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Revision of Pazala Moore, 1888: The Graphium (Pazala) mandarinus (Oberthür, 1879) Group, with Treatments of Known Taxa and Descriptions of New Species and New Subspecies (Lepidoptera: Papilionidae)

Hu, Shao-Ji; Cotton, Adam M.; Condamine, Fabien L.; Duan, Kuang; Wang, Rong-Jiang; Hsu, Yu-Feng; Zhang, Xin; Cao, Jun

Hu, Shao-Ji, Cotton, Adam M., Condamine, Fabien L., Duan, Kuang, Wang, Rong-Jiang, Hsu, Yu-Feng, Zhang, Xin, Cao, Jun (2018): Revision of Pazala Moore, 1888: The Graphium (Pazala) mandarinus (Oberthür, 1879) Group, with Treatments of Known Taxa and Descriptions of New Species and New Subspecies (Lepidoptera: Papilionidae). Zootaxa 4441 (3): 401-446, DOI:
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  • Alpheraky, S. (1897) Lepidopteres des provinces chinoises Se-Tchouen et Kham recueillis, en 1893, par M-r G. N. Potanine. In: Romanoff, N.M. (Ed.), Memoires sur les Lepidopteres. Vol. 9. P.P. Soikine, St.-Petersbourg, pp. 83-149.

  • Ayres, D.L., Darling, A., Zwickl, D.J., Beerli, P., Holder, M.T., Lewis, P.O., Huelsenbeck, J.P., Ronquist, F., Swofford, D.L., Cummings, M.P., Rambaut, A. & Suchard, M.A. (2012) BEAGLE: an application programming interface and highperformance computing library for statistical phylogenetics. Systematic Biology, 61, 170-173.


  • Baker, D.B. (1995) A review of the Asian species of the genus Euaspis Gerstacker (Hymenoptera: Apoidea: Megachilidae). Zoologische Mededelingen, Leiden, 69, 281-302.

  • Bertheau, C., Schuler, H., Krumbock, S., Arthofer, W. & Stauffer, C. (2011) Hit or miss in phylogenetic analyses: the case of the cryptic NUMTs. Molecular Ecology Resources, 11, 1056-1059.


  • Borkhausen, M.B. (1788) Naturgeschichte der Europaischen Schmetterlinge nach Systematischer Ordnung. 1. Varrentrapp und Wenner, Frankfurt, 36 + 288 pp.

  • Bozano, G.C. (2008) Guide to the Butterflies of the Palearctic Region, Nymphalidae part III, Subfamily Limenitidinae, Tribe Neptini. Omnes Artes, Milano, 77 pp.

  • Bryk, F. (1946) Type Lycaena ferra for Satsuma Murray nec. Adams (trans.). Arkiv for Zoologi, 38, 50.

  • Chou, I. (1994) Monographia Rhopalocerorum Sinensium. Henan Science and Technology Publishing House, Zhengzhou, 9 + 852 pp.

  • Condamine, F.L., Sperling, F.A.H., Wahlberg, N., Rasplus, J.-Y. & Kergoat, G.J. (2012) What causes latitudinal gradients in species diversity? Evolutionary processes and ecological constraints on swallowtail biodiversity. Ecology Letters, 15, 267- 277.


  • Condamine, F.L., Sperling, F.A.H. & Kergoat, G.J. (2013a) Global biogeographical pattern of swallowtail diversification demonstrates alternative colonization routes in the Northern and Southern hemispheres. Journal of Biogeography, 40, 9- 23.


  • Condamine, F.L., Toussaint, E.F.A., Cotton, A.M., Genson, G.S., Sperling, F.A.H. & Kergoat, G.J. (2013b) Fine-scale biogeographical and temporal diversification processes of peacock swallowtails (Papilio subgenus Achillides) in the Indo- Australian Archipelago. Cladistics, 29, 88-111.


  • Cotton, A.M. & Racheli, T. (2007) A preliminary annotated checklist of the Papilionidae of Laos with notes on taxonomy, phenology, distribution and variation (Lepidoptera, Papilionoidea). Fragmenta Entomologica, 38, 279-378.


  • de Niceville, L. (1886) On some new Indian butterflies. Journal of the Asiatic Society of Bengal, 55, Part II (3), 249-256, pl. XI.

