Arhopala

Arhopala is a very large genus of gossamer-winged butterflies (Lycaenidae). They are the type genus of the tribe Arhopalini. In the relatively wide circumscription used here, it contains over 200 species collectively known as oakblues. They occur from Japan throughout temperate to tropical Asia south and east of the Himalayas to Australia and the Solomon Islands of Melanesia. Like many of their relatives, their caterpillars are attended and protected by ants (myrmecophily). Sexual dichromatism is often prominent in adult oakblues.[1]

Arhopala
Arhopala japonica is of unclear affiliations. When Narathura was considered distinct, it was placed there.
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Lepidoptera
Family:	Lycaenidae
Tribe:	Arhopalini
Genus:	Arhopala Boisduval, 1832
Type species
Arhopala phryxus Boisduval, 1832
Diversity
About 220 species
Synonyms
Several, see text

Underwing pattern of unidentified Arhopala species, Selangor on Peninsular Malaysia

The genus' delimitation versus Amblypodia and Flos has proven to be problematic; not all issues are resolved and the assignment of species to these genera must be considered somewhat provisional.

Systematics

As circumscribed here, this genus includes many formerly independent genera. Junior synonyms of Arhopala are:[1]

• Acesina Moore, 1884
• Aurea Evans, 1957
• Daranasa Moore, 1884 (lapsus)
• Darasana Moore, 1884
• "Iois" Doherty, 1899 (nomen nudum)
• Narathura Moore, 1879
• Nilasera Moore, 1881
• Panchala Moore, 1882
• Satadra Moore, 1884

The species have been provisionally sorted into groups of presumed closest relatives, but many species remain insufficiently studied even for such a preliminary assessment at present. For example, A. phryxus – the type species used by Jean Baptiste Boisduval when he described Arhopala in 1832 – was established at the same time as and specifically for this genus. It is considered to be a valid species of unclear affiliations, but it is suspected that Boisduval's taxon is a junior synonym of A. thamyras – the namesake of its species group – which had been described as Papilio thamyras by Carl Linnaeus already in 1764.[2]

Molecular phylogenetic studies have only sampled a fraction of the known diversity of oakblues, but as it seems at least some of the groups represent clades that could justifiably be treated as subgenera. It is also to be seen, however, if the genus is monophyletic in the loose sense as used here, or would need to be split up again.[1]

Species

Species are listed alphabetically, while the groups are listed in the presumed phylogenetic sequence:[2]

Bright oakblue (Arhopala madytus: centaurus group)

Anthelus bushblue (Arhopala anthelus: anthelus group)

Male opal oakblue (Arhopala opalina: camdeo group)

Female Tytler's rosy oakblue (Arhopala allata allata: oenea group) upperside (left) and underside

Arhopala alitaeus mirabella (alitaeus group)

Male de Niceville's oakblue (Arhopala agrata: agrata group)

Male Sylhet oakblue (Arhopala silhetensis: cleander group)

Female Arhopala horsfieldi (eumolphus group)

Malayan oakblue (Arhopala ammon: ganesa group)

centaurus group (subgenus Nilasera if valid) Arhopala acron Arhopala adherbal Grose-Smith, 1902 Arhopala admete (Hewitson, 1863) Arhopala aexone (Hewitson, [1863]) Arhopala alkisthenes Fruhstorfer, 1914 Arhopala amantes – large oakblue Arhopala ander Arhopala araxes C. & R.Felder, [1865] Arhopala azenia Arhopala centaurus – centaur oakblue, dull oakblue Arhopala eucolpis Arhopala eurisus Arhopala kirwinii Bethune-Baker, 1903 Arhopala leander (Evans, 1957) Arhopala madytus – bright oakblue Arhopala meander Boisduval, 1832 Arhopala micale Arhopala philander C. & R.Felder, [1865] Arhopala pseudocentaurus (Doubleday, 1847) Arhopala sophrosyne Arhopala styx Arhopala wanda agesias group Arhopala agesias (Hewitson, 1862) Arhopala ijanensis Bethune-Baker, 1897 (tentatively placed here) Arhopala kinabala Druce, 1895 Arhopala similis Druce, 1895 anthelus group (subgenus Narathura if valid) Arhopala achelous (Hewitson, 1862) Arhopala antharita Grose-Smith, 1894 Arhopala anthelus – anthelus bushblue Arhopala auxesia Arhopala aedias (Hewitson, 1862) (tentatively placed here) Arhopala hypomuta (Hewitson, 1862) (tentatively placed here) camdeo group camdeo subgroup Arhopala anarte (Hewitson, 1862) – magnificent oakblue Arhopala belphoebe Doherty, 1889 Arhopala camdana Corbet, 1941 Arhopala camdeo (Moore, [1858]) – lilac oakblue Arhopala dispar Riley & Godfrey, 1921 Arhopala hayashihisakazui Seki & Treadaway, 2013 Arhopala hellada Fruhstorfer, 1914 Arhopala johoreana Corbet, 1941 Arhopala opalina – opal oakblue Arhopala semperi Bethune-Baker, 1896 Arhopala varro Fruhstorfer, 1914 myrzala subgroup Arhopala bazaloides – Tamil oakblue Arhopala myrzala (Hewitson, 1869) Arhopala myrzalina Corbet, 1941 oenea group Arhopala allata Arhopala allata suffusa – Tytler's rosy oakblue Arhopala khamti Doherty, 1891 – Khamti oakblue, Doherty's dull oakblue Arhopala oenea (Hewitson, 1869) – Hewitson's oakblue epimuta group Arhopala atosia (Hewitson, [1863]) Arhopala epimuta (Moore, [1858]) Arhopala lurida Corbet, 1941 amphimuta group amphimuta subgroup Arhopala alica (Evans, 1957) Arhopala amphimuta (C. & R.Felder, 1860) Arhopala avathina Corbet, 1941 Arhopala baluensis Bethune-Baker, 1904 Arhopala dajagaka Bethune-Baker, 1896 Arhopala delta (Evans, 1957) Arhopala inornata (C. & R.Felder, 1860) (tentatively placed here) Arhopala kurzi (Distant, 1885) Arhopala major (Staudinger, 1889) Arhopala moolaiana (Moore, [1879]) Arhopala norda (Evans, 1957) Arhopala sceva Bethune-Baker, 1903 Arhopala stubbsi Eliot, 1962 Arhopala zylda Corbet, 1941 muta subgroup Arhopala moorei Bethune-Baker, 1896 Arhopala muta (Hewitson, 1862) Arhopala tropaea Corbet, 1941 perimuta subgroup (subgenus Darasana if valid) Arhopala antimuta C. & R.Felder, [1865] Arhopala metamuta (Hewitson, [1863]) Arhopala perimuta (Moore, [1858]) – yellow-disk tailless oakblue agesilaus group Arhopala agesilaus (Staudinger, 1889) Arhopala avatha de Nicéville, [1896] Arhopala democritus (Fabricius, 1793) (tentatively placed here) alitaeus group Arhopala aida de Nicéville, 1889 Arhopala alitaeus Arhopala ariana (Evans, [1925]) Arhopala arianaga Corbet, 1941 Arhopala denta (Evans, 1957) Arhopala elopura Druce, 1894 Arhopala havilandi Bethune-Baker, 1896 Arhopala pseudomuta (Staudinger, 1889) Arhopala sintanga Corbet, 1948

agrata group Arhopala ace de Nicéville, [1893] – Tytler's oakblue Arhopala agrata – de Niceville's oakblue Arhopala azinis de Nicéville, [1896] cleander group cleander subgroup Arhopala aruana (Evans, 1957) Arhopala ate (Hewitson, 1863) Arhopala athada (Staudinger, 1889) Arhopala cleander (C.Felder, 1860) Arhopala silhetensis – Sylhet oakblue Arhopala zambra Swinhoe, [1911] alea subgroup Arhopala aenigma Eliot, 1972 Arhopala agaba – purple-glazed oakblue Arhopala alea – rosy oakblue Arhopala aroa (Hewitson, [1863]) Arhopala aurelia (Evans, [1925]) Arhopala evansi Corbet, 1941 Arhopala milleri Corbet, 1941 Arhopala normani Eliot, 1972 Arhopala phaenops C. & R.Felder, [1865] (including A. detrita) Arhopala phanda Corbet, 1941 Arhopala ralanda Corbet, 1941 Arhopala selta (Hewitson, 1869) Arhopala sublustris Bethune-Baker, 1904 Arhopala vihara (C. & R. Felder, 1860) (tentatively placed here) eumolphus group Arhopala acta (Evans, 1957) Arhopala asma Arhopala bazalus – powdered oakblue Arhopala chamaeleona Bethune-Baker, 1903 Arhopala corinda (Hewitson, 1869) (tentatively placed here) Arhopala critala Arhopala eumolphus – green oakblue Arhopala florinda Arhopala hellenore Doherty, 1889 – Doherty's green oakblue Arhopala horsfieldi Arhopala irma Fruhstorfer, 1914 Arhopala nobilis (Felder, 1860) Arhopala overdijkinki Corbet, 1941 Arhopala pagenstecheri Arhopala tameanga Bethune-Baker, 1896 Arhopala wildei rama group Arhopala arvina (Hewitson, [1863]) – purple-brown tailless oakblue Arhopala buddha Bethune-Baker, 1903 Arhopala paralea – glazed oakblue Arhopala rama – dark Himalayan oakblue agelastus group Arhopala agelastus (Hewitson, 1862) Arhopala alaconia (Hewitson, 1869) Arhopala alesia – pallid oakblue Arhopala barami Bethune-Baker, 1903 Arhopala cardoni Corbet, 1941 Arhopala epimete (Staudinger, 1889) Arhopala labuana Bethune-Baker, 1896 Arhopala wildeyana Corbet, 1941 fulla group Arhopala disparilis Arhopala fulla (Hewitson, 1862) – spotless oakblue aurea group (subgenus Aurea if valid) Arhopala aurea (Hewitson, 1862) Arhopala borneensis Bethune-Baker, 1896 Arhopala caeca (Hewitson, [1863]) Arhopala stinga (Evans, 1957) Arhopala trogon (Distant, 1884) ganesa group (subgenus Panchala if valid) Arhopala ammon – Malayan oakblue Arhopala ammonides (Doherty, 1891) Arhopala ariel (Doherty, 1891) Arhopala elizabethae (Eliot, 1959) Arhopala ganesa – tailless bushblue Arhopala paraganesa (de Nicéville, 1882) – dusky bushblue Arhopala tomokoae (H. Hayashi, 1976) abseus group Arhopala abseus – aberrant oakblue, aberrant bushblue Arhopala anella de Nicéville, [1895] thamyras group Arhopala arta Arhopala axina Arhopala axiothea Arhopala doreena Arhopala helianthes Arhopala thamyras (Linnaeus, 1764) hercules group Arhopala hercules (Hewitson, 1862) Arhopala herculina Staudinger, 1888 Arhopala leo Arhopala sophilus Arhopala tyrannus

