''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 between t ...

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 redu ...

subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end subfamily botanical names with "-oideae", and zoologi ...

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 A ...

, or leafwing butterflies. Like the large and conspicuous forest queens (subgenus ''

Euxanthe ''Euxanthe'' is a genus of butterflies in the subfamily Charaxinae. Species There are six species all found in Afrotropical forests. The popular name is forest queen. *Subgenus ''Euxanthe'' Hübner, 819/small> **'' Euxanthe crossleyi'' (Ward, ...

''), 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 language, English is in the discipline of anthropology. This definition is contested, in p ...

Charaxini __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 souther ...

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 and ...

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 an ...

, and from

China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...

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 se ...

(

Fiji Fiji ( , ,; fj, Viti, ; Fiji Hindi: फ़िजी, ''Fijī''), officially the Republic of Fiji, is an island country in Melanesia, part of Oceania in the South Pacific Ocean. It lies about north-northeast of New Zealand. Fiji consists ...

New Caledonia ) , anthem = "" , image_map = New Caledonia on the globe (small islands magnified) (Polynesia centered).svg , map_alt = Location of New Caledonia , map_caption = Location of New Caledonia , mapsize = 290px , subdivision_type = Sovereign st ...

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 genu ...

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, the ...

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 DNA sequencing is the process of determining the nucleic acid sequence – the order of nucleotides in DNA. It includes any method or technology that is used to determine the order of the four bases: adenine, guanine, cytosine, and thymine. Th ...

. However the genus ''

'' comprises many morphologically very different groups such as the subgenera ''

'' and ''Polyura''. A

phylogenomic Phylogenomics is the intersection of the fields of evolution and genomics. The term has been used in multiple ways to refer to analysis that involves genome data and evolutionary reconstructions. It is a group of techniques within the larger fields ...

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''. 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 Taxonomy is the practice and science of categorization or classification. A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...

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, i ...

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 recen ...

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 ecologica ...

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 ''Charaxes agrarius'', the anomalous nawab, is a butterfly found in Asia that belongs to the rajahs and nawabs group, that is, the Charaxinae subfamily of the brush-footed butterflies family. The name is based on their resemblance to the common n ...

'' (Swinhoe, 1887) – Anomalous nawab – southern India, Burma * ''Charaxes alphius'' (Staudinger, 1886) – Staudinger's nawab – Sulawesi, Timor * ''Charaxes arja'' (C. & R. Felder,

867 __NOTOC__ Year 867 (Roman numerals, DCCCLXVII) was a common year starting on Wednesday (link will display the full calendar) of the Julian calendar. Events By place Byzantine Empire * September 23 – Emperor Michael III is murde ...

– Pallid nawab – north-eastern India, Sikkim, Assam, Burma, Thailand, Indochina * ''Charaxes athamas'' (Drury,

773 __NOTOC__ Year 773 ( DCCLXXIII) was a common year starting on Friday (link will display the full calendar) of the Julian calendar. The denomination 773 for this year has been used since the early medieval period, when the Anno Domini calendar e ...

– Common nawab * '' Charaxes attalus'' (C. & R. Felder, 1867) - Java, Sumatra * '' Charaxes bharata'' (Leech, 1891) – India * ''Charaxes hebe'' (Butler, 866 – Plain nawab * ''Charaxes jalysus'' (C. & R. Felder,

– 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, 10th edition of ''Systema Naturae'', the sta ...

) – 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