Taxonomic changes suggested by the genomic analysis of Hesperiidae (Lepidoptera)

Our expanded efforts in genomic sequencing to cover additional skipper butterfly (Lepidoptera: Hesperiidae) species and populations, including primary type specimens, call for taxonomic changes to restore monophyly and correct misidentifications by moving taxa between genera and proposing new names. Reconciliation between phenotypic characters and genomic trees suggests three new tribes, two new subtribes, 23 new genera, 17 new subgenera and 10 new species that are proposed here: Psolosini Grishin, new tribe (type genus Psolos Staudinger, 1889), Ismini Grishin, new tribe (type genus Isma Distant, 1886), Eetionini Grishin, new tribe (type genus Eetion de Nicéville, 1895), Orphina Grishin, new subtribe (type genus Orphe Godman, 1901), Carystoidina Grishin, new subtribe (type genus Carystoides Godman, 1901), Fulvatis Grishin, new genus (type species Telegonus fulvius Plötz, 1882), Adina Grishin, new genus (type species Nascus adrastor Mabille and Boullet, 1912), Ornilius Grishin, new genus (type species Ornilius rotundus Grishin, new species), Tolius Grishin, new genus (type species Antigonus tolimus Plötz, 1884), Lennia Grishin, new genus (type species Leona lena Evans, 1937), Trida Grishin, new genus (type species Cyclopides barberae Trimen, 1873), Noxys Grishin, new genus (type species Oxynthes viricuculla Hayward, 1951), Gracilata Grishin, new genus (type species Enosis quadrinotata Mabille, 1889), Hermio Grishin, new genus (type species Falga ? hermione Schaus, 1913), Eutus Grishin, new genus (type species Cobalus rastaca Schaus, 1902), Gufa Grishin, new genus (type species Phlebodes gulala Schaus, 1902), Godmia Grishin, new genus (type species Euroto chlorocephala Godman, 1900), Rhomba Grishin, new genus (type species Eutychide gertschi Bell, 1937), Rectava Grishin, new genus (type species Megistias ignarus Bell, 1932), Contrastia Grishin, new genus (type species Hesperia distigma Plötz, 1882), Mit Grishin, new genus (type species Mnasitheus badius Bell, 1930), Picova Grishin, new genus (type species Vorates steinbachi Bell, 1930), Lattus Grishin, new genus (type species Eutocus arabupuana Bell, 1932), Gubrus Grishin, new genus (type species Vehilius lugubris Lindsey, 1925), Koria Grishin, new genus (type species Hesperia kora Hewitson, 1877), Corta Grishin, new genus (type species Eutychide lycortas Godman, 1900), Calvetta Grishin, new genus (type species Hesperia calvina Hewitson, 1866), Oz Grishin, new genus (type species Astictopterus ozias Hewitson, 1878), Praxa Grishin, new subgenus (type species Nascus prax Evans, 1952), Bron Grishin, new subgenus (type species Papilio broteas Cramer, 1780), Turis Grishin, new subgenus (type species Pyrgus (Scelothrix) veturius Plötz, 1884), Tiges Grishin, new subgenus (type species Antigonus liborius Plötz, 1884), Ocrypta Grishin, new subgenus (type species Notocrypta caerulea Evans, 1928), Tixe Grishin, new subgenus (type species Cobalus quadrata Herrich-Schäffer, 1869), Nycea Grishin, new subgenus (type species Pamphila hycsos Mabille, 1891), Nausia Grishin, new subgenus (type species Oenus [sic] nausiphanes Schaus, 1913), Flor Grishin, new subgenus (type species Stomyles florus Godman, 1900), Geia Grishin, new subgenus (type species Pamphila geisa Möschler, 1879), Rotundia Grishin, new subgenus (type species Enosis schausi Mielke and Casagrande, 2002), Volus Grishin, new