  • de Niceville, L. (1900) On new and little-known Lepidoptera from the Oriental Region. Journal of Bombay Natural History Society, 13 (1), 157-176, pls. CC-EE.

  • Eliot, J.N. (1963) The Heliophorus epicles (Godart, 1823) (Lepidoptera: Lycaenidae) complex. Entomologist, 96, 169-180.

  • Elwes, H.J. & Edwards, J. (1893) A revision of the genus Ypthima, with especial reference to the characters afforded by the male genitalia. Transactions of the Entomological Society of London, 41, 1-54.


  • Fabricius, J.C. (1807) Die neueste gattungs-eintheilung der schmetterlinge aus den Linneichen Gattungen Papilio und Sphinx. Magazin fur Insektenkunde, 6, 277-285.

  • Fisher, K.J. (1948) Some geographical notes on West China localities. Entomologist, 81, 213-219.

  • Folmer, O., Black, M.B., Hoch, W., Lutz, R.A. & Vrijehock, R.C. (1994) DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology, 3, 294-299.

  • Geyer, C. (1832) Zutrage zur Sammlung exotischer Schmettlinge. Vol. 4. im Verlag der Hubner'schen Werke, Augsburg, 48 pp. [in German]

  • Gray, G.R. (1831) Description of eight new species of Indian butterflies, (Papilio, Lin.) from the collection of General Hardwicke. Zoological Miscellany, 1, 32-33.

  • Gray, G.R. [1853] Catalogue of Lepidopterous Insects in the Collection of the British Museum. Part I. Papilionidae. Taylor and Francis, London, iii + 84 pp.

  • Guindon, S. & Gascuel, O. (2003) A simple, fast and accurate method to estimate large phylogenies by maximum-likelihood. Systematic Biology, 53, 696-704.


  • Hajibabaei, M., Singer, G.A.C., Hebert, P.D.N. & Hickey, D.A. (2007) DNA barcoding: how it complements taxonomy, molecular phylogenetics and population genetics. Trends in Genetics, 23, 167-172.


  • Hall, T.A. (1999) BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nuclear Acids Symposium Series, 41, 95-98.

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

  • Hebert, P.D.N., Cywinska, A., Ball, S.L. & de Waard, J.R. (2003) Biological identifications through DNA barcodes. Proceedings of the Royal Society of London, 270, 313-321.


  • Hebert, P.D.N., Penton, E.H., Burns, J.M., Janzen, D.H. & Hallwachs, W. (2004) Ten species in one: DNA barcoding reveals cryptic species in the neotropical skipper butterfly Astraptes fulgerator. Proceedings of the National Academy of Sciences of the United States of America, 101, 14812-14817.


  • Hemming, F. (1934) Revisional notes on certain species of Rhopalocera (Lepidoptera). Stylops, 3, 193-200.


  • Huang, H. (1999) Some new butterflies from China 1 (Rhopalocera). Lambillionea, 94, 642-676.

  • Huang, H. (2002) Some new nymphalids from the valley of Nujiang and Dulongjiang, China (Lepidoptera, Nymphalidae). Atalanta, 33, 339-360.

  • Huang, H. & Sun, W.H. (2016) Ahlbergia bijieensis spec. nov. from Guizhou, China. Atalanta, 47, 151-160.

  • Huang, H., Wang, J.Y. & Chen, Z. (2016) A new species of the Euthalia subgenus Limbusa from the Yarlung Tsangpo Grand Canyon, South-Eastern Tibet (Lepidoptera, Nymphalidae). Atalanta, 47, 205-210.

  • Huang, H. & Zhu, J.Q. (2016) Ahlbergia maoweiweii sp. n. from Shaanxi, China with revisional notes on similar species (Lepidoptera: Lycaenidae). Zootaxa, 4114 (4), 409-433.


