Long-tailed skipper Urbanus proteus, copyright Andreas Kay.

Belongs within: Obtectomera.
Contains: Hasora, Pyrginae, Toxidia, Hesperilla, TrapezitesPamphila, Hesperiinae.

The Hesperiidae, skippers, are a group of butterfly-like Lepidoptera with an erratic, quick flight. They have their antennal bases spaced relatively far apart and the antennal club usually has a small apical hook. The fore wing lacks forked peripheral veins (Braby 2000). Many skippers are dully coloured and species may be very difficult to distinguish.

Several subfamilies are currently recognised, including the Megathyminae, giant skippers, of arid regions of North America that have sometimes been recognised as a distinct family. The Trapezitinae of Australasia are relatively small, stout-bodied skippers whose larvae feed on monocotyledons such as grasses or sedges. The Coeliadinae, awls, are relatively large, robust hesperiids found in the Afrotropical, Oriental and Australian regions. Coeliadine larvae are brightly coloured and feed on dicotyledons, particularly Leguminosae (Braby 2000). Urbanus is a Neotropical genus of skippers with a long tail extending from each of the hind wings.

Characters (from Braby 2000): Adults small to medium-sized, head broad, body stout, wings relatively short, usually sombre brown but sometimes brightly coloured; antennae widely separated at base, apiculus of club curved or sharply bent with tip blunt or pointed; fore wing with five branches on radial vein, all arising separately from cell, and one anal vein; hind wing with humeral vein usually present, two anal veins, tornus usually rounded or sometimes slightly produced but rarely tailed; both sexes with six normal functional legs, tarsi with a pair of apical claws; fore tibia with epiphysis on inner side. Egg hemispherical or dome-shaped, smooth but often with series of longitudinal ribs. Final instar larva cylindrical, tapering at each end, finely spinulose with numerous secondary setae; head comparatively large, larger than prothorax; osmeterium absent; crochets of prolegs in a circle; anal comb usually present. Pupa usually concealed inside larval shelter, attached by cremaster and sometimes also by central girdle.

<==Hesperiidae [Hesperioidea]
    |  `--Megathymus [Megathyminae, Megathymidae]KP19
    |       `--M. yuccaeKP19
    `--Urbanus [Eudamiinae]KP19
         |--U. dorantes (Stoll 1791)FS54
         |    |--U. d. dorantesPP72
         |    `--U. d. galapagensisPP72
         |--U. eurycles (Latreille 1723)FS54
         |    |--U. e. euryclesFS54
         |    `--U. e. latipennis (Marille & Vuillot 1891)FS54
         |--U. gracilicauda (Plötz 1880)FS54
         |--U. proteus (Linnaeus 1758)FS54
         `--U. undulatus (Hewitson 1867)FS54

