. W. Benson and G. W. Fernandes.) pp. 315–340. (Wiley: New York, USA.)

Felsenstein J. 1985 Confidence limits on phylogenies: an approach using the boot strap. Evolution; International Journal of Organic Evolution 39 783 791

Felsenstein J. 1988 Phylogenies from molecular sequences: inference and realiability. Annual Review of Genetics 22 521 565

Felsenstein J. (2004). ‘Inferring Phylogenies.’ (Sinauer Associates Inc.: Sunderland, MA, USA.)

Fiedler K. 1991 Systematic, evolutionary, and ecological implications of myrmecophily within the Lycaenidae (Insecta: Lepidoptera: Papilionoidea). Bonner Zoologische Monographien 31 1 197

Folmer O. Black M. Hoeh W. Lutz R. Vrijenhoek R. 1994 DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology 3 294 299

Ford E. B. 1944 Studies on the chemistry of pigments in the Lepidoptera, with reference to their bearing on systematics. 4. The classification of the Papilionidae. Transactions of the Royal Entomological Society of London 94 201 223

Fordyce J. A. Nice C. C. 2003 Contemporary patterns in a historical context: phylogeographic history of the Pipevine Swallowtail, Battus philenor (Papilionidae). Evolution; International Journal of Organic Evolution 57 1089 1099

Friedlander T. P. Regier J. C. Mitter C. 1994 Phylogenetic information content of five nuclear gene sequences in animals: initial assessment of character sets from concordance and divergence studies. Systematic Biology 43 511 525

Friedlander T. P. Horst K. R. Regier J. C. Mitter C. Peigler R. S. Fang Q. Q. 1998 Two nuclear genes yield concordant relationships within Attacini (Lepidoptera: Saturnidae). Molecular Phylogenetics and Evolution 9 131 140

Gaunt M. W. Miles M. A. 2002 An insect molecular clock dates the origin of the insects and accords palaeontological and biogeographic landmarks. Molecular Biology and Evolution 19 748 761

Grimaldi D. 1999 The co-radiations of pollinating insects and angiosperms in the Cretaceous. Annals of the Missouri Botanical Garden 86 373 406

Grote A. R. 1900 The descent of the pierids. Proceedings of the American Philosophical Society 39 4 67

Hall J. P. W. Robbins R. K. Harvey D. J. 2004 Extinction and biogeography in the Caribbean: new evidence from a fossil riodinid butterfly in Dominican amber. Proceedings of the Royal Society of London. Series B. Biological Sciences 271 797 801

Hall R. (1998). The plate tectonics of Cenozoic SE Asia and the distribution of land and sea. In ‘Biogeography and Geological Evolution of SE Asia’. (Eds R. Hall and J. D. Holloway.) pp. 99–131. (Backhuys Publishers: Leiden, The Netherlands.)

Hall R. (2001). Cenozoic reconstructions of SE Asia and the SW Pacific: changing patterns of land and sea. In ‘Faunal and Floral Migrations and Evolution in SE Asia-Australia’. (Eds I. Metcalfe, J. M. B. Smith, M. Morwood and I. Davidson.) pp. 35–56. (A.A. Balkema Publishers: Lisse.)

Hancock D. L. 1980 The status of the genera Atrophaneura Reakirt and Pachliopta Reakirt (Lepidoptera: Papilionidae). Australian Entomological Magazine 7 27 32

Hancock D. L. 1983 Classification of the Papilionidae (Lepidoptera): a phylogenetic approach. Smithersia 2 1 48

Hancock D. L. 1988 A revised classification of the genus Atrophaneura Reakirt (Lepidoptera: Papilionidae). Australian Entomological Magazine 15 7 16

Hancock D. L. 1991 Notes on the phylogeny and biogeography of Ornithoptera Boisduval (Lepidoptera: Papilionidae). Tyô to Ga 42 17 36

Hancock D. L. Orr A. G. 1997 Ornithoptera euphorion (Gray) (Lepidoptera: Papilionidae): species or subspecies? Australian Entomologist 24 165 168

Haugum J. , and Low A. M. (1978–1979). ‘A Monograph of the Birdwing Butterflies. Vol. 1 (1–3).’ (Scandinavian Science Press: Klampenborg, Denmark.)

