Hesperiidae

Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Protostomia
Cladus: Ecdysozoa
Cladus: Panarthropoda
Phylum: Arthropoda
Subphylum: Hexapoda
Classis: Insecta
Cladus: Dicondylia
Subclassis: Pterygota
Cladus: Metapterygota
Infraclassis: Neoptera
Cladus: Eumetabola
Cladus: Endopterygota
Superordo: Panorpida
Cladus: Amphiesmenoptera
Ordo: Lepidoptera
Subordo: Glossata
Cladus: Coelolepida
Cladus: Myoglossata
Cladus: Neolepidoptera
Infraordo: Heteroneura
Cladus: Eulepidoptera
Cladus: Ditrysia
Cladus: Apoditrysia
Cladus: Obtectomera
Superfamilia: Papilionoidea

Familia: Hesperiidae
Subfamiliae (12, per Zhang et al., 2019):
Barcinae – Chamundinae – Coeliadinae – Eudaminae – Euschemoninae – Hesperiinae – Heteropterinae – Katreinae – Megathyminae – Pyrginae – Pyrrhopyginae – Tagiadinae – Trapezitinae

Overview of genera

Abantis – Abraximorpha – Acada – Acerbas – Achlyodes – Acleros – Actinor – Adlerodea – Adopaeoides – Aegiale – Aeromachus – Aethilla – Afrogegenes – Agara – Agathymus – Aguna – Agyllia – Aides – Albiphasma – Alenia – Allora – Alychna – Amblyscirtes – Amenis – Ampittia – Amysoria – Anastrus – Anatrytone – Anaxas – Ancistroides – Ancyloxypha – Andronymus – Anisynta – Anisyntoides – Ankola – Anthoptus – Apallaga – Apaustus – Apostictopterus – Appia – Apyrrhothrix – Ardaris – Argemma – Argon – Argopteron – Arnetta – Aroma – Arotis – Arrhenella – Arrhenes – Arteurotia – Artines – Artitropa – Artonia – Asbolis – Aspitha – Astictopterus – Astraptes – Atalopedes – Atarnes – Atrytone – Atrytonopsis – Augiades – Aurina – Aurivittia – Autochton – Austinus – Avestia – Azonax – Badamia – Banta – Baoris – Baracus – Barca – Barrolla – Bibasis – Bolla – Borbo – Bruna – Bralus – Brownus – Brusa – Bungalotis – Burara – Burca – Burmaozephyrus – Burnsius – Butleria – Buzella – Buzyges – Cabirus – Caenides – Caligulana – Calleagris – Callimormus – Calpodes – Caltoris – Camptopleura – Canesia – Cantha – Capila – Caprona – Carcharodus – Carterocephalus – Carystina – Carystoides – Carystus – Cecropterus – Celaenorrhinus – Cephise – Cephrenes – Ceratrichia – Ceratricula – Cerba – Chaetocneme – Chalcone – Chalypyge – Chamunda – Charidia – Chioides – Chiomara – Chiothion – Chirgus – Chitta – Chloeria – Choaspes – Chondrolepis – Choranthus – Clito – Clytius – Cobaloides – Cobalopsis – Cobalus – Codatractus – Coeliades – Cogia – Coladenia – Conga – Conognathus – Coolus – Copaeodes – Corra – Corticea – Cravera – Crenda – Creonpyge – Creteus – Crinifemur – Croniades – Ctenoptilum – Cumbre – Cupitha – Cyanopyge – Cycloglypha – Cyclopyge – Cyclosemia – Cyclosma – Cymaenes – Cynea – Daimio – Dalla – Damas – Dardarina – Daron – Darpa – Decinea – Doberes – Dion – Dotta – Drephalys – Dubia – Dubiella – Duroca – Dyscophellus – Eagris – Eantis – Ebrietas – Eburuncus – Ebusus – Echelatus – Ectomis – Eetion – Elbella – Emmelus – Enosis – Entheus – Eogenes – Epargyreus – Ephyriades – Eretis – Erionota – Ernsta – Erynnis – Euphyes – Euriphellus – Euschemon – Eutocus – Eutychide – Evansiella – Exometoeca – Falga – Favria – Festivia – Fidius – Flaccilla – Flandria – Flattoides – Freemaniana – Fresna – Fulda – Galerga – Gallio – Gamia – Gangara – Ge – Gegenes – Gerosis – Gesta – Gindanes – Ginungagapus – Gomalia – Gorgopas – Gorgyra – Gorgythion – Grais – Granila – Gretna – Gunayan – Gyrogra – Haemactis – Halotus – Halpe – Halpemorpha – Hansa – Hasora – Haza – Helias – Heliopetes – Herila – Hesperia – Hesperilla – Hesperopsis – Heteropterus – Hidari – Hollandus – Holguinia – Hoodus – Hovala – Hyalothyrus – Hyarotis – Hylephila – Hypoleucis – Iambrix – Idmon – Igapophilus – Iliana – Ilma – Incisus – Inglorius – Isma – Ismene – Isoteinon – Iton – Jemadia – Jera – Joanna – Jonaspyge – Jongiana – Justinia – Katreus – Kedestes – Kobrona – Koruthaialos – Ladda – Lamponia – Larsenia – Leona – Lento – Lepella – Leptalina – Lerema – Lerodea – Leucochitonea – Libra – Librita – Lindra – Lobocla – Lobotractus – Lissia – Livida – Lon – Lotongus – Loxolexis – Lucida – Ludens – Lurida – Lycas – Lychnuchoides – Lychnuchus – Marela – Malaza – Matapa – Megaleas – Megathymus – Melanopyge – Melphina – Melphinyet – Metardaris – Methion – Methionopsis – Metisella – Metron – Meza – Microceris – Mictris – Mielkeus – Milanion – Miltomiges – Mimardaris – Mimene – Mimia – Mimoniades – Miraja – Misius – Mnaseas – Mnasicles – Mnasilus – Mnasinous – Mnasitheus – Mnestheus – Moeris – Moeros – Molla – Molo – Moltena – Monca – Monza – Mooreana – Mopala – Morvina – Morys – Mucia – Muschampia – Mylon – Myrinia – Mysarbia – Myscelus – Mysoria – Naevolus – Narcosius – Nascus – Nastra – Neochlodes – Neohesperilla – Neomorphuncus – Neoxeniades – Neposa – Nerula – Nervia – Netrobalane – Netrocoryne – Nicephellus – Niconiades – Nisoniades – Noctuana – Noctulana – Nosphistia – Notamblyscirtes – Notocrypta – Nyctelius – Nyctus – Oarisma – Ocella – Ochlodes – Ochropyge – Ocybadistes – Odina – Odontoptilum – Oechydrus – Oenides – Oeonus – Oerane – Oileides – Oligoria – Onenses – Onespa – Onophas – Onryza – Oreisplanus – Oriens – Orphe – Orses – Ortholexis – Orthos – Osmodes – Osphantes – Ouleus – Oxynetra – Oxynthes – Pachyneuria – Pamba – Panca – Panoquina – Papias – Paracarystus – Parachoranthus – Paracleros – Paramimus – Parasovia – Paratrytone – Pardaleodes – Parelbella – Pares – Parnara – Paronymus – Parosmodes – Parphorus – Passova – Pastria – Peba – Pedesta – Pellicia – Pelopidas – Pemara – Penicula – Perichares – Perrotia – Perus – Phanes – Phanus – Phareas – Phemiades – Pheraeus – Phlebodes – Phocides – Pholisora – Pintara – Pirdana – Piruna – Pithauria – Plastingia – Platylesches – Ploetzia – Poanes – Polites – Polyctor – Polygonus – Polytremis – Pompeius – Porphyrogenes – Potamanaxas – Potanthus – Prada – Praescobura – Praethoressa – Problema – Procampta – Propapias – Propertius – Prosopalpus – Protelbella – Proteides – Prusiana – Pseudoborbo – Pseudocoladenia – Pseudocopaeodes – Pseudocroniades – Pseudodrephalys – Pseudokerana – Pseudonascus – Pseudopirdana – Pseudorphe – Pseudosarbia – Psolos – Psoralis – Pteroteinon – Pudicitia – Punta – Pyroneura – Pyrrhocalles – Pyrrhochalcia – Pyrrhopyge – Pyrrhopygopsis – Pythonides – Quadrus – Quasimellana – Quedara – Quinta – Racta – Radiatus – Ralis – Remella – Repens – Rhabdomantis – Rhinthon – Ridens – Rigga – Rufocumbre – Sabera – Sacrator – Salanoemia – Salantoia – Salatis – Saniba – Santa – Sarbia – Sarmientoia – Satarupa – Saturnus – Scobura – Sebastonyma – Semalea – Semperium – Serdis – Seseria – Sodalia – Sophista – Sostrata – Sovia – Spathilepia – Spialia – Spicauda – Stallingsia – Staphylus – Stimula – Stinga – Styriodes – Suada – Suastus – Sucova – Suniana – Synale – Synapte – Tagiades – Tapena – Talides – Taractrocera – Tarsoctenus – Tava – Tekliades – Telegonus – Telemiades – Telicota – Telles – Tellona – Teniorhinus – Testia – Thargella – Theagenes – Thespieus – Thoon – Thoressa – Thracides – Thymelicus – Tiacellia – Tiana – Tigasis – Timochares – Tirynthia – Tirynthoides – Tisias – Tosta – Trapezites – Tricrista – Triskelionia – Tromba – Troyus – Tsitana – Tsukiyamaia – Turesis – Turmosa – Turnerina – Typhedanus – Udaspes – Udranomia – Unkana – Urbanus – Vacerra – Veadda – Vehilius – Venada – Venas – Vernia – Vertica – Vettius – Vidius – Vinpeius – Viola – Virga – Viridina – Viuria – Wahydra – Wallengrenia – Willema – Windia – Xanthodisca – Xanthoneura – Xanthonymus – Xeniades – Xispia – Yanguna – Yania – Yanoancistroides – Zalomes – Zariaspes – Zela – Zenis – Zenonia – Zenonoida – Zera – Zestusa – Zetka – Zeutus – Zinaida – Zobera – Zographetus – Zonia – Zophopetes – Zopyrion
Name