  • Hubner, J. (1818) Zutrage zur Sammlung exotischer Schmettlinge. Vol. 1. bey dem Verfasser zu finden, Augsburg, 40 pp., 35 pls. [in German]

  • Hubner, J. [1819] Verzeichniss Bekannter Schmettlinge. Vol. 3. bey dem Verfasser zu finden, Augsburg, 16 pp. [in German]

  • Hubner, J. [1819] Verzeichniss Bekannter Schmettlinge. Vol. 6. bey dem Verfasser zu finden, Augsburg, 16 pp. [in German]

  • Hubner, J. [1821] Sammlung Exotischer Schmetterlinge. Vol. 2. im Verlag der Hubner'schen Werke, Augsburg, 24 pp. [in German]

  • Johnson, K. (1992) The Palaearctic "elfin" butterflies (Lycaenidae, Theclinae). Neue Entomologische Nachrichten, 29, 5-141.

  • Katayama, T. (1988) Notes on Pazala glycerion (Gray), with description of a new subspecies from North-west India. Gekkan- Mushi, 203, 8-9, pl. 1. [in Japanese with English description]

  • Kimura, M. (1980) A simple method for estimating evolutionary rates of base substitutions through comparative studies of nucleotide sequences. Journal of Molecular Evolution, 16, 111-120.


  • Koiwaya, S. (1993) Description of three new genera, eleven new species and seven new subspecies of butterflies from China. Studies of Chinese Butterflies, 2, 43-111, pl. 9-27. [in Japanese with English descriptions]

  • Leech, J.H. (1893-94) Butterflies from China, Japan, and Corea. Part II. R. H. Porter, London, 385 pp., 15 pls. [pp. 297-681, pls. 29-43]

  • Lepage, T., Bryant D., Philippe H. & Lartillot N. (2007) A general comparison of relaxed molecular clock models. Molecular Biology and Evolution, 24, 2669-2680.


  • Li, K.Q., Wang, Y.Z., Dong, D.Z. & Zhang, L.K. (2015) Catalog of insect type specimens preserved at the Kunming Insititute of Zoology, Chinese Academy of Science with corrections of some specimens. Zoological Research, 36, 263-284.


  • Lu, C.C., Wu, L.W., Jiang, G.F., Deng, H.L., Wang, L.H., Yang, P.S. & Hsu, Y.F. (2009) Systematic status of Agehana elwesi f. cavaleriei based on morphological and molecular evidence. Zoological Studies, 48, 270-279.

  • Lu, J., Hu, S.J., Ma, X.Y., Chen, J.M., Li, Q.Q. & Ye, H. (2014) Origin and expansion of the Yunnan shoot borer, Tomicus yunnanensis (Coleoptera: Scolytinae): a mixture of historical natural expansion and contemporary human-mediated relocation. PLoS ONE, 9, e111940.


  • Mell, R. (1923) Noch unbeschriebene Lepidopteren aus Sudchina. II. Deutsche Entomologische Zeitschrift, 1923, 153-160.

  • Mell, R. (1935) Noch unbeschriebene chinesische Lepidopteren. IV. Mitteilungen der Deutschen Entomologischen Gesellschaft, 6, 36-38.

  • Miller, J. S. (1987) Phylogenetic studies in the Papilioninae (Lepidoptera: Papilionidae). Bulletin of the American Museum of Natural History, 186, 365-512.

  • Miller, M.A., Schwartz, T., Pickett, B.E., He, S., Klem, E.B., Scheuermann, R.H., Passarotti, M., Kaufman, S. & O'Leary, M.A. (2015) A RESTful API for access to phylogenetic tools via the CIPRES science gateway. Evolutionary Bioinformatics Online, 11, 43-48.


  • Moritz, C. & Cicero, C. (2004) DNA barcoding: promise and pitfalls. PLoS Biology, 2, e354.


  • Moore, F. (1888) Additional Species Family Papilionidae. In: Hewitson & Moore (1879-1888), Description of New Indian Lepidopterous Insects from the Collection of the Late Mr. W.S. Atkinson, M.A., F.L.S., & c., Director of the Public Instruction, Bengal. Rhopalocera. Part III. Asiatic Society of Bengal, Calcutta, pp. 283-285.

  • Mutanen, M., Kivela, S.M., Vos, R.A., Doorenweerd, C., Ratnasingham, S., Hausmann, A., Huemer, P., Dinca, V., van Nieukerken, E.J., Lopez-Vaamonde, C., Vila, R., Aarvik, L., Decaens, T., Efetov, K.A., Hebert, P.D.N., Johnsen, A., Karsholt O., Pentinsaari, M., Rougerie, R., Segerer, A., Tarmann, G., Zahiri, R. & Godfray, H.C.J. (2016) Species-level para- and polyphyly in DNA barcode gene trees: strong operational bias in European Lepidoptera. Systematic Biology, 65, 1024-1040.