Hay W. W. , DeConto R. M. , Wold C. N. , Willson K. M. , Voigt S. , et al. (1999). An alternative global Cretaceous paleogeography. In ‘Evolution of the Cretaceous Ocean-climate System. Special Paper 332’. (Eds E. Barrera and C. Johnson.) pp. 1–48. (Geological Society of America: Boulder, CO, USA.)

Heatwole H. (1987). Major components and distributions of the terrestrial fauna. In ‘Fauna of Australia. Volume 1A, General Articles’. (Eds G. R. Dyne and D. W. Walton.) pp. 101–135. (Australian Government Publishing Service: Canberra, Australia.)

Hedges S. B. Kumar S. 2003 Genomic clocks and evolutionary timescales. Trends in Genetics 19 200 206

Hill R. S. (1994). The Australian fossil plant record: an introduction. In ‘History of the Australian Vegetation: Cretaceous to Recent’. (Ed R. S. Hill.) pp. 1–4. (Cambridge University Press: Cambridge, UK.)

Hill R. S. , Truswell E. M. , McLoughlin S. , and Dettman M. E. (1999). Evolution of the Australian flora: fossil evidence. In ‘Flora of Australia. Volume 1. Introduction’. (Eds A. E. Orchard and H. S. Thompson.) pp. 251–320. (Australian Biological Resources Study: Canberra, Australia.)

Hillis D. M. , Mable B. K. , Larson A. , Davis S. K. , and Zimmer E. A. (1996). Nucleic Acids IV. Sequencing and cloning. In ‘Molecular Systematics’. (Eds D. M. Hillis, C. Moritz and B. K. Mable.) pp. 321–384. (Sinauer Associates: Sunderland, MA, USA.)

Holloway J. D. 1969 A numerical investigation of the biogeography of the butterfly fauna of India, and its relation to continental drift. Biological Journal of the Linnean Society 1 373 385

Holloway J. D. 1973 The affinities within four butterfly groups (Lepidoptera: Rhopalocera) in relation to general patterns of butterfly distribution in the Indo-Australian area. Transactions of the Royal Entomological Society of London 125 125 176

Holloway J. D. (1974 a). The biogeography of Indian butterflies. In ‘Ecology and Biogeography in India. Monographiae Biologicae. Volume 23’. (Ed M. S. Mani.) pp. 473–499. (Dr. W. Junk bv Publishers: The Hague, The Netherlands.)

Holloway J. D. 1974 b The endemic Satyridae (Lepidoptera: Rhopalocera) of New Caledonia. Journal of Entomology (B) 43 89 101

Holloway J. D. (1979). ‘A Survey of the Lepidoptera, Biogeography and Ecology of New Caledonia.’ (Dr. W. Junk B.V. Publishers: The Hague, The Netherlands.)

Holloway J. D. (1986). Origins of Lepidopteran faunas in high mountains of the Indo-Australian tropics. In ‘High Altitude Tropical Biogeography’. (Eds F. Vuilleumier and M. Monasterio.) pp. 533–556. (Oxford University Press: New York, USA.)

Holloway J. D. , and Hall R. (1998). SE Asian geology and biogeography: an introduction. In ‘Biogeography and Geological Evolution of SE Asia’. (Eds R. Hall and J. D. Holloway.) pp. 1–23. (Backhuys Publishers: Leiden, The Netherlands.)

Holloway J. D. Jardine N. 1968 Two approaches to zoogeography: a study based on the distributions of butterflies, birds and bats in teh Indo-Australian area. Proceedings of the Linnean Society of London 179 153 188

Holloway J. D. , and Nielsen E. S. (1999). Biogeography of the Lepidoptera. In ‘Lepidoptera, Moths and Butterflies. Volume 1. Evolution, Systematics and Biogeography’. (Ed N. P. Kristensen.) pp. 423–462. (de Gruyter: Berlin, Germany.)

Huelsenbeck J. P. 1998 Systematic bias in phylogenetic analysis: is the Strepsiptera problem solved? Systematic Biology 47 519 537

Janz N. Nylin S. 1998 Butterflies and plants: a phylogenetic study. Evolution 52 486 502

Jell P. A. , and Duncan P. M. (1986). Invertebrates, mainly insects, from the freshwater Lower Cretaceous, Koonwarra Fossil Bed (Korumburra Group), South Gippsland, Victoria. In ‘Plants and Invertebrates from the Lower Cretaceous Koonwarra Fossil Bed, South Gippsland, Victoria’. (Eds P. A. Jell and J. Roberts.) pp. 111–205. (Association of Australasian Palaeontologists: Sydney, Australia.)

Johnson K. , and Coates S. (1999). ‘Nabokov’s Blues. The Scientific Odyssey of a Literary Genius.’ (Zoland Books: Cambridge, MS, USA.)

Kamie K. Taira H. Ooura H. Kakuta A. Matsumoto S. Ejiri S.-i. Katsumata T. 1993 Nucleotide sequence of the cDNA encoding silk gland elongation factor 1α. Nucleic Acids Research 21 742

Keto A. , Scott K. , and Fox A. (1985). An overview. In ‘Rainforests of Australia’. (Ed. P. Figgis.) pp. 17–32. (Weldons: Sydney, Australia.)

Kitching R. L. , and Dunn K. L. (1999). The biogeography of Australian butterflies. In ‘Biology of Australian Butterflies. Monographs of Australian Lepidoptera. Volume 6’. (Eds R. L. Kitching, R. E. Jones, E. Scheermeyer and N. E. Pierce.) pp. 53–74. (CSIRO Publishing: Melbourne, Australia.)

Kitching R. L. , Eastwood R. G. , and Hurley K. (2001). Butterflies and Wallace’s Line: faunistic patterns and explanatory hypotheses within the south-east Asian butterflies. In ‘Faunal and Floral Migrations and Evolution in SE Asia-Australasia’. (Eds I. Metcalfe, C. R. Smith, M. Morwood and I. Davidson.) pp. 269–286. (A.A. Balkema Publishers: Lisse, The Netherlands.)

Klitzke C. F. Brown K. S. 2000 The occurrence of aristolochic acids in Neotropical troidine swallowtails (Lepidoptera: Papilionidae). Chemoecology 10 99 102

Klots A. B. 1933 A generic classification of the Pieridae (Lepidoptera) together with a study of the male genitalia. Entomologica America 12 139 242

Kondo K. Shinkawa T. Matsuka H. 2003 Molecular systematics of birdwing butterflies (Papilionidae) inferred from mitochondrial ND5 gene. Journal of the Lepidopterists’ Society 57 17 24

Krause D. W. , Hartman J. H. , and Wells N. A. (1997). Late Cretaceous vertebrates from Madagascar. Implications for biotic changes in deep time. In ‘Natural Change and Human Impact in Madagascar’. (Eds S. M. Goodman and B. D. Patterson.) pp. 3–43. (Smithsonian Institution Press: Washington, DC, USA.)

Krause D. W. Rogers R. R. Forster C. A. Hartman J. H. Buckley G. A. Sampson S. D. 1999 The Late Cretaceous vertebrate fauna of Madagascar: implications for Gondwanan Paleobiogeography. GSA Today 9 1 7

Kristensen N. P. 1976 Remarks on the family-level phylogeny of butterflies (Insecta, Lepidoptera, Rhopalocera). Zeitschrift für zoologische Systematik und Evolutionsforschung 14 25 33

Kristensen N. P. , and Skalski A. W. (1999). Phylogeny and Palaeontology. In ‘Lepidoptera, Moths and Butterflies. Volume 1: Evolution, Systematics and Biogeography’. (Ed N. P. Kristensen.) pp. 7–25. (de Gruyter: Berlin, Germany.)

Kukalová-Peck J. (1991). Fossil history and the evolution of hexapod structures. In ‘The Insects of Australia’. (Ed. I. D. Naumann.) pp. 141–179. (Melbourne University Press: Melbourne, Australia.)