Hesperiidae Latreille, 1809.
Synonymy

Hesperiidae Latreille, 1809: 187, 207.
Hesperiadae Doherty, 1886: 111; Meyrick, E. & Lower, O., 1902: 38.
Megathymidae Comstock & Comstock, 1895: 365; Freeman, [1969]: 2.

References
Primary references

Bridges, C.A. 1994. Catalogue of the Family-Group, Genus-Group and Species-Group Names of the Hesperioidea (Lepidoptera) of the World. Bridges, Urbana, Illinois. BHL Reference page.
Comstock, J.H. & Comstock, A.B. 1895. A manual for the study of insects. Comstock Publishing Company, Ithaca, N.Y. 701pp. Reference page.
Cong, Q., Zhang, J., Shen, J.H. & Grishin, N.V. 2019. Fifty new genera of Hesperiidae (Lepidoptera). Insecta Mundi 0731: 1–56. Zoobank. Reference page.
Doherty, W., 1886. A list of butterflies taken in Kumaon. J. Asiat. Soc. Beng. 55(2): 103–140. BHL. Reference page.
Freeman, H.A., 1959. A revision of the genera of the Megathymidae, with the description of Three new genera. The Lepidopterists' News 12(3-4): 81-92. (26 Jan 1959) Reference page.
Latreille, P.A. 1809. Genera crustaceorum et insectorum secundum ordinem naturalem in familias disposita, iconibus exemplisque plurimis explicata. Tomus quartus et ultimus. Parisiis: A. Koenig, 399 pp. BHL Reference page.
Li, W., Cong. Q., Shen, J.H., Zhang, J., Hallwachs, W., Janzen, D.H. & Grishin, N.V. 2019. Genomes of skipper butterflies reveal extensive convergence of wing patterns. PNAS 116(13): 6232–6237. DOI: 10.1073/pnas.1821304116. Text. SI Appendix. Reference page.
Warren, A.D. 2006. The Higher Classification of the Hesperiidae. Oregon State University. PDF. Reference page.
Warren, A.D., Ogawa, J.R. & Brower, A.V.Z. 2009. Revised classification of the family Hesperiidae (Lepidoptera: Hesperioidea) based on combined molecular and morphological data. Systematic entomology 34(3): 467–523. Reference page.
Warren, B.C.S., 1926. Monograph of the Tribe Hesperiidi (European Species) with Revised Classification of the Subfamily Hesperiinae (Palaearctic Species) Based on the Genital Armature of the Males. Transactions of the Entomological Society of London 74(1): 1–167. DOI: 10.1111/j.1365-2311.1926.tb01184.x Reference page.
Zhang, J., Cong. Q., Shen, J.H., Brockmann, E. & Grishin, N.V. 2019. Three new subfamilies of skipper butterflies (Lepidoptera, Hesperiidae). ZooKeys 861: 91–105. DOI: 10.3897/zookeys.861.34686 Reference page.

