Polyura

Life-forms

Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Protostomia
Cladus: Ecdysozoa
Cladus: Panarthropoda
Phylum: Arthropoda
Subphylum: Hexapoda
Classis: Insecta
Cladus: Dicondylia
Subclassis: Pterygota
Cladus: Metapterygota
Infraclassis: Neoptera
Cladus: Eumetabola
Cladus: Endopterygota
Superordo: Panorpida
Cladus: Amphiesmenoptera
Ordo: Lepidoptera
Subordo: Glossata
Cladus: Coelolepida
Cladus: Myoglossata
Cladus: Neolepidoptera
Infraordo: Heteroneura
Cladus: Eulepidoptera
Cladus: Ditrysia
Cladus: Apoditrysia
Cladus: Obtectomera
Superfamilia: Papilionoidea

Familia: Nymphalidae
Subfamilia: Charaxinae
Tribus: Charaxini
Genus: Polyura
Name

Polyura Billberg, 1820

Type species: Papilio pyrrhus Linnaeus,
Synonymy

Eulepis Scudder, 1875, (Nec Eulepis Billberg, 1820: 80) TS: Papilio athamas Drury.
Murwareda Moore, [1896]: 263, TS: Murwareda dolon [sic] Westwood.
Pareriboea Roepke, 1938, TS: Papilio athamas Drury.

List of Species

P. aeolus – P. agrarius – P. alphius – P. andrewsi – P. arja – P. athamas – P. athamis – P. attila – P. bharata – P. buruensis – P. caphontis – P. charata – P. clitarchus – P. cognatus – P. dehanii – P. delphis – P. dolon – P. eleganta – P. epigenes – P. eudamippus – P. gamma – P. gilolensis – P. hamastiformis – P. hebe – P. inopinatus – P. jalysus – P. jupiter – P. luzonica – P. moori – P. mutata – P. narcaeus – P. nepenthes – P. noko – P. paulettae – P. posidonius – P. pyrrhus – P. pyrrichia – P. sacco – P. schreiber – P. sempronius – P. smilesi – P. walshae – P. weismanni
Taxa In Synonymy

P. aborica – P. abrupta – P. acuminata – P. acuta (Polyura athamas) – P. admiralitatis – P. aemiliani – P. albanus – P. aloranus – P. andamanica (Polyura athamas) – P. antigonus – P. aristophanes – P. arna – P. arnoldi – P. assamensis (Polyura schreiber) – P. attalus (Polyura athamas) – P. australis – P. babbericus – P. bandanus – P. batavianus – P. baweanicus – P. buruanus – P. canomaculatus – P. carolus – P. celetis – P. centralis – P. chalazias – P. chersonesus – P. chlorus – P. chota – P. clavata – P. clitiphon – P. concha – P. cupidinius – P. cygnus – P. delphinion – P. dexippus (Polyura athamas) – P. editha – P. entheatus – P. ephebus – P. falculus – P. faliscus – P. fallacides – P. fallax – P. formosana – P. fruhstorferi – P. fugator – P. galaxia – P. ganymedes – P. glauca – P. grandis – P. hamasta – P. heracles – P. intermedia – P. jamblichus – P. javanus – P. jovis – P. juta – P. kaba – P. kadenii – P. kailicus – P. kalaonicus – P. kangeanus – P. kannegieteri (Polyura athamas) – P. keianus – P. kiangsiensis – P. kronos – P. kuangtungensis – P. lemoulti – P. lettianus – P. lissainei – P. lombokianus – P. madeus – P. magniplagus – P. major – P. malayica (Polyura schreiber) – P. mandarinus – P. marginalis – P. marginepunctatus – P. meghaduta – P. menaius – P. menedemus – P. niasica (Polyura schreiber) – P. nigra – P. nigrobasalis – P. nikias – P. niveus – P. obiensis – P. oitylus – P. othonis – P. palawanica (Polyura athamas) – P. pallida – P. peninsularis – P. phrixus – P. piepersianus – P. plautus – P. praedictus – P. pyrrhulus – P. monochromus – P. quaesita – P. rectifascia – P. richthofeni – P. roeberi – P. romanus – P. rothschildi – P. saida – P. samatha – P. sandakanus – P. satyrina – P. scipio – P. seitzi – P. sinica – P. smerdis – P. splendens – P. stratiocus – P. sulthan – P. sumatrana – P. sumbaensis – P. thawgawa – P. thibetanus – P. tiberius – P. tisamenus (Polyura schreiber) – P. triphonius – P. tyrtaeus – P. uraeus (Polyura athamas) – P. valesius – P. vernus – P. wardii (Polyura schreiber) – P. watubela – P. whiteheadi
References