Labandeira C. C. Dilcher D. L. Davis D. R. Wagner D. L. 1994 Ninety-seven million years of angiosperm-insect association: paleobiological insights into the meaning of coevolution. Proceedings of the National Acadamy of Sciences, USA 91 12278 12282

Ladiges P. Y. Udovicic F. Nelson G. 2003 Australian biogeographical connections and the phylogeny of large genera in the plant family Myrtaceae. Journal of Biogeography 30 989 998

Lanave C. Preparata G. Saccone C. Serio G. 1984 A new method for calculating evolutionary substitution rates. Journal of Molecular Evolution 20 86 93

Lattke J. E. 2003 Biogeographic analysis of the ant genus Gnamptogenys Roger in South-east Asia-Australasia (Hymenoptera: Formicidae: Ponerinae). Journal of Natural History 37 1879 1897

Lee C. Grasso C. Sharlow M. F. 2002 Multiple sequence alignment using partial order graphs. Bioinformatics (Oxford, England) 18 452 464 DOI

Lees D. C. Smith N. G. 1991 Foodplant associations of the Uraniinae (Uraniidae) and their systematic, evolutionary, and ecological significance. Journal of the Lepidopterists’ Society 45 296 347

Lidgard S. Crane P. R. 1988 Quantitative analysis of the early angiosperm radiation. Nature 331 344 346

Maddison W. P. , and Maddison D. R. (1999). ‘MacClade: Analysis of Phylogeny and Character Evolution, version 3.08a for Macintosh.’ (Sinauer Associates: Sunderland, MA, USA.)

Magallón S. A. 2004 Dating lineages: molecular and paleontological approaches to the temporal framework of clades. International Journal of Plant Sciences 165 S7 S21 DOI

Martins Neto R. G. Kucera-Santos J. C. de Moraes Vieira F. R. de Campos Fragoso L. M. 1993 Nova espécie de borboleta (Lepidoptera: Nymphalidae: Satirynae [sic]) da Formçao Tremembé, Oligoceno do Estado de Sao Paulo. Acta Geologica Leopoldensia 16 5 16

Miller J. S. 1987 a Host-plant relationships in the Papilionidae (Lepidoptera): parallel cladogenesis or colonization? Cladistics 3 105 120

Miller J. S. 1987 b Phylogenetic studies in the Papilioninae. Bulletin of the American Museum of Natural History 186 365 512

Miller J. Y. Brown F. M. 1989 A new oligocene fossil butterfly Vanessa amerindica (Lepidoptera: Nymphalidae) from the Florissant formation, Colorado. Bulletin of the Allyn Museum No. 126 1 9

Miller L. D. (1968). The higher classification, phylogeny and zoogeography of the Satyridae (Lepidoptera). Memoirs of the American Entomological Society 24, iv+174 pp.

Miller L. D. Miller J. Y. 1997 Gondwanan butterflies: the Africa-South America connection. Metamorphosis 3 42 51

Mitchell A. Cho S. Regier J. C. Mitter C. Poole R. W. Matthews M. 1997 Phylogenetic utility of Elongation Factor-1α in Noctuoidea (Insecta: Lepidoptera): the limits of synonymous substitution. Molecular Biology and Evolution 14 381 390

Mitchell A. Mitter C. Regier J. C. 2000 More taxa or more characters revisted: combining data from nuclear protein-encoding genes for phylogenetic analysis of Noctuoidea (Insecta: Lepidoptera). Systematic Biology 49 202 224

Monteiro A. Pierce N. E. 2001 Phylogeny of Bicyclus (Lepidoptera: Nymphalidae) inferred from COI, COII, and EF-1a gene sequences. Molecular Phylogenetics and Evolution 18 264 281 DOI

Morinaka S. Nakazawa T. 1999 A study of the Delias eichhorni-complex from New Guinea Island (Lepidoptera; Pieridae) (IV) phylogenetic estimation using morphological characters. Biogeography 1 69 80

Morinaka S. Maeyama T. Maekawa K. Erniwati D. Prijono S. N. Ginarsa I. K. Nakazawa T. Hidaka T. 1999 Molecular phylogeny of birdwing butterflies based on the representatives in most genera of the tribe Troidini (Lepidoptera: Papilionidae). Entomological Science 2 347 358

Morinaka S. Minaka N. Sekiguchi M. Erniwati D. Prijono S. N. Ginarsa I. K. Miyata T. Hidaka T. 2000 Molecular phylogeny of birdwing butterflies of the tribe Troidini (Lepidoptera: Papilionidae) – using all species of the genus Ornithoptera. Biogeography 2 103 111

Morrone J. J. Crisci J. V. 1995 Historical biogeography: introduction to methods. Annual Review of Ecology and Systematics 26 373 401

Munroe E. 1961 The classification of the Papilionidae (Lepidoptera). The Canadian Entomologist 1 51 Supplement 17

Munroe E. Ehrlich P. R. 1960 Harmonization of concepts of higher classification of the Papilionidae. Journal of the Lepidopterists’ Society 14 169 175

Nelson G. , and Platnick N. I. (1981). ‘Systematics and Biogeography: Cladistics and Vicariance.’ (Columbia University Press: New York.)

Orr A. G. (1999). The Big Greasy, Cressida cressida (Papilionidae). In ‘Biology of Australian Butterflies. Monographs of Australian Lepidoptera. Volume 6’. (Eds R. L. Kitching, R. E. Jones, E. Scheermeyer and N. E. Pierce.) pp. 115–134. (CSIRO Publishing: Melbourne, Australia.)

Palumbi S. R. (1996). Nucleic acids II: the polymerase chain reaction. In ‘Molecular Systematics’. (Eds D. M. Hillis, C. Moritz and B. K. Mable.) pp. 205–247. (Sinauer Associates: Sunderland, MA, USA.)

Parsons M. J. 1996 a Gondwanan evolution of the Troidine Swallowtails (Lepidoptera: Papilionidae): cladistic reappraisals using mainly immature stage characters, with focus on the Birdwings Ornithoptera Boisduval. Bulletin of the Kitakyushu Museum of Natural History 15 43 118

Parsons M. J. 1996 b New species of Aristolochia and Pararistolochia (Aristolochiaceae) from Australia and New Guinea. Botanical Journal of the Linnean Society 120 199 238

Parsons M. J. 1996 c A phylogenetic reappraisal of the birdwing genus Ornithoptera (Lepidoptera: Papilionidae: Troidini) and a new theory of its evolution in relation to Gondwanan vicariance biogeography. Journal of Natural History 30 1707 1736

Parsons M. J. (1998). ‘The Butterflies of Papua New Guinea. Their Systematics and Biology.’ (Academic Press: London, UK.)

Penz C. M. Peggie D. 2003 Phylogenetic relationships among Heliconiinae genera based on morphology (Lepidoptera: Nymphalidae). Systematic Entomology 28 451 479

Pole M. 1994 The New Zealand flora - Entirely long-distance dispersal? Journal of Biogeography 21 625 635

Posada D. Crandall K. A. 1998 MODELTEST: testing the model of DNA substitution. Bioinformatics (Oxford, England) 14 817 818

Rabinowitz P. D. Coffin M. F. Falvey D. 1983 The separation of Madagascar and Africa. Science 220 67 69

Raxworthy C. J. Forstner M. R. J. Nussbaum R. A. 2002 Chameleon radiation by oceanic dispersal. Nature 415 784 787

Reed R. D. Sperling F. A. H. 1999 Interaction of process partitions in phylogenetic analysis: an example from the swallowtail butterfly genus Papilio. Molecular Biology and Evolution 16 286 297

Reid C. A. M. 1992 Notes on the concept of “Original (or ‘primitive’) state = most abundant” in biogeography and karyology. Chrysomela 27 5

Riek E. F. (1971). Origin of the Australian insect fauna. In ‘Second Gondwana Symposium. Proceedings and Papers’. pp. 593–598. (Council for Scientific and Industrial Research, Scientia: Pretoria, South Africa.)

Riek E. F. 1976 A new collection of insects from the Upper Triassic of South Africa. Annals of the Natal Museum 22 791 820

Robbins R. K. 1982 How many butterfly species? News of the Lepidopterists’ Society 3 40 41

Rodríguez F. Oliver J. L. Marín A. Medina J. R. 1990 The general stochastic model of nucleotide substitution. Journal of Theoretical Biology 142 485 501

Roger A. J. Sandblom O. Doolittle W. F. Philippe H. 1999 An evaluation of Elongation Factor 1α as a phylogenetic marker for Eukaryotes. Molecular Biology and Evolution 16 218 233

Ronquist F. 1997 Dispersal–vicariance analysis: a new approach to quantification of historical biogeography. Systematic Biology 46 195 203

Ronquist F. Huelsenbeck J. P. 2003 MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics (Oxford, England) 19 1572 1574

Rothschild M. (1972). Secondary plant substances and warning colouration in insects. In ‘Insect Plant Relationships. Symposium of the Royal Entomological Society of London. Number 6’. (Ed H. F. van Emden.) pp. 59–83. (Royal Entomological Society of London: London, UK.)