Hesperiidae incertae sedis:
  Phareas Westwood 1852 [=Phareus (l. c.) non Simon 1879]K03
  Thymele fulminans Herrich-Schäffer 1869FS54
  Nascus phocus (Cramer 1779)S55
  Cogia calchas (Herrich-Schäffer 1869)FS54
  Spathilepia clonius (Cramer 1779)FS54
  Phanus vitreus (Cramer 1782)S55
    |--P. bromias (Godman & Salvin 1894)FS54
    `--P. polyctor (Prittwitz 1868)S55
    |--M. brasilicola (Strand 1920)S55
    `--M. lassia (Hewitson 1868)FS54
  Antigonus erosus (Hübner 1820)FS54
  Copaeodes aurantiaca (Hewitson 1868)FS54
  Calpodes ethlius (Cramer 1782)FS54
  Panoquina vala (Mabille 1891)FS54
    |--C. carbo Mabille 1890M90b
    |--C. corvinus Mabille 1890M90b
    |--C. debilisM90b
    |--C. duplex Mabille 1890M90b
    |--C. subcordataFS54
    `--C. subfacatus Mabille 1890M90b
  PyrrhopygeGE05 [PyrrhopyginaeB2000]
    |--P. creonGE05
    `--P. josephina Draudt 1921S55
  Mysoria thasus (Cramer 1782)S55
  Jemadia suzetta Mabille & Boullet 1908S55
    |--G. decurtatus Herrich-Schäffer 1869S55
    |--G. eurycles Latreille 1823S55
    |--G. leucogramma Mabille 1888S55
    `--G. undulatus Hewitson 1867S55
  Chioides catillus (Cramer 1782)S55
    |--T. creteus (Cramer 1782)S55
    `--T. mephitis Hewitson 1876S55
  Paches loxus (Westwood & Hewitson 1852)S55
  Eudamidas ozema (Butler 1870)S55
  Ebrietas infanda (Butler 1877)S55
    |--T. athenion (Hübner 1816)S55
    `--T. lineolus Ochsenheimer 1808B08
  Carystus ebusa (Cramer 1782)S55
  Pyrrhopygopsis cleanthes (Latreille 1823)S55
  Elbella etnaGE05
  Pamphilites abditaGE05
  Erionota thraxMHG04, VHK02
    |--P. icteriaB01a
    `--P. numa Druce 1901D01
    |--A. biguttulus Mabille 1890M90b
    |--A. instabilis Mabille 1890M90b
    |--A. leucopygaM90b
    |--A. mackeniiB01a
    |--A. placidusB01a
    `--A. ploetzi Mabille 1890M90b
  Andronymus philanderB01a
  Kedestes wallengreniiB01a
    |--C. punctulataB01a
    |--C. quaterna Mabille 1890M90a
    `--C. stellataB01a
  Eretis lugensB01b
  Gomalia elmaB01b
  Cyclopides quadrisignataB01b
  Padraona zenoB01b
  Gegenes letterstedtiB01b
  Rhopalocampta forestanB01b
    |--Allora Waterhouse & Lyell 1914B2000
    |    |--A. doleschallii (Felder 1860)B2000
    |    `--A. major (Rothschild 1915)B2000
    |    |--I. jainaS72
    |    `--I. ladonWM66
    `--Badamia Moore 1881B2000
         |--B. atrox (Butler 1877)B2000
         |    |--B. a. atroxPP02
         |    `--B. a. subflavaPP02
         `--B. exclamationis (Fabricius 1775)B12
    |--Felicina Waterhouse 1932B2000
    |--Hewitsoniella Shepard 1931B2000
    |--Rachelia Hemming 1964B2000
    |    `--R. extrusus (Felder & Felder 1867)B2000
    |--Pasma Waterhouse 1932B2000
    |    `--P. tasmanicus (Miskin 1889)B2000
    |--Dispar Waterhouse & Lyell 1914B2000
    |    `--D. compacta (Butler 1882)B2000
    |--Proeidosa Atkins 1973B2000
    |    `--P. polysema (Lower 1908)B12
    |--Signeta Waterhouse & Lyell 1914B2000
    |    |--S. flammeata (Butler 1882)B2000
    |    `--S. tymbophora (Meyrick & Lower 1902)B2000
