Polyura

Polyura
	Polyura schreiber
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Lepidoptera
Family:	Nymphalidae
Tribe:	Charaxini
Genus:	Polyura; Billberg, 1820
Species
	Many, see text

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, 1820; Enum. Ins. Mus. Billb. : 79
Smiles, R. L. (1982). The taxonomy and phylogeny of the genus Polyura Billberg (Lepidoptera: Nymphalidae). British Museum (Natural History).
Aduse-Poku, K., Vingerhoedt, E., & Wahlberg, N. (2009). Out-of-Africa again: A phylogenetic hypothesis of the genus< i> Charaxes</i>(Lepidoptera: Nymphalidae) based on five gene regions. Molecular Phylogenetics and Evolution, 53(2), 463-478.
Toussaint EFA, Morinière J, Müller CJ, 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.
New Insights into the Systematics of the Genus Polyura Billberg, 1820 (Nymphalidae, Charaxinae) with an Emphasis on the P. athamas Group », Journal of the Lepidopterists' Society, vol. 70, no 2, 1er juin 2016, p. 145–152
Toussaint, E.F., Morinière, J., Lam, A. and Turlin, B., 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), pp.241-256.

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[:0-1] Billberg, 1820; Enum. Ins. Mus. Billb. : 79

[:1-2] Smiles, R. L. (1982). The taxonomy and phylogeny of the genus Polyura Billberg (Lepidoptera: Nymphalidae). British Museum (Natural History).

[:2-3] Aduse-Poku, K., Vingerhoedt, E., & Wahlberg, N. (2009). Out-of-Africa again: A phylogenetic hypothesis of the genus< i> Charaxes</i>(Lepidoptera: Nymphalidae) based on five gene regions. Molecular Phylogenetics and Evolution, 53(2), 463-478.

[:3-4] Toussaint EFA, Morinière J, Müller CJ, 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.

[:4-5] New Insights into the Systematics of the Genus Polyura Billberg, 1820 (Nymphalidae, Charaxinae) with an Emphasis on the P. athamas Group », Journal of the Lepidopterists' Society, vol. 70, no 2, 1er juin 2016, p. 145–152

[:5-6] Toussaint, E.F., Morinière, J., Lam, A. and Turlin, B., 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), pp.241-256.