Rozefelds A. C. 1988 a Insect leaf mines from the Eocene Anglesea locality, Victoria, Australia. Alcheringa 12 1 6

Rozefelds A. C. 1988 b Lepidoptera mines in Pachypteris leaves (Corystospermaceae: Pteridospermophyta) from the Upper Jurassic/Lower Cretaceous Battle Camp Formation, north Queensland. Proceedings of the Royal Society of Queensland 99 77 81

Sampson S. D. Witmer L. M. Forster C. A. Krause D. W. O’Connor P. M. Dodson P. Ravoavy F. 1998 Predatory dinosaur remains from Madagascar: implications for the Cretaceous biogeography of Gondwana. Science 280 1048 1051

Sanderson M. J. 2002 Estimating absolute rates of molecular evolution and divergence times: a penalized likelihood approach. Molecular Biology and Evolution 19 101 109

Sands D. P. A. , and Scott S. (2002). Editorial. In ‘Conservation of Birdwing Butterflies’. (Eds D. P. A. Sands and S. Scott.) pp. 48. (Science Communication and Editorial Services and The Hut Environmental and Community Association Inc.: Brisbane, Australia.)

Sanmartín I. Ronquist F. 2004 Southern hemisphere biogeography inferred by event-based models: plant verses animal patterns. Systematic Biology 53 216 243

Schodde R. 1989 Origins, radiations and sifting in the Australasian biota - changing concepts from new data and old. Australian Systematic Botanical Society Newsletter 60 2 11

Scoble M. J. 1986 The structure and affinities of the Hedyloidea: a new concept of butterflies. Bulletin of the British Museum (Natural History). Entomology 53 251 286

Scoble M. J. 1990 An identification guide to the Hedylidae (Lepidoptera: Hedyloidea). Entomologica Scandinavica 21 121 158

Scoble M. J. (1992). ‘Guía de las mariposas hedílidas de Costa Rica (Lepidoptera: Hedylidae).’ (Instituto Nacional de Biodiversidad, INBiol.: Heredia, Costa Rica.)

Scotese C. R. (2001). ‘Atlas of Earth History.’ (University of Texas: Arlington, TX, USA.)

Scott J. A. 1985 The phylogeny of butterflies (Papilionoidea and Hesperioidea). The Journal of Research on the Lepidoptera 23 241 281

Scott J. A. (1986). ‘The Butterflies of North America. A Natural History and Field Guide.’ (Stanford University Press: Stanford, CA, USA.)

Scott J. A. , and Wright D. M. (1990). Butterfly phylogeny and fossils. In ‘Butterflies of Europe. Volume 2’. (Ed. O. Kudrna.) pp. 152–208. (Aula Verlag: Wiesbaden, Germany.)

Scriber J. M. , Tsubaki Y. , and Lederhouse R. C. (1995). ‘Swallowtail Butterflies: Their Ecology and Evolutionary Biology.’ (Scientific Publishers: Gainesville, FL, USA.)

Sequeira A. S. Farrell B. D. 2001 Evolutionary origins of Gondwanan interactions: how old are Araucaria beetle herbivores? Biological Journal of the Linnean Society 74 459 474

Shapiro A. M. 1994 Why are there so few butteflies in the high Andes? The Journal of Research on the Lepidoptera 31 25 56

Shapiro A. M. (1996). Foreword. In ‘Las Mariposas de Chile’. (Eds L. E. Peña and A. J. Ugarté.) pp. 12–15. (Editorial Universitaria: Santiago, Chile.)

Shields O. 1976 Fossil butterflies and the evolution of Lepidoptera. The Journal of Research on the Lepidoptera 15 132 143

Shields O. 1985 Zoogeography of the Libytheidae (Snouts or Beaks). Tokurana 9 1 58

Shields O. 1988 Mesozoic history and neontology of Lepidoptera in relation to Trichoptera, Mecoptera, and angiosperms. Journal of Paleontology 62 251 258

Shields O. 1989 World numbers of butterflies. Journal of the Lepidopterists’ Society 43 178 183

Shields O. Dvorak S. K. 1979 Butterfly distribution and continental drift between the Americas, the Caribbean and Africa. Journal of Natural History 13 221 250