    |--Croitana Waterhouse 1932B2000
    |    |--C. aestiva Edwards 1979B2000
    |    |--C. arenaria Edwards 1979B2000
    |    `--C. croites (Hewitson 1874) [incl. C. croites pindar (Waterhouse 1932)]B2000
    |--Neohesperilla Waterhouse & Lyell 1914B2000
    |    |--N. croceus (Miskin 1889)B2000
    |    |--N. senta (Miskin 1891)B2000
    |    |--N. xanthomera (Meyrick & Lower 1902)B2000
    |    `--N. xiphiphora (Lower 1911)B2000
    |--Oreisplanus Waterhouse & Lyell 1914B2000
    |    |--O. munionga (Olliff 1890)B2000
    |    |    |--O. m. muniongaB2000
    |    |    `--O. m. larana Couchman 1962B2000
    |    `--O. perornata (Kirby 1893)B2000
    |--Herimosa Atkins 1994B2000
    |    `--H. albovenata (Waterhouse 1940) [=Anisynta albovenata]B2000
    |         |--H. a. albovenataB2000
    |         |--H. a. fuscata (Parson 1965)B2000
    |         `--H. a. weemala (Couchman 1954)B2000
    |--Mesodina Meyrick 1901B2000
    |    |--M. aeluropis Meyrick 1901B2000
    |    |--M. cyanophracta Lower 1911B2000
    |    |--M. gracillima Edwards 1987B2000
    |    |--M. halyzia (Hewitson 1868)B2000
    |    `--M. hayi Edwards & Graham 1995B2000
    |--Motasingha Watson 1893B2000
    |    |--M. dirphia (Hewitson 1868)B2000
    |    |--M. dominulaP27
    |    `--M. trimaculata (Tepper 1882)B2000
    |         |--M. t. trimaculataB2000
    |         |--M. t. dilata Waterhouse 1932 [incl. M. trimaculata dea Waterhouse 1933]B2000
    |         `--M. t. occidentalis Moulds & Atkins 1986B2000
    |--Antipodia Atkins 1984B2000
    |    |--A. atralba (Tepper 1882)B2000
    |    |--A. chaostola (Meyrick 1888) [=Hesperilla chaostola]B2000
    |    |    |--A. c. chaostolaB2000
    |    |    |--A. c. chares (Waterhouse 1933)B2000
    |    |    `--A. c. leucophaea (Couchman 1946)B2000
    |    `--A. dactyliota (Meyrick 1888) [=A. atralba dactyliota]B2000
    |         |--A. d. dactyliotaB2000
    |         `--A. d. nila (Waterhouse 1932)B2000
    `--Anisynta Lower 1911B2000
         |--A. cynone (Hewitson 1874)B2000
         |    |--A. c. cynone [incl. A. cynone gracilis (Tepper 1882), A. cynone grisea Waterhouse 1932]B2000
         |    `--A. c. gunneda Couchman 1954B2000
         |--A. dominula (Plötz 1884)B2000
         |    |--A. d. dominula (see below for synonymy)B2000
         |    `--A. d. pria (Waterhouse 1932)B2000
         |--A. monticolae (Olliff 1890)B2000
         |--A. sphenosema (Meyrick & Lower 1902)B2000
         `--A. tillyardi Waterhouse & Lyell 1912B2000
  Oxypalpus rusoB1900
    |--B. dognini Mabille 1889M89b
    |--B. polydesma Mabille 1889M89b
    |--B. quadristiga Mabille 1889M89b
    `--B. riza Mabille 1889M89b
  Steropes furvus Mabille 1890M90a
  ‘Choristoneura’ Mabille 1890 non Lederer 1859M90a
    `--*C. apicalis Mabille 1890M90a
  Chetoneura callixenusM90a
  Hyda Mabille 1890M90c
    `--*H. micacea Mabille 1890M90c
  Stethotrix Mabille 1890M90c
    `--*S. heterogyna Mabille 1890M90c
  Dis Mabille 1890M90c
    `--*D. annulatus Mabille 1890M90c

Anisynta dominula dominula [incl. A. dominula drachmophora (Meyrick 1885), A. dominula draco Waterhouse 1938, A. dominula dyris Waterhouse 1938]B2000

*Type species of generic name indicated


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[B2000] Braby, M. F. 2000. Butterflies of Australia: their identification, biology and distribution vol. 1. CSIRO Publishing: Collingwood (Victoria).

[B12] Braby, M. F. 2012. The butterflies of El Questro Wilderness Park, with remarks on the taxonomy of the Kimberley fauna, Australia. Records of the Western Australian Museum 27 (2): 161–175.

[B1900] Butler, A. G. 1900. On a second collection of butterflies obtained by Mr. Edward M. de Jersey in Nyasaland. Annals and Magazine of Natural History, series 7, 5: 59–63.

[B01a] Butler, A. G. 1901a. An account of a collection of butterflies made by the Rev. K. St. Aubyn Rogers between Mombasa and the forests of Taveta. Annals and Magazine of Natural History, series 7, 7: 22–35.

[B01b] Butler, A. G. 1901b. An account of a collection of butterflies obtained by Lord Delamere, chiefly at Munisu, near Mount Kenya. Annals and Magazine of Natural History, series 7, 7: 197–203.

[D01] Druce, H. 1901. Descriptions of some new species of Lepidoptera from east Africa and tropical America. Annals and Magazine of Natural History, series 7, 7: 432–444.

[E66] Evans, H. E. 1966. The Comparative Ethology and Evolution of the Sand Wasps. Harvard University Press: Cambridge (Massachusetts).

[FS54] Franz, E., & H. Schröder. 1954. Tagfalter (Lep. Rhopalocera) aus El Salvador. Senckenbergiana Biologica 35: 75–87.

[GE05] Grimaldi, D., & M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press: New York.

[J03] Jong, R. de. 2003. Are there butterflies with Gondwanan ancestry in the Australian region? Invertebrate Systematics 17: 143–156.

[KP19] Kawahara, A. Y., D. Plotkin, M. Espeland, K. Meusemann, E. F. A. Toussaint, A. Donath, F. Gimnich, P. B. Frandsen, A. Zwick, M. dos Reis, J. R. Barber, R. S. Peters, S. Liu, X. Zhou, C. Mayer, L. Podsiadlowski, C. Storer, J. E. Yack, B. Misof & J. W. Breinholt. 2019. Phylogenomics reveals the evolutionary timing and pattern of butterflies and moths. Proceedings of the National Academy of Sciences of the USA 116 (45): 22657–22663.

[K03] Kury, A. B. 2003. Annotated catalogue of the Laniatores of the New World (Arachnida, Opiliones). Revista Ibérica de Aracnología, volumen especial monográfico 1: 1–337.

[M89a] Mabille, P. 1889a. Description de quelques hespérides nouvelles du genre Pamphila. Annales de la Société Entomologique de France, 6e série 9: lxxxiv–lxxxv.

[M89b] Mabille, P. 1889b. Descriptions de quelques espèces nouvelles d’hespérides du genre Butleria. Annales de la Société Entomologique de France, 6e série 9: xci–xciii.

[M90a] Mabille, P. 1890a. Lépidoptères (hespérides) nouveaux (2e partie). Annales de la Société Entomologique de France, 6e série 9: clv–clvi.

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[M90c] Mabille, P. 1890c. Lépidoptères (hespérides) nouveaux (4e partie). Annales de la Société Entomologique de France, 6e série 9: clxxxiii–clxxxiv.

[MHG04] Maynard, G. V., J. G. Hamilton & J. F. Grimshaw. 2004. Quarantine—phytosanitary, sanitary and incursion management: an Australian entomological perspective. Australian Journal of Entomology 43 (3): 318–328.

[PP72] Parkin, P., D. T. Parkin, A. W. Ewing & H. A. Ford. 1972. A report on the arthropods collected by the Edinburgh University Galapagos Islands expedition, 1968. Pan-Pacific Entomologist 48 (2): 100–107.

[PP02] Patrick, B., & H. Patrick. 2002. Butterflies of Fiji. Weta 24: 5–12.

[P27] Philpott, A. 1927. The maxillae in the Lepidoptera. Transactions and Proceedings of the New Zealand Institute 57: 721–746.

[S55] Schröder, H. 1955. Eine Falter-Ausbeute aus dem westlichen Bolivien. (Ins. Lepid. Rhopal.) Senckenbergiana Biologica 36: 329–338.

[S72] Shields, O. 1972. Flower visitation records for butterflies (Lepidoptera). Pan-Pacific Entomologist 48 (3): 189–203.

[VHK02] Verma, M., M. Hayat & S. I. Kazmi. 2002. The species of Elasmus from India (Hymenoptera: Chalcidoidea: Eulophidae). Oriental Insects 36: 245–306.

[WM66] Wallace, A. R., & F. Moore. 1866. List of lepidopterous insects collected at Takow, Formosa, by Mr. R. Swinhoe. Proceedings of the Zoological Society of London 1866: 355–365.

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