Abang, F., Treadaway, C.G. & Schröder, H.G., 2004. A contribution to the butterfly fauna of the Island of Balambangan (Malaysia, Sabah) (Insecta: Lepidoptera). Futao 47: 6–13. Reference page.
Billberg, G.J. 1820. Enumeratio insectorum in museo Gust. Joh. Billberg. Stockholm, 138 pp. BHL Reference page.
Fang, L.J. & Zhang, Y.L. 2007. Taxonomy of the genus Polyura Billberg (Lepidoptera: Nymphalidae) with description of a new species of China. Acta Zootaxonomica Sinica 33 (2). Reference page.
Fang & Zhang, Y.L. 2018. Review of the genus Polyura Billberg (Lepidoptera: Nymphalidae) from China with description of a new species. Entomotaxonomia 40(1): 27–45. Full article (PDF). Reference page.
Hanafusa, H., 1991. Five new subspecies of Indonesian butterflies (Lep.: Papilionidae, Nymphalidae, Riodinidae). Futao 8: 17–19. Reference page.
Hanafusa, H. 1993. A list of butterflies from Mentawai Islands, Indonesia (3). Futao 14: 8–25. Reference page.
Hanafusa, H., 1994. Eight new butterflies from Indonesia (Lepidoptera: Papilionidae, Pieridae, Satyridae, Nymphalidae, Riodinidae). Futao 17: 18–23. Reference page.
Hanafusa, H., 1995. A new subspecies of Polyura schreiber from Sumbawa Island, Indonesia (Lepidoptera: Nymphalidae). Futao 20: 16–18. Reference page.
Hanafusa, H., 1998. A new subspecies of Polyura galaxia (Butler, 1866) from Sawu Is., Indonesia (Lepidoptera : Nymphalidae). Futao 30: 1. Reference page.
Moore, F., 1896–1899. Lepidoptera Indica Vol. 3. Rhopalocera: Nymphalidae: Nymphalinae, Potamina, Euthaliina, Limenitina. 254 pp., pls 191-286. Reference page.
Shakespeare, T.J.; Shakespeare, Z.J.; Shakespeare, T.P. 2013: An observation of Polyura sempronius (Fabricius) (Lepidoptera: Nymphalidae) feeding on bandicoot droppings in south-east Queensland. Australian entomologist 40(4): 175-176. [Not seen] Reference page.
Smiles, R., 1982. The taxonomy and phylogeny of the genus Polyura Billberg (Lepidoptera: Nymphalidae). Bulletin of th British Museum (Natural History); Entomology. 44(3): 115-237. Reference page.
Toussaint, E.F.A., Morinière, J., Müller, C.J., Kunte, K., Turlin, B., Hausmann, A. & Balke, M. 2015. Comparative molecular species delimitation in the charismatic Nawab butterflies (Nymphalidae, Charaxinae, Polyura). Molecular Phylogenetics and Evolution 91: 194–209. DOI: 10.1016/j.ympev.2015.05.015. Reference page.
Turlin, B. & H. Sato 1995. Description of new subspecific taxa in the genus Polyura Billberg (Lepidoptera: Nymphalidae) from the Moluccas and Solomons Archipelagos. Futao 20: 7–15. Reference page.

Polyura is a subgenus of butterflies also referred to as Nawab butterflies[1][2] and belonging to the brush-footed butterfly subfamily Charaxinae, or leafwing butterflies. Like the large and conspicuous forest queens (subgenus Euxanthe), they belong to the genus Charaxes, unique genus of the tribe Charaxini.[3][4]

Distribution

Polyura butterflies are native to the Indomalayan and Australasian realms. They are widespread from Pakistan to Okinawa Island, and from China to Pacific Islands (Fiji, New Caledonia, Vanuatu).
Systematics

The subgenus Polyura was described by the Swedish naturalist Gustaf Johan Billberg in 1820.[1]
The type species is Polyura pyrrhus (Linnaeus).

Taxonomy

The subgenus was revised in 1982 by Robert Leslie Smiles based on morphological characters.[2]

The genus Polyura was synonymized with the genus Charaxes in 2009 in a study investigating phylogenetic relationships among Charaxini using DNA sequencing.[3] However the genus Charaxes comprises many morphologically very different groups such as the subgenera Euxanthe and Polyura. A phylogenomic study is ongoing to resolve the phylogenetic placements of these different groups. The sister-group to Polyura seems to comprise the African species Charaxes paphianus and Charaxes pleione.[3][4] Southeast Asian species of the genus Charaxes do not seem to be closely related to species of the subgenus Polyura.[3]

A comprehensive molecular phylogeny of the subgenus as well as several taxonomic studies have allowed a complete revision of the group.[4][5][6] A follow-up study investigating the biogeographical history of the subgenus suggested an origin in the Miocene about 12 million years ago in mainland Asia. The Australasian region was colonized later through dispersal.
Species

The subgenus is divided in three morphological groups supported by molecular phylogenetics:[4][5][6]