  • Murayama, S. (1982) Notes on some butterflies of North Thailand and China. New Entomologist, 31, 1-6.

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

  • O'Neill, S.L., Giordano, R., Colbert, A.M.E., Karr, T.L. & Robertson, H.M. (1992) 16S rRNA phylogenetic analysis of the bacterial endosymbionts associated with cytoplasmic incompatibility in insects. Proceedings of the National Academy of Sciences of the United States of America, 89, 2699-2702.


  • Oberthur, C. (1876) Lepidopteres nouveaux de la Chine. Etudes d'Entomologie, 2, 13-34, pl. 1-4. [in French]

  • Oberthur, C. (1879) Catalogue raissone des Papilionidae de la collection de Ch. Oberthur. Etudes d'Entomologie, 4, 19-102, 107-117, pls. I-VI. [in French]

  • Okano, K. (1986) Two new subspecies of butterflies from Thailand. Tokurana, 11, 1-8.

  • Posada, D. (2008) jModelTest: phylogenetic model averaging. Molecular Biology and Evolution, 25, 1253-1256.


  • Racheli, T. & Cotton, A.M. (2009) Guide to the Butterflies of the Palearctic Region. Papilionidae. Part I. Omnes Artes, Milano, 70 pp.

  • Rehm, P., Borner, J., Meusemann, K., von Reumont, B.M., Simon, S., Hadrys, H., Misof, B. & Burmester, T. (2011) Dating the arthropod tree based on large-scale transcriptome data. Molecular Phylogenetics and Evolution, 61, 880-887.


  • Riley, N.D. (1929) Revisional notes on the genus Heliophorus with descriptions of new forms. Journal of the Bombay Natural History Society, 33, 384-402.

  • Ronquist, F., Teslenko, M., Van Der Mark, P., Ayres, D.L., Darling, A., Hohna, S., Larget, B., Liu, L., Suchard, M.A. & Huelsenbeck, J.P. (2012) MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology, 61, 539-542.


  • Saigusa, T., Nakanishi, A., Shima, H. & Yata, O. (1982) Phylogeny and Geographical Distribution of the Swallow-Tail Subgenus Graphium (Lepidoptera: Papilionidae). Entomologia Generalis, 8, 59-69.

  • Sanmartin, I., Enghoff, H. & Ronquist, F. (2001) Patterns of animal dispersal, vicariance and diversification in the Holarctic. Biological Journal of the Linnean Society, 73, 345-390.


  • Schenk, J.J. (2016) Consequences of secondary calibrations on divergence time estimates. PLoS ONE, 11, e0148228.


  • Scopoli, J.A. (1777) Introductio ad historiam naturalem sistens genera lapidum, plantarum, et animalium. Gerle, Prague, x + 506 pp.

  • Shiraiwa, K., Cong, Q. & Grishin, N.V. (2014) A new Heraclides swallowtail (Lepidoptera, Papilionidae) from North America is recognised by the pattern on its neck. Zookeys, 468, 85-135.


  • Shirozu, T. (1979) On the natural groups and their phylogenetic relationships of the genus Ypthima Hubner mainly from Asia (Lepidoptera: Satyridae). Sieboldia, 4, 231-295.

  • Smith, C.R. & Vane-Wright, R.I. (2001) A review of the Afrotropical species of the genus Graphium (Lepidoptera: Rhopalocera: Papilionidae). Bulletin of the British Museum (Natural History) Entomology, 70, 503-719.

  • Song, H., Buhay, J.E., Whiting, M.F. & Crandall, K.A. (2008) Many species in one: DNA barcoding overestimates the number of species when nuclear mitochondrial pseudogenes are coamplified. Proceedings of the National Academy of Sciences of the United States of America, 105, 13486-13491.