Additional references

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Mielke, O.H.H., Carneiro, E. & Casagrande, M.M. 2012. Hesperiidae (Lepidoptera, Hesperioidea) from Ponta Grossa, Paraná, Brazil: 70 years of records with special reference to faunal composition of Vila Velha State Park. Revista Brasileira de Entomologia 56(1): 59–66. Reference page.
Miller, L.D. 1971. Descriptions of new species and notes on other Hesperiidae of Africa. Bulletin of the Allyn Museum 2: 1–17. PDF. Reference page.
Miller, L.D. & Miller, J.Y. 1972. New high-altitude Hesperiinae from Mexico and Ecuador (Hesperiidae). Bulletin of the Allyn Museum 7: 1–7. Reference page.
Sahoo, R.K., Warren, A.D., Wahlberg, N., Brower, A.V.Z., Lukhtanov, V.A. & Kodandaramaiah, U. 2016. Ten genes and two topologies: an exploration of higher relationships in skipper butterflies (Hesperiidae). PeerJ 4: e2653. DOI: 10.7717/peerj.2653 Reference page.
Sharma, S. & Sharma, N., 2020. On the rediscovery and new additions of Skippers (Lepidoptera: Papilionoidea: Hesperiidae) from Jammu and Kashmir, India. Revista Chilena de Entomología 46(4): 591-599, figa 1-9. Reference page.
Bivar de Sousa, A. & Mendes, L.F. 2007. On the genus Eagris Guenée, 1863 in Angola (Lepidoptera: Hesperiidae). Shilap Revista de Lepidopterologia 35(139): 311–316. Reference page.
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Vernacular names
беларуская: Тоўстагалоўкі
English: Skipper, Skipper Butterfly
suomi: Paksupäät
français: hespéridés
日本語: セセリチョウ科
polski: powszelatkowate
தமிழ்: தாவிகள், துள்ளிகள் (ஹெஸ்பெரிடே)
中文: 弄蝶科

Skippers are a family of the Lepidoptera (moths and butterflies) named the Hesperiidae. Being diurnal, they are generally called butterflies. They were previously placed in a separate superfamily, Hesperioidea; however, the most recent taxonomy places the family in the superfamily Papilionoidea, the butterflies. They are named for their quick, darting flight habits. Most have their antenna tips modified into narrow, hook-like projections. Moreover, skippers mostly have an absence of wing-coupling structure available in most moths.[1] More than 3500 species of skippers are recognized, and they occur worldwide, but with the greatest diversity in the Neotropical regions of Central and South America.[2]

Description and systematics
Plate from Biologia Centrali-Americana showing Pyrginae (3 at right center — black and blue-brown) and Eudaminae (the others)

Traditionally, the Hesperiidae were placed in a monotypic superfamily Hesperioidea, because they are morphologically distinct from other Rhopalocera (butterflies), which mostly belong to the typical butterfly superfamily Papilionoidea. The third and rather small butterfly superfamily is the moth-butterflies (Hedyloidea), which are restricted to the Neotropics, but recent phylogenetic analyses suggest the traditional Papilionoidea are paraphyletic, thus the subfamilies should be reorganised to reflect true cladistic relationships.[3][4]

Collectively, these three groups of butterflies share many characteristics, especially in the egg, larval, and pupal stages.[2] Nevertheless, skippers have the antennae clubs hooked backward like a crochet hook, while the typical butterflies have club-like tips to their antennae, and moth-butterflies have feathered or pectinate (comb-shaped) antennae similar to moths. Skippers also have generally stockier bodies and larger compound eyes than the other two groups, with stronger wing muscles in the plump thorax, in this resembling many moths more than the other two butterfly lineages do. Unlike, for example, the Arctiinae, though, their wings are usually small in proportion to their bodies. Some have larger wings, but only rarely as large in proportion to the body as in other butterflies. When at rest, skippers keep their wings usually angled upwards or spread out, and only rarely fold them up completely.[2]
Signeta flammeata (Trapezitinae)
Mating pair of Oberthür's Grizzled Skipper (Pyrgus armoricanus)