Silva-Brandão K. L. Lucci Freitas A. V. Brower A. V. Z. Solferini V. N. 2005 Phylogenetic relationships of the New World Troidini swallowtails (Lepidoptera: Papilionidae) based on COI, COII, and EF-1α genes. Molecular Phylogenetics and Evolution
in press

Sime K. R. Feeny P. P. Haribal M. M. 2000 Sequestration of aristolochic acids by the pipevine swallowtail, Battus philenor (L.): evidence and ecological implications. Chemoecology 10 169 178

Simon C. Frati F. Beckenbach A. Crespi B. J. Liu H. Flook P. 1994 Evolution, weighting, and phylogenetic utility of mitochondrial gene sequences and a compilation of conserved polymerase chain reaction primers. Annals of the Entomological Society of America 87 651 701

Simmons M. P. Pickett K. M. Miya M. 2004 How Meaningful Are Bayesian Support Values? Molecular Biology and Evolution 21 188 199

Simpson G. G. 1977 Too many lines: the limits of the Oriental and Australian zoogeographic regions. Proceedings of the American Philosophical Society 121 107 120

Skalski A. W. 1990 An annotated review of fossil records of lower Lepidoptera. Bulletin of the Sugadeira Montane Research Centre 11 125 128
Tsukuba University

Smith A. G. , Smith D. G. , and Funnell B. M. (1994). ‘Atlas of Mesozoic and Cenozoic Coastlines.’ (Cambridge University Press: Cambridge, UK.)

Sorenson M. D. (1999). ‘TreeRot, version 2.’ (Boston University: Boston, MA, USA.)

Su Z. H. Tominaga O. Okamoto M. Osawa S. 1998 Origin and diversification of hindwingless Damaster ground beetles within the Japanese islands as deduced from mitochondrial ND5 gene sequences (Coleoptera, Carabidae). Molecular Biology and Evolution 15 1026 1039

Sullivan J. Swofford D. L. 1997 Are guinea pigs rodents? The importance of adequate models in molecular phylogenetics. Journal of Mammalian Evolution 4 77 86

Suzuki Y. Glazko G. V. Nei M. 2002 Overcredibility of molecular phylogenies obtained by Bayesian phylogenetics. Proceedings of the National Acadamy of Sciences 99 16138 16143 DOI

Swofford D. L. (2002). ‘PAUP*: Phylogenetic Analysis Using Parsimony (* and Other Methods), version 4.0b10 for Macintosh.’ (Sinauer Associates: Sunderland, MA, USA.)

Talbot G. (1928–1937). ‘A Monograph of the Pierine Genus Delias. Parts I–VI.’ (British Museum (Natural History): London, UK.)

The Angiosperm Phylogeny Group* 2003 An update of the Angiosperm Phylogeny Group: classification for the orders and families of flowering plants: APG II. Botanical Journal of the Linnean Society 141 399 436 DOI

Tindale N. B. 1980 Origin of the Lepidoptera, with description of a new mid-Triassic species and notes on the origin of the butterfly stem. Journal of the Lepidopterists’ Society 34 263 285

Tindale N. B. (1981). The origin of the Lepidoptera relative to Australia. In ‘Ecological Biogeography of Australia’. (Ed A. Keast.) pp. 957–976. (Dr. W. Junk bv Publishers: The Hague, The Netherlands.)

Torres E. Lees D. C. Vane-Wright R. I. Kremen C. Leonard J. A. Wayne R. K. 2001 Examining monophyly in a large radiation of Madagascan butterflies (Lepidoptera: Satyrinae: Mycalesina) based on mitochondrial DNA data. Molecular Phylogenetics and Evolution 20 460 473

Trueman J. W. H. 1996 Permutation tests and outgroups. Cladistics 12 253 261 DOI

Truswell E. M. , Kershaw A. P. , and Sluiter I. R. (1987). The Australian-Southeast Asian connection; evidence from the palaeobotanical record. In ‘Biogeographical Evolution of the Malay Archipelago’. (Ed T. C. Whitmore.) pp. 32–49. (Clarendon Press: Oxford, UK.)

Tyler H. A. , Brown K. S. Jr , and Wilson K. H. (1994). ‘Swallowtail Butterflies of the Americas. A Study in Biological Dynamics, Ecological Diversity, Biosystematics, and Conservation.’ (Scientific Publishers: Gainesville, FL, USA.)