  • Takahashi, M. (2000) A revision of the Ypthima sakra group (Lepidoptera, Satyridae) in Taiwan, China, with description of a new species. Transactions of the Lepidopterological Society of Japan, 51, 1-18. [in Japanese with English summary]

  • Tamura, K., Stecher, G., Peterson, D., Filipski, A. & Kumar, S. (2013) MEGA6: Molecular evolutionary genetics analysis version 6.0. Molecular Biology and Evolution, 30, 2725-2729.


  • Thompson, J.D., Higgins, D.G. & Gibson, T.J. (1994) CLUSTAL W: improving the sensitivity of progressive multiple sequence alignment through sequence weighting, position-specific gap penalties and weight matrix choice. Nuclear Acids Research, 22, 4673-4680.


  • Thorne, J.L. & Kishino, H. (2002) Divergence time and evolutionary rate estimation with multilocus data. Systematic Biology, 51, 689-702.


  • Thorne, J.L., Kishino, H. & Painter, I.S. (1998) Estimating the rate of evolution of the rate of molecular evolution. Molecular Biology and Evolution, 15, 1647-1657.


  • Tsao, W.C. & Yeh, W.B. (2008) DNA-Based discrimination of subspecies of swallowtail butterflies (Lepidoptera: Papilioninae) from Taiwan. Zoological Studies, 47, 633-643.

  • Wang, X. & Gu, M.B. (2017) Species of butterflies and its host in Nanling National Nature Reserve. Journal of Zhejiang Forestry Science and Technology, 36, 37-45.

  • Will, K.W. & Rubinoff, D. (2004) Myth of the molecule: DNA barcodes for species cannot replace morphology for identification and classification. Cladistics, 20, 47-55.


  • Xing, Y. & Ree, R.H. (2017) Uplift-driven diversification in the Hengduan Mountains, a temperate biodiversity hotspot. Proceedings of the National Academy of Sciences of the United States of America, 114, E3444-E3451.


  • Yago, M. (2002) Comparative morphology and identification of the subgenus Kulua, with description of a new species from Vietnam (Lepidoptera, Lycaenidae, Heliophorus). Tijdschrift voor Entomologie, 145, 145-171.


  • Yago, M. (2003) A new species of Heliophorus Geyer from Nepal, with a key to the Nepalese species (Lepidoptera, Lycaenidae). Bulletin de la Societe Entomologique de France, 108, 27-34.

  • Yago, M., Saigusa, T. & Nakanishi, A. (2000) Rediscovery of Heliophorus yunnani D'Abrera and its systematic position with intrageneric relationship in the genus Heliophorus (Lepidoptera: Lycaenidae). Entomological Science, 3, 81-100.

  • Yago, M., Saigusa, T. & Nakanishi, A. (2002) A Revision of the Heliophorus kohimensis group (Lepidoptera: Lycaenidae). Entomological Science, 5, 375-388.

  • Yokochi, T. (2010) Revision of the subgenus Limbusa Moore, [1897] (Lepidoptera, Nymphalidae, Adoliadini) Part 1. Systematic arrangement and taxonomic list. Bulletin of the Kitakyushu Museum of Natural History and Human History, Series A, 8, 19-67.

  • Yokochi, T. (2011) Revision of the subgenus Limbusa Moore, [1897] (Lepidoptera, Nymphalidae, Adoliadini) Part 2. Group division and description of species (1). Bulletin of the Kitakyushu Museum of Natural History and Human History, Series A, 9, 9-106.

  • Yokochi, T. (2012) Revision of the subgenus Limbusa Moore, [1897] (Lepidoptera, Nymphalidae, Adoliadini) Part 3. Description of species (2). Bulletin of the Kitakyushu Museum of Natural History and Human History, Series A, 10, 9-100.

  • Zhao, J., Hu, S.J., He, Q.J., Yi, C.H., He, J., Feng, Z.W., Yang, J.H. & Chen, P. (2017) Genetic diversity of Bhutanitis thaidina Blanchard, based on mitochondrial COI and ND1. Journal of Fujian Agricultural and Forestry University, Natural Science Edition, 46, 387-391. [in Chinese with English abstract]

  • Zhu, L.X. & Wu, X.B. (2011) Phylogenetic evaluation of the taxonomic status of Papilio maackii and P. syfanius (Lepidoptera: Papilionidae). Zoological Research, 32, 248‾254.


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