subgenus (type species Eutocus volasus Godman, 1901), Pseudopapias Grishin, new subgenus (type species Papias tristissimus Schaus, 1902), Septia Grishin, new subgenus (type species Justinia septa Evans, 1955), Brasta Grishin, new subgenus (type species Lychnuchus brasta Evans, 1955), Bina Grishin, new subgenus (type species Cobalus gabina Godman, 1900), Balma Grishin, new subgenus (type species Carystoides balza Evans, 1955), Ornilius rotundus Grishin, new species (type locality in Brazil: Santa Catarina), Salantoia metallica Grishin, new species (type locality in Guyana: Acarai Mts.), Dyscophellus australis Grishin, new species (type locality in Paraguay: Sapucay), Dyscophellus basialbus Grishin, new species (type locality in Brazil: Rondônia), Telegonus subflavus Grishin, new species (type locality in Ecuador: Riobamba), Decinea colombiana Grishin, new species (type locality in Colombia: Bogota), Lerema lucius Grishin, new species (type locality in Panama: Colón), Cynea rope Grishin, new species (type locality in Nicaragua: Chontales), Lerodea sonex Grishin, new species (type locality in Peru: Cuzco), and Metiscus goth Grishin, new species (type locality in Costa Rica). Lectotypes are designated for the following 17 taxa: Telegonus gildo Mabille, 1888, Netrocoryne damias Plötz, 1882, Telegonus erythras Mabille, 1888, Telegonus galesus Mabille, 1888, Eudamus cretellus Herrich-Schäffer, 1869, Leucochitonea chaeremon Mabille, 1891, Antigonus aura Plötz, 1884, Pamphila voranus Mabille, 1891, Hesperia pupillus Plötz, 1882, Cobalus lumina Herrich-Schäffer, 1869, Cobalus stigmula Mabille, 1891, Megistias isus Godman, 1900, Cobalopsis latonia Schaus, 1913, Pamphila nubila Mabille, 1891, Metiscus atheas Godman, 1900, Mnasalcas amatala Schaus, 1902, and Hesperia ina Plötz, 1882. The lectotype of Hesperia infuscata Plötz, 1882 is invalid because it does not agree with the original description and illustration by Plötz, is not from the locality listed in the original description, and therefore is not a syntype. Neotypes are designated for the following five taxa: Telegonus corentinus Plötz, 1882, Hesperia dido Plötz, 1882, Hesperia distigma Plötz, 1882, Hesperia infuscata Plötz, 1882, and Hesperia pruinosa Plötz, 1882. As a result, the following five taxa are junior objective synonyms: Telegonus diophorus Möschler, 1883 of Telegonus corentinus Plötz, 1882, Pamphila puxillius Mabille, 1891 of Hesperia pupillus Plötz, 1882, Cobalus stigmula Mabille, 1891 of Hesperia distigma Plötz, 1882, Mnasalcas amatala Schaus, 1902 of Hesperia infuscata Plötz, 1882, and Hesperia pruinosa Plötz, 1882 of Hesperia uza Hewitson, 1877. Morys valerius valda Evans, 1955 is fixed as the type species of Morys Godman, 1900, and Pamphila compta Butler, 1877 is reaffirmed as the type species of Euroto Godman, 1900. Furthermore, the following taxonomic changes are suggested. Prosopalpus Holland, 1896, Lepella Evans, 1937, and Creteus de Nicéville, 1895 are placed in Aeromachini Tutt, 1906. Triskelionia Larsen and Congdon, 2011 is transferred from Celaenorrhinini Swinhoe, 1912 to Tagiadini Mabille, 1878. Kobelana Larsen and Collins, 2013 is transferred from Tagiadini Mabille, 1878 to Celaenorrhinini Swinhoe, 1912. The following nine genus-group names are resurrected from synonymy and treated as valid genera: Abaratha Moore, 1881 (not in Caprona Wallengren, 1857), Bibla Mabille, 1904 (not in Taractrocera Butler, 1870), Kerana Distant, 1886 and