Incertae sedis

Male Burmese bushblue (Arhopala birmana)

Hooked oakblue (Arhopala paramuta)

Arhopala aberrans (de Nicéville, [1889]) – pale bushblue Arhopala acetes Arhopala aeeta de Nicéville, [1893] Arhopala alax – silky oakblue Arhopala alexandrae Schröder & Treadaway, 1978 Arhopala anamuta Semper, 1890 Arhopala annulata (Felder, 1860) Arhopala argentea Staudinger, 1888 Arhopala aronya (Hewitson, 1869) Arhopala arsenius (C. & R.Felder, [1865]) Arhopala asinarus C. & R.Felder, [1865] Arhopala asopia (Hewitson, [1869]) – plain tailless oakblue Arhopala atrax – dark broken-band oakblue, Indian oakblue (type species of Satadra) Arhopala bella Bethune-Baker, 1896 Arhopala birmana – Burmese bushblue Arhopala brooksiana Corbet, 1941 Arhopala canaraica (Moore, 1884) Arhopala comica de Nicéville, 1900 – comic oakblue Arhopala curiosa (Evans, 1957) Arhopala davaona Semper, 1890 Arhopala dodonaea – pale Himalayan oakblue Arhopala dohertyi Bethune-Baker, 1903 Arhopala ellisi Evans, 1914 – Ellis's bushblue Arhopala eridanus Arhopala grandimuta Seki, 1993 Arhopala halma Arhopala halmaheira Arhopala hellenoroides Chou & Gu, 1994 Arhopala hesba (Hewitson, 1869) Arhopala hinigugma Takanami, 1985 Arhopala hylander Arhopala ilocana Osada & Hashimoto, 1987 Arhopala irregularis Bethune-Baker, 1903

Arhopala japonica – Japanese oakblue Arhopala lata (Evans, 1957) Arhopala luzonensis Takanami & Ballantine, 1987 Arhopala matsutaroi (H. Hayashi, 1979) Arhopala mindanensis Arhopala mizunumai Arhopala myrtha (Staudinger, 1889) Arhopala nakamotoi Arhopala nicevillei Bethune-Baker, 1903 Arhopala ocrida (Hewitson, 1869) Arhopala ormistoni – Ormiston's oakblue Arhopala pabihira Arhopala paramuta – hooked oakblue Arhopala phryxus Boisduval, 1832 Arhopala pseudovihara (H. Hayashi, 1981) Arhopala qiongdaoensis Chou & Gu, 1994 Arhopala quercoides Arhopala rudepoema Seki, [1995] Arhopala sakaguchii (H. Hayashi, 1981) Arhopala sangira Bethune-Baker, 1897 Arhopala schroederi Arhopala siabra Arhopala simoni Schröder & Treadaway, 1999 Arhopala singla (de Nicéville, 1885) – yellow-disk oakblue Arhopala staudingeri Semper, 1890 Arhopala straatmani Arhopala tephlis Arhopala theba (Hewitson, [1863]) Arhopala tindongani Nuyda & Takanami, 1990 Arhopala trionaea Semper, 1890 (camdeo group?) Arhopala weelii Arhopala zeta

Footnotes

1. Brower (2008)
2. Brower (2008), and see Savela (2008) for additional references

References

• Brower, Andrew V. Z. (2008): Tree of Life Web Project – Arhopala. Version of 2008-APR-09. Retrieved 2009-MAR-21.
• Savela, Markku (2008): Markku Savela's Lepidoptera and Some Other Life Forms – Arhopala. Version of 2008-AUG-06. Retrieved 2009-MAR-21.
• Evans, W H, 1957 A revision of the Arhopala group of Oriental Lycaenidae (Lepidoptera: Rhopalocera) Bulletin of the British Museum (Natural History), London, Entomology Volume 5 : 85- 141 online

External links

• Images representing Arhopala at Consortium for the Barcode of Life