''Polyura'' is a

subgenus In biology, a subgenus (plural: subgenera) is a taxonomic rank directly below genus. In the International Code of Zoological Nomenclature, a subgeneric name can be used independently or included in a species name, in parentheses, placed betw ...

of butterflies also referred to as Nawab butterflies and belonging to the

brush-footed butterfly The Nymphalidae are the largest family of butterflies, with more than 6,000 species distributed throughout most of the world. Belonging to the superfamily Papilionoidea, they are usually medium-sized to large butterflies. Most species have a red ...

subfamily

Charaxinae __NOTOC__ The Charaxinae, the leafwings, are a nymphalid subfamily of butterflies that includes about 400 species, inhabiting mainly the tropics, although some species extend into temperate regions in North America, Europe, China, and southern ...

, or leafwing butterflies. Like the large and conspicuous forest queens (subgenus '' Euxanthe''), they belong to the genus ''

Charaxes The rajah and pasha butterflies, also known as emperors in Africa and Australia, (genus ''Charaxes'') make up the huge type genus of the brush-footed butterfly subfamily Charaxinae, or leafwing butterflies. They belong to the tribe Charaxini, ...

'', unique genus of the

tribe The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide usage of the term in English is in the discipline of anthropology. This definition is contested, in part due to confl ...

Charaxini.

Distribution

''Polyura'' butterflies are native to the

Indomalayan The Indomalayan realm is one of the eight biogeographic realms. It extends across most of South and Southeast Asia and into the southern parts of East Asia. Also called the Oriental realm by biogeographers, Indomalaya spreads all over the Indi ...

and

Australasian realm The Australasian realm is a biogeographic realm that is coincident with, but not (by some definitions) the same as, the geographical region of Australasia. The realm includes Australia, the island of New Guinea (comprising Papua New Guinea an ...

s. They are widespread from

Pakistan Pakistan ( ur, ), officially the Islamic Republic of Pakistan ( ur, , label=none), is a country in South Asia. It is the world's List of countries and dependencies by population, fifth-most populous country, with a population of almost 24 ...

Okinawa Island is the largest of the Okinawa Islands and the Ryukyu (''Nansei'') Islands of Japan in the Kyushu region. It is the smallest and least populated of the five main islands of Japan. The island is approximately long, an average wide, and has a ...

, and from China to

Pacific Islands Collectively called the Pacific Islands, the islands in the Pacific Ocean are further categorized into three major island groups: Melanesia, Micronesia, and Polynesia. Depending on the context, the term ''Pacific Islands'' may refer to one of ...

( Fiji, New Caledonia,

Vanuatu Vanuatu ( or ; ), officially the Republic of Vanuatu (french: link=no, République de Vanuatu; bi, Ripablik blong Vanuatu), is an island country located in the South Pacific Ocean. The archipelago, which is of volcanic origin, is east of no ...

Systematics

* The subgenus ''Polyura'' was described by the Swedish naturalist

Gustaf Johan Billberg Gustaf Johan Billberg (14 June 1772, Karlskrona – 26 November 1844, Stockholm) was a Swedish botanist, zoologist and anatomist, although professionally and by training he was a lawyer and used science and biology as an avocation. The plant g ...

in 1820. * The type species is ''Polyura pyrrhus'' (

Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, t ...

Taxonomy

The subgenus was revised in 1982 by Robert Leslie Smiles based on morphological characters. The genus ''Polyura'' was synonymized with the genus ''

'' in 2009 in a study investigating phylogenetic relationships among Charaxini using DNA sequencing. However the genus ''

'' 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 ''Charaxes paphianus'', the falcate red charaxes, is a butterfly in the family Nymphalidae. It is found in Sierra Leone, Guinea, Liberia, Ivory Coast, Ghana, Togo, Nigeria, Cameroon, Gabon, the Republic of the Congo, the Central African Republi ...

'' and '' Charaxes pleione''. Southeast Asian species of the genus ''

'' do not seem to be closely related to species of the subgenus ''Polyura''. A comprehensive

molecular phylogeny Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...

of the subgenus as well as several taxonomic studies have allowed a complete revision of the group. A follow-up study investigating the

biogeographical Biogeography is the study of the distribution of species and ecosystems in geographic space and through geological time. Organisms and biological communities often vary in a regular fashion along geographic gradients of latitude, elevation, ...

history of the subgenus suggested an origin in the

Miocene The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recent" ...

about 12 million years ago in mainland Asia. The

Australasia Australasia is a region that comprises Australia, New Zealand and some neighbouring islands in the Pacific Ocean. The term is used in a number of different contexts, including geopolitically, physiogeographically, philologically, and ecolo ...

n region was colonized later through dispersal.

Species

The subgenus is divided in three morphological groups supported by molecular phylogenetics: ''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, 867 – Pallid nawab – north-eastern India, Sikkim, Assam, Burma, Thailand, Indochina * ''Charaxes athamas'' (Drury, 773 – Common nawab * '' Charaxes attalus'' (C. & R. Felder, 1867) - Java, Sumatra * ''

Charaxes bharata ''Polyura athamas'', the common nawab, is a species of fast-flying canopy butterfly found in tropical Asia. It belongs to the Charaxinae (rajahs and nawabs) in the brush-footed butterfly family (Nymphalidae). It occurs in the Himalayas from Kas ...

'' (Leech, 1891) – India * ''Charaxes hebe'' (Butler,

866 __NOTOC__ Year 866 ( DCCCLXVI) was a common year starting on Tuesday (link will display the full calendar) of the Julian calendar. Events By place Byzantine Empire * April 21 – Bardas, the regent of the Byzantine Empire, is murd ...

– Plain nawab * ''Charaxes jalysus'' (C. & R. Felder, 867 – 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,

824 __NOTOC__ Year 824 ( DCCCXXIV) was a leap year starting on Friday (link will display the full calendar) of the Julian calendar. Events By place Europe * November 11 – The '' Constitutio Romana'' establishes the authority of the ...

– 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'' (

1758 Events January–March * January 1 – Swedish biologist Carl Linnaeus (Carl von Linné) publishes in Stockholm the first volume (''Animalia'') of the 10th edition of ''Systema Naturae'', the starting point of modern zoological ...

) – 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

{{Taxonbar, from=Q2703245 Charaxinae Insect subgenera Taxa named by Gustaf Johan Billberg