P. athamas group:

Charaxes agrarius (Swinhoe, 1887) – Anomalous nawab – southern India, Burma
Charaxes alphius (Staudinger, 1886) – Staudinger's nawab – Sulawesi, Timor
Charaxes arja (C. & R. Felder, [1867]) – Pallid nawab – north-eastern India, Sikkim, Assam, Burma, Thailand, Indochina
Charaxes athamas (Drury, [1773]) – Common nawab
Charaxes attalus (C. & R. Felder, 1867) - Java, Sumatra
Charaxes bharata (Leech, 1891) – India
Charaxes hebe (Butler, [1866]) – Plain nawab
Charaxes jalysus (C. & R. Felder, [1867]) – Malaysia, Sunda Islands, Thailand
Charaxes luzonicus (Rothschild, 1899) – Philippines
Charaxes moori (Distant, 1883) – Malayan nawab – Malaysia, Sunda Islands
Charaxes paulettae (Toussaint, 2015) – India, Pakistan, Thailand
Charaxes schreiber (Godart, [1824]) – Blue nawab

P. eudamippus group:

Charaxes delphis (Doubleday, 1843) – Jewelled nawab – Assam, Burma, Malaya, Sumatra, Borneo
Charaxes dolon (Westwood, 1847) – Stately nawab – Tibet, China, Kulu, Assam, Burma
Charaxes eudamippus (Doubleday, 1843) – Great nawab – central China, Kumaon, Assam, Burma
Charaxes narcaeus (Hewitson, 1854) – China nawab – China, Naga Hills, Abor Valley, northern Burma
Charaxes nepenthes (Grose-Smith, 1883) – – eastern China
Charaxes posidonius (Leech, 1891) – Tibet, China
Charaxes weismanni – (Fritze 1894) – Okinawa Island.Sometimes subspecies of Polyura eudamippus

P. pyrrhus group:

Charaxes andrewsi (Butler, 1900) – Christmas emperor – Christmas Island
Charaxes bicolor (Turlin & Sato, 1995) – Solomon Islands
Charaxes caphontis (Hewitson, 1863) – Fiji
Charaxes clitarchus (Hewitson, 1874) – New Caledonia
Charaxes cognatus Vollenhoven, 1861 – Sulawesi blue nawab – Sulawesi
Charaxes dehanii (Westwood, 1850) – Java, Sumatra
Charaxes epigenes (Godman & Salvin, 1888) – Solomon Islands
Charaxes gamma (Lathy, 1898) – New Caledonia
Charaxes gilolensis (Butler, 1869) – Moluccas
Charaxes inopinatus (Röber, 1939) – New Britain
Charaxes jupiter (Butler, 1869) – New Guinea
Charaxes pyrrhus (Linnaeus, 1758) – Tailed nawab, tailed emperor – Ambon, Ceram
Charaxes sacco Smart, 1977 – Vanuatu
Charaxes smilesi (Toussaint, 2015) – Ceram
Charaxes sempronius (Fabricius, 1793) – Tailed emperor, four tail – Australia, Lesser Sunda Islands

References

Billberg, Gustaf Johan (1820). Enumeratio insectorum in Museo. p. 79. doi:10.5962/bhl.title.49763.
Smiles, Robert L. (1982). "The taxonomy and phylogeny of the genus Polyura Billberg (Lepidoptera: Nymphalidae)". Bull. Br. Mus. Nat. Hist. (Ent.). 44 (3): 115–237.
Aduse-Poku, Kwaku; Vingerhoedt, Eric; Wahlberg, Niklas (2009). "Out-of-Africa again: A phylogenetic hypothesis of the genus Charaxes (Lepidoptera: Nymphalidae) based on five gene regions". Molecular Phylogenetics and Evolution. 53 (2): 463–478. doi:10.1016/j.ympev.2009.06.021. PMID 19580878.
Toussaint, Emmanuel F.A.; Morinière, Jérôme; Müller, Chris J.; Kunte, Krushnamegh; Turlin, Bernard; Hausmann, Axel; Balke, Michael (2015). "Comparative molecular species delimitation in the charismatic Nawab butterflies (Nymphalidae, Charaxinae, Polyura)". Molecular Phylogenetics and Evolution. 91: 194–209. doi:10.1016/j.ympev.2015.05.015. PMID 26021440.
Toussaint, Emmanuel F.A.; Morinière, Jérôme; Lam, Athena; Balke, Michael (2016). "New Insights into the Systematics of the GenusPolyura Billberg, 1820 (Nymphalidae, Charaxinae) with an Emphasis on theP. Athamas Group". Journal of the Lepidopterists' Society. 70 (2): 145–152. doi:10.18473/lepi.70i2.a10. S2CID 54091020.
Toussaint, Emmanuel F.A.; Morinière, Jérôme; Lam, Athena; Turlin, Bernard; Fls, Michael Balke (2016). "Bayesian Poisson tree processes and multispecies coalescent models shed new light on the diversification of Nawab butterflies in the Solomon Islands (Nymphalidae, Charaxinae, Polyura)". Zoological Journal of the Linnean Society. 178 (2): 241–256. doi:10.1111/zoj.12413.