:''for the monotypic genus of plants, see

Polyura geminata ''Polyura geminata'' is a species of flowering plant in the family Rubiaceae. It is the sole species in genus ''Polyura''. It is a subshrub native to the eastern Himalayas, including Bhutan, Arunachal Pradesh, and Assam Assam (, , ) is a s ...

'' ''Polyura'' is a

subgenus In biology, a subgenus ( 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 the ge ...

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 ...

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 botanical subfamily names with "-oideae", and zo ...

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 ''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 Nymphalidae, brush-footed butterfly subfamily Charaxinae, or leafwing butterflies. They belong to the tribe ...

'', 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 use of the term in English is in the discipline of anthropology. The definition is contested, in part due to conflict ...

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 southern A ...

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 Ind ...

and

Australasian realm The Australasian realm is one of eight biogeographic realms 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 Ne ...

s. They are widespread from

Pakistan Pakistan, officially the Islamic Republic of Pakistan, is a country in South Asia. It is the List of countries and dependencies by population, fifth-most populous country, with a population of over 241.5 million, having the Islam by country# ...

Okinawa Island , officially , is the largest of the Okinawa Islands and the Ryukyu Islands, Ryukyu (''Nansei'') Islands of Japan in the Kyushu region. It is the smallest and least populated of the five Japanese archipelago, main islands of Japan. The island is ...

, and from

China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...

Pacific Islands The Pacific islands are a group of islands in the Pacific Ocean. They 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 several ...

(

Fiji Fiji, 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 of an archipelago of more than 330 islands—of which about ...

New Caledonia New Caledonia ( ; ) is a group of islands in the southwest Pacific Ocean, southwest of Vanuatu and east of Australia. Located from Metropolitan France, it forms a Overseas France#Sui generis collectivity, ''sui generis'' collectivity of t ...

Vanuatu Vanuatu ( or ; ), officially the Republic of Vanuatu (; ), is an island country in Melanesia located in the South Pacific Ocean. The archipelago, which is of volcanic origin, is east of northern Australia, northeast of New Caledonia, east o ...

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 Sweden, Swedish botanist, zoologist and anatomist, although professionally and by training he was a lawyer and used science and biology as an avocation. The pl ...

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

Linnaeus Carl Linnaeus (23 May 1707 – 10 January 1778), also known after ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...

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, thymine, cytosine, and guanine. The ...

. 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 280px, Generalized scheme of taxonomy Taxonomy is a practice and science concerned with classification or categorization. Typically, there are two parts to it: the development of an underlying scheme of classes (a taxonomy) and the allocation ...

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 epoch (geology), 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 mea ...

about 12 million years ago in mainland Asia. The

Australasia Australasia is a subregion of Oceania, comprising Australia, New Zealand (overlapping with Polynesia), and sometimes including New Guinea and surrounding islands (overlapping with Melanesia). The term is used in a number of different context ...

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 ...

'' (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 ( DCCCLXVII) was a common year starting on Wednesday of the Julian calendar. Events By place Byzantine Empire * September 24 – Emperor Michael III is murdered, by order of his co-emperor Basil I. Basil become ...

– 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 of the Julian calendar, the 773rd year of the Common Era (CE) and Anno Domini (AD) designations, the 773rd year of the 1st millennium, the 73rd year of the 8th century, and t ...

– 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 Ka ...

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

866 __NOTOC__ Year 866 ( DCCCLXVI) was a common year starting on Tuesday of the Julian calendar. Events By place Byzantine Empire * April 21 – Bardas, the regent of the Byzantine Empire, is murdered by Basil the Macedonian at Miletu ...

– 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 of the Julian calendar. Events By date * February 8 – The Tenchō era begins in Japan at the beginning of the reign of the Emperor Junna, bringing an end to 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 zoologic ...

) – Tailed nawab, tailed emperor – Ambon, Ceram * ''Charaxes sacco'' Smart, 1977 – Vanuatu * '' Charaxes smilesi Toussaint nec Rydon'' (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 Taxa described in 1820