Vane-Wright R. I. (2003). Evidence and identity in butterfly systematics. In ‘Butterflies: Ecology and Evolution Taking Flight’. (Eds C. L. Boggs, W. B. Watt and P. R. Ehrlich.) pp. 477–513. (University of Chicago Press: Chicago, IL, USA.)

Vane-Wright R. I. 2004 Butterflies at that awkward age. Nature 428 477 479 DOI

Vences M. Freyhof J. Sonnenberg R. Kosuch J. Veith M. 2001 Reconciling fossils and molecules: Cenozoic divergence of cichlid fishes and the biogeography of Madagascar. Journal of Biogeography 28 1091 1099 DOI

Vickers-Rich P. , Monaghan J. M. , Baird R. F. , and Rich T. H. (1991). ‘Vertebrate Palaeontology of Australasia.’ (Pioneer Design Studio: Melbourne, Australia.)

Viloria A. L. (2003). Historical biogeography and the origins of the satyrine butterflies of the Tropical Andes (Insecta: Lepidoptera, Rhopalocera). In ‘Una perspectiva latinoamericana de la biogeografía México’. (Eds J. J. Morrone and J. Llorente-Bousquets.) pp. 247–261. (D. F. Las Prensas de Ciencias, Facultad de Ciencias, UNAM: Mexico City, Mexico.)

Wahlberg N. Weingartner E. Nylin S. 2003 Towards a better understanding of the higher systematics of Nymphalidae (Lepidoptera: Papilionoidea). Molecular Phylogenetics and Evolution 28 473 484 DOI

Wahlberg N. Braby M. F. Brower A. Z. de Jong R. Lee M.-M. Nylin S. Pierce N. E. Sperling F. A. H. Vila R. U. Warren A. D. Zakharov E. 2005 Synergistic effects of combining morphological and molecular data in resolving the phylogeny of butterflies and skippers. Proceedings of the Royal Society of London B in press

Weintraub J. D. (1995). Host plant association patterns and phylogeny in the tribe Troidini (Lepidoptera: Papilionidae). In ‘Swallowtail Butterflies: Their Ecology and Evolutionary Biology’. (Eds J. M. Scriber, Y. Tsubaki and R. C. Lederhouse.) pp. 307–316. (Scientific Publishers: Gainesville, FL, USA.)

Whalley P. 1986 A review of the current fossil evidence of Lepidoptera in the Mesozoic. Biological Journal of the Linnean Society 28 253 271

White M. E. (1994). ‘After the Greening. The Browning of Australia.’ (Kangaroo Press: Sydney, Australia.)

White M. E. (1997). ‘Listen...Our Land is Crying. Australia’s Environment: Problems and Solutions.’ (Kangaroo Press: Sydney, Australia.)

White M. E. (1998). ‘The Greening of Gondwana.’ 3rd edn. (Kangaroo Press: Sydney, Australia.)

Wiegmann B. M. Mitter C. Regier J. C. Friedlander T. P. Wagner D. L. Nielsen E. S. 2000 Nuclear genes resolve Mesozoic-aged divergences in the insect order Lepidoptera. Molecular Phylogenetics and Evolution 15 242 259

Wikström N. Savolainen V. Chase M. W. 2001 Evolution of the angiosperms: calibrating the family tree. Proceedings of the Royal Society of London. Series B. Biological Sciences 268 2211 2220 DOI

Yagi T. Sasaki G. Takebe H. 1999 Phylogeny of Japanese papilionid butterflies inferred from nucleotide sequences of the mitochondrial ND5 gene. Journal of Molecular Evolution 48 42 48

Zakharov E. V. Caterino M. S. Sperling F. A. H. 2004 Molecular phylogeny, historical biogeography, and divergence time estimates for swallowtail butterflies of the genus Papilio (Lepidoptera: Papilionidae). Systematic Biology 53 193 215

Zeuner F. E. 1943 Studies in the systematics of Troides Hübner (Lepidoptera: Papilionidae) and its allies; distribution and phylogeny in relation to the geological history of the Australasian Archipelago. Transactions of the Zoological Society of London 25 107 184

Zeuner F. E. (1962). Notes on the evolution of the Rhopalocera (Lep.). 11th International Congress of Entomology 1, 310–313.

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