Tamela Swinhoe, 1913 (not in Ancistroides Butler, 1874), Metrocles Godman, 1900 (not in Metron Godman, 1900), Alerema Hayward, 1942 (not in Tigasis Godman, 1900), Metiscus Godman, 1900 (not in Enosis Mabille, 1889), Vistigma Hayward, 1939 (not in Phlebodes Hübner, [1819]), and Mnasalcas Godman, 1900 (not in Mnasitheus Godman, 1900). The genus-group names Daimio Murray, 1875 and Pterygospidea Wallengren, 1857 are resurrected from synonymy and treated as valid subgenera of Tagiades Hübner, [1819]. We confirm Apallaga Strand, 1911 as a valid genus. The following 24 genera are placed as subgenera, new status: Pseudonascus Austin, 2008 of Nascus Watson, 1893; Albiphasma Huang, Chiba, Wang and Fan, 2016 of Pintara Evans, 1932; Ctenoptilum de Nicéville, 1890 of Tapena Moore, [1881]; Odontoptilum de Nicéville, 1890 of Abaratha Moore, 1881; Caprona Wallengren, 1857 of Abantis Hopffer, 1855; Timochreon Godman and Salvin, 1896 of Zopyrion Godman and Salvin, 1896; Pulchroptera Hou, Fan and Chiba, 2021 of Heteropterus Duméril, 1806; Stimula de Nicéville, 1898 of Koruthaialos Watson, 1893; Udaspes Moore, [1881] and Notocrypta de Nicéville, 1889 of Ancistroides Butler, 1874; Cravera de Jong, 1983 of Xeniades Godman, 1900; Cobaloides Hayward, 1939 of Oligoria Scudder, 1872; Saniba O. Mielke and Casagrande, 2003 of Psoralis Mabille, 1904; Quinta Evans, 1955 of Cynea Evans, 1955; Styriodes Schaus, 1913 and Remella Hemming, 1939 of Mnasicles Godman, 1901; Repens Evans, 1955 of Eprius Godman, 1901; Morys Godman, 1900 of Lerema Scudder, 1872; Enosis Mabille, 1889 of Lychnuchus Hübner, [1831]; Penicula Evans, 1955 of Vistigma Hayward, 1939; Mnasinous Godman, 1900 of Methionopsis Godman, 1901; and Moeros Evans, 1955, Argon Evans, 1955, and Synale Mabille, 1904 of Carystus Hübner, [1819]. The following 20 genera are treated as junior subjective synonyms: Leucochitonea Wallengren, 1857 of Abantis Hopffer, 1855; Sapaea Plötz, 1879 and Netrobalane Mabille, 1903 of Caprona Wallengren, 1857; Parasovia Devyatkin, 1996 of Sebastonyma Watson, 1893; Pemara Eliot, 1978 of Oerane Elwes and Edwards, 1897; Ankola Evans, 1937 of Pardaleodes Butler, 1870; Arotis Mabille, 1904 of Mnaseas Godman, 1901; Chalcone Evans, 1955, Hansa Evans, 1955, and Propertius Evans, 1955 of Metrocles Godman, 1900; Jongiana O. Mielke and Casagrande, 2002 of Cobaloides Hayward, 1939; Pamba Evans, 1955 of Psoralis Mabille, 1904; Brownus Grishin, 2019 of Styriodes Schaus, 1913; Mnasilus Godman, 1900 of Papias Godman, 1900; Sucova Evans, 1955 of Mnasitheus Godman, 1900; Pyrrhocalles Mabille, 1904 and Asbolis Mabille, 1904 of Choranthus Scudder, 1872; Miltomiges Mabille, 1903 of Methionopsis Godman, 1901; Sacrator Evans, 1955 of Thracides Hübner, [1819]; and Lychnuchoides Godman, 1901 of Perichares Scudder, 1872. Arunena Swinhoe, 1919 is a junior subjective synonym of Stimula de Nicéville, 1898 (not of Koruthaialos Watson, 1893). The following 27 names are species-level taxa (some in new combinations) reinstated from synonymy: Salantoia gildo (Mabille, 1888) (not Salatis cebrenus (Cramer, 1777)), Bungalotis corentinus (Plötz, 1882) (not Bungalotis midas (Cramer, 1775)), Telegonus cretellus (Herrich-Schäffer, 1869) (not Telegonus cassander (Fabricius, 1793)), Santa palica (Mabille, 1888) (not Chiothion asychis (Stoll, 1780)), Camptopleura cincta Mabille and Boullet, 1917 (not Camptopleura auxo (Möschler, 1879)), Camptopleura orsus (Mabille, 1889) (not Nisoniades mimas (Cramer, 1775)), Metron voranus (Mabille, 1891) and Metron fasciata (Möschler, 1877) (not Metron zimra (Hewitson, 1877)), Limochores catahorma (Dyar, 1916) (not Limochores pupillus (Plötz, 1882)), Pares viridiceps (Mabille, 1889) (not Thoon modius (Mabille, 1889)), Tigasis wellingi (Freeman, 1969) (not Tigasis arita (Schaus, 1902)), Rectava sobrinus (Schaus, 1902) (not Papias phainis Godman, 1900), Nastra subsordida (Mabille, 1891) (not Adlerodea asema (Mabille, 1891), previously in Eutychide Godman, 1900), Lerema pattenii Scudder, 1872 (not Lerema accius (J. E. Smith, 1797)), Lerema (Morys) ancus (Möschler, 1879) (not Cymaenes tripunctus theogenis (Capronnier, 1874)), Cobalopsis zetus (Bell, 1942) (not Cobalopsis nero (Herrich-Schäffer, 1869)), Lerema (Geia) etelka (Schaus, 1902) (not Lerema (Geia) geisa (Möschler, 1879), previously in Morys Godman, 1900), Cymaenes isus (Godman, 1900) (not Cymaenes trebius (Mabille, 1891)), Vehilius labdacus (Godman, 1900) (not Vehilius inca (Scudder, 1872)), Papias amyrna (Mabille, 1891) (not Papias allubita (Butler, 1877), previously in Mnasilus Godman, 1900), Papias integra (Mabille, 1891) (not Papias subcostulata (Herrich-Schäffer, 1870)), Metiscus atheas Godman, 1900 (not Hesperia achelous Plötz, 1882), Dion agassus (Mabille, 1891) (not Dion uza (Hewitson, 1877), previously in Enosis Mabille, 1889), Picova incompta (Hayward, 1942) (not Lerema (Morys) micythus (Godman, 1900), previously in Morys Godman, 1900), Lucida melitaea (Draudt, 1923) (not Lucida lucia (Capronnier, 1874)), Methionopsis modestus Godman, 1901 (not Methionopsis ina (Plötz, 1882)), and Thargella (Volus) volasus (Godman, 1901) (not Eutocus facilis (Plötz, 1884)). The following 57 taxa are elevated from subspecies to species, new status (some in new combinations): Dyscophellus doriscus (Hewitson, 1867) (not Dyscophellus porcius (C. Felder and R. Felder, 1862), Phocides vida (A. Butler, 1872) (not Phocides urania (Westwood, 1852)), Tagiades (Daimio) ceylonica Evans, 1932 (not Tagiades litigiosa Möschler, 1878), Tagiades (Daimio) tubulus Fruhstorfer, 1910 (not Tagiades sambavana Elwes and Edwards, 1897), Tagiades (Daimio) kina Evans, 1934, Tagiades (Daimio) sheba Evans, 1934, Tagiades (Daimio) martinus Plötz, 1884, Tagiades (Daimio) sem Mabille, 1883, and Tagiades (Daimio) neira Plötz, 1885 (not Tagiades trebellius (Hopffer, 1874)), Tagiades (Daimio) korela Mabille, 1891 and Tagiades (Daimio) presbyter Butler, 1882 (not Tagiades nestus (C. Felder, 1860)), Tagiades obscurus Mabille, 1876, Tagiades ravi (Moore, [1866]), Tagiades atticus (Fabricius, 1793), Tagiades titus Plötz, 1884, Tagiades janetta Butler, 1870, Tagiades inconspicua Rothschild, 1915, and Tagiades hovia Swinhoe, 1904 (not Tagiades japetus (Stoll, [1781])), Tagiades silvia Evans, 1934 and Tagiades elegans Mabille, 1877 (not Tagiades gana (Moore, [1866])), Tapena bornea Evans, 1941 and Tapena minuscula Elwes and Edwards, 1897 (not Tapena thwaitesi Moore, [1881]), Darpa dealbata (Distant, 1886) (not Darpa pteria (Hewitson, 1868)), Perus manx (Evans, 1953) (not Perus minor (Schaus, 1902)), Canesia pallida (Röber, 1925) (not Carrhenes canescens (R. Felder, 1869)), Carrhenes conia Evans, 1953 (not Carrhenes fuscescens (Mabille, 1891)), Anisochoria extincta Hayward, 1933 and Anisochoria polysticta Mabille, 1876 (not Anisochoria pedaliodina (Butler, 1870)), Anisochoria verda Evans, 1953 (not Anisochoria minorella Mabille, 1898), Bralus alco (Evans, 1953) (not Bralus albida (Mabille, 1888)), Ephyriades jamaicensis (Möschler, 1879) (not Ephyriades brunnea (Herrich-Schäffer, 1865)), Koruthaialos (Stimula) frena Evans, 1949 (not Koruthaialos focula (Plötz, 1882)), Euphyes kiowah (Reakirt, 1866) (not Euphyes vestris (Boisduval, 1852)), Mnaseas inca Bell, 1930 (not Mnaseas bicolor (Mabille, 1889)), Metron hypochlora (Draudt, 1923) (not Metrocles schrottkyi (Giacomelli, 1911), previously in Metron Godman, 1900), Decinea huasteca (H. Freeman, 1969), Decinea denta Evans, 1955, and Decinea antus (Mabille, 1895) (not Decinea decinea (Hewitson, 1876)), Xeniades pteras Godman, 1900 (not Xeniades chalestra (Hewitson, 1866)), Xeniades difficilis Draudt, 1923 (not Xeniades orchamus (Cramer, 1777)), Xeniades hermoda (Hewitson, 1870) (not Tisias quadrata (Herrich-Schäffer, 1869)), Hermio vina (Evans, 1955) (not Hermio hermione (Schaus, 1913), previously in Lento Evans, 1955), Cymaenes loxa Evans, 1955, (not Cymaenes laureolus (Schaus, 1913)), Niconiades peri (Evans, 1955) (not Rhinthon bajula (Schaus, 1902), previously in Neoxeniades Hayward, 1938), Gallio danius (Bell, 1941) (not Vehilius seriatus (Mabille, 1891)), Gallio massarus (E. Bell, 1940) (not Gallio garima (Schaus, 1902) previously in Tigasis Godman, 1900), Cymaenes edata (Plötz, 1882), Cymaenes miqua (Dyar, 1913) and Cymaenes aequatoria (Hayward, 1940) (not Cymaenes odilia (Burmeister, 1878)), Lychnuchus (Enosis) demon (Evans, 1955) (not Lychnuchus (Enosis) immaculata (Hewitson, 1868), previously in Enosis Mabille, 1889), Naevolus naevus Evans, 1955 (not Naevolus orius (Mabille, 1883)), Lucida scopas (Mabille, 1891), Lucida oebasus (Godman, 1900), and Lucida leopardus (Weeks, 1901) (not Lucida lucia (Capronnier, 1874)), Corticea schwarzi (E. Bell, 1941) and Corticea sylva (Hayward, 1942) (not Corticea mendica (Mabille, 1898)), and Choranthus orientis (Skinner, 1920) (not Choranthus antiqua (Herrich-Schäffer, 1863), previously in Pyrrhocalles Mabille, 1904). Borbo impar bipunctata (Elwes and J. Edwards, 1897) is a valid subspecies, not a synonym of Borbo impar tetragraphus (Mabille, 1891), here placed in synonymy with Lotongus calathus (Hewitson, 1876), new synonym. We confirm the species status of Telegonus cassius (Evans, 1952) and Lerema (Morys) valda Evans, 1955. Euphyes chamuli Freeman, 1969 is placed as a subspecies of Euphyes kiowah (Reakirt, 1866), new status. The following 41 taxa are junior subjective synonyms, either newly proposed or transferred from synonymy with other species or subspecies: Telegonus mutius Plötz, 1882 of Euriphellus phraxanor (Hewitson, 1876), Telegonus erythras Mabille, 1888 of Dyscophellus damias (Plötz, 1882), Aethilla jaira Butler, 1870 of Telegonus cretellus (Herrich-Schäffer, 1869), Paches era Evans, 1953 of Santa palica (Mabille, 1888), Antigonus alburnea Plötz, 1884 of Tolius tolimus robigus (Plötz, 1884) (not of Echelatus sempiternus simplicior (Möschler, 1877)), Echelatus depenicillus Strand, 1921 of E. sempiternus simplicior (not of T. tolimus robigus), Antigonus aura Plötz, 1884 of Theagenes dichrous (Mabille, 1878) (not of Helias phalaenoides palpalis (Latreille, [1824])), Achlyodes impressus Mabille, 1889 of Camptopleura orsus (Mabille, 1889), Augiades tania Schaus, 1902 of Metron voranus (Mabille, 1891), Pamphila verdanta Weeks, 1906 of Metron fasciata (Möschler, 1877), Niconiades viridis vista Evans, 1955 of Niconiades derisor (Mabille, 1891), Pamