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Type: Article
Published: 2018-06-28
Page range: 401–446
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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)

Yunnan Key Laboratory of International Rivers and Transboundary Eco-security, Yunnan University, Kunming, 650500, China Institute of International Rivers and Eco-security, Yunnan University, Kunming, 650500, China School of Agriculture, Yunnan University, Kunming, 650500, China
86/2 Moo 5, Tambon Nong Kwai, Hang Dong, Chiang Mai, Thailand
CNRS, UMR 5554 Institut des Sciences de l’Evolution, Université de Montpellier, Place Eugène Bataillon 34095 Montpellier, France
School of Agriculture, Yunnan University, Kunming, 650500, China
College of Life Sciences, Peking University, Beijing, 100871, China
Department of Life Sciences, National Taiwan Normal University, Taipei, 116, Taiwan, China
Laboratory for Animal Genetic Diversity and Evolution of Higher Education in Yunnan Province, Yunnan University, Kunming, 650091, China
Laboratory of Ecology and Evolutionary Biology, Yunnan University, Kunming, 650500, China
Lepidoptera Leptocircini Pazala new species India China Vietnam


The previously recognised closely related species Graphium (Pazala) mandarinus (Oberthür, 1879) and G. (P.) sichuanica (Koiwaya, 1993) are shown to comprise seven species as a result of both molecular and morphological analysis. Molecular dating analysis is also performed on the mandarinus group in order to investigate the divergence time of the taxa. Two taxa, G. (P.) garhwalica (Katayama, 1988) stat. nov. and G. (P.) paphus (de Nicéville, 1886) stat. nov., are raised from subspecific to specific status; G. (P.) hoeneanus Cotton & Hu nom. nov., stat. rev. is separated from sichuanica at species level; and two previously unrecognised new species, G. (P.) daiyuanae Hu, Zhang & Cotton sp. nov. and G. (P.) confucius Hu, Duan & Cotton sp. nov. are described from Vietnam and China respectively, the latter being sympatric with nominate G. (P.) mandarinus. The identity of the lectotype of G. (P.) mandarinus is confirmed and a lectotype is designated for the taxon Papilio Glycerion Gray, 1831. A new subspecies of G. (P.) mandarinus is described from western Yunnan and northern Myanmar, G. (P.) mandarinus stilwelli Cotton & Hu ssp. nov.



  1. Alphéraky, S. (1897) Lépidoptères des provinces chinoises Sé-Tchouen et Kham recueillis, en 1893, par M-r G. N. Potanine. In: Romanoff, N.M. (Ed.), Mémoires sur les Lépidoptères. Vol. 9. P.P. Soïkine, St.-Pétersbourg, 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 high-performance computing library for statistical phylogenetics. Systematic Biology, 61, 170–173.

    Baker, D.B. (1995) A review of the Asian species of the genus Euaspis Gerstäcker (Hymenoptera: Apoidea: Megachilidae). Zoölogische Mededelingen, Leiden, 69, 281–302.

    Bertheau, C., Schuler, H., Krumböck, 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 Europäischen 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 för 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 Nicéville, L. (1886) On some new Indian butterflies. Journal of the Asiatic Society of Bengal, 55, Part II (3), 249–256, pl. XI.

    de Nicéville, 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 Linnéichen Gattungen Papilio und Sphinx. Magazin für 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) Zuträge zur Sammlung exotischer Schmettlinge. Vol. 4. im Verlag der Hübner’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.

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

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

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

    Hübner, J. [1821] Sammlung Exotischer Schmetterlinge. Vol. 2. im Verlag der Hübner’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.

    Lü, 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 Südchina. 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., Kivelä, S.M., Vos, R.A., Doorenweerd, C., Ratnasingham, S., Hausmann, A., Huemer, P., Dincă, V., van Nieukerken, E.J., Lopez-Vaamonde, C., Vila, R., Aarvik, L., Decaëns, 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.

    Oberthür, C. (1876) Lépidoptères nouveaux de la Chine. Études d’Entomologie, 2, 13–34, pl. 1–4. [in French]

    Oberthür, C. (1879) Catalogue raissoné des Papilionidae de la collection de Ch. Oberthür. Études 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., Höhna, 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.

    Sanmartín, 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.

    Shirôzu, T. (1979) On the natural groups and their phylogenetic relationships of the genus Ypthima Hübner 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 Société 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.