The wings are usually well-rounded with more or less sharply tipped forewings. Some have prominent hindwing tails, and others have more angled wings; the skippers' basic wing shapes vary not much by comparison to the Papilionoidea, though. Most have a fairly drab coloration of browns and greys; some are more boldly black-and-white. Yellow, red, and blue hues are less often found, but some largely brown species are quite richly colored, too. Green colors and metallic iridescence are generally absent. Sexual dichromatism is present in some; males may have a blackish streak or patch of scent scales on their forewings.[2]
Skipper on a flower in Tokyo

Many species of skippers look very alike. For example, some species in the genera Amblyscirtes, Erynnis (duskywings), and Hesperia (branded skippers) cannot currently be distinguished in the field even by experts. The only reliable method of telling them apart involves dissection and microscopic examination of the genitalia, which have characteristic structures that prevent mating except between conspecifics.[2]
Subfamilies
The regent skipper (Euschemon rafflesia) is the most distinct skipper, forming a subfamily of its own.

The roughly 3500 species of skippers are now classified in these subfamilies:[5]

Coeliadinae – awls, awlets, and policemen (about 75 species)
Euschemoninae – regent skipper (monotypic)
Eudaminae – dicot skippers
Pyrginae – spread-winged skippers and firetips (including Pyrrhopyginae)
Heteropterinae – skipperlings (about 150 species)
Hesperiinae – grass skippers (over 2000 species)
Megathyminae – giant skippers (about 18 species; doubtfully distinct from Hesperiinae)
Trapezitinae – Australian skippers (about 60 species)

References

"Skipper | lepidopteran family". Encyclopedia Britannica. Retrieved 2020-10-02.
Ackery et al. (1999)
Heikkilä et al. (2012)
Kawahara & Breinholt (2014)

Brower & Warren (2008)

Ackery, P. R.; de Jong, R. & Vane-Wright, R. I. (1999). "The Butterflies: Hedyloidea, Hesperioidea and Papilionoidae". In: Kristensen, N. P. (ed.): Handbook of Zoology: A Natural History of the Phyla of the Animal Kingdom. Volume IV Arthropoda: Insecta, Part 35: Lepidoptera, Moths and Butterflies Vol.1: Evolution, Systematics, and Biogeography: 263–300. Walter de Gruyter, Berlin, New York.
Brower, Andrew V. Z. & Warren, Andrew (April 7, 2008). "Hesperiidae". Tree of Life Web Project. Retrieved December 24, 2009.
Brower, Andrew V. Z. & Warren, Andrew (2006). "The higher classification of the Hesperiidae (Lepidoptera: Hesperioidea)". Retrieved October 26, 2012.
Evans, W. H. (1951). "A Catalogue of the Hesperiidae indicating the classification and nomenclature adopted in the British Museum (Natural History). Part I. Pyrrhophyginae". London, British Museum. 92 pp. + p15. 1–9.
Evans, W. H. (1952). "A Catalogue of the Hesperiidae indicating the classification and nomenclature adopted in the British Museum (Natural History). Part II. Pyrginae. Section I". London, British Museum. 178 pp. + pls. 10–25.
Evans, W. H. (1953). "A Catalogue of the Hesperiidae indicating the classification and nomenclature adopted in the British Museum (Natural History). Part III. Pyrginae. Section II". London, British Museum. 246 pp. + pls. 26–53.
Evans, W. H. (1955). "A Catalogue of the Hesperiidae indicating the classification and nomenclature adopted in the British Museum (Natural History). Part IV. Hesperiinae and Megathyminae". London, British Museum. 499 pp. + pls. 54–88.
Heikkilä, M.; Kaila, L.; Mutanen, M.; Peña, C. & Wahlberg, N. (2012). "Cretaceous origin and repeated tertiary diversification of the redefined butterflies". Proceedings of the Royal Society B: Biological Sciences. 279(1731), 1093–1099.
Kawahara, A. Y., & Breinholt, J. W. (2014). "Phylogenomics provides strong evidence for relationships of butterflies and moths". Proceedings of the Royal Society B: Biological Sciences. 281 (1788), 20140970.
Korolev, Vladimir A. (2014). "Catalogus on the collection of Lepidoptera. Part I. Hesperiidae". Moscow, 310 p. ISBN 978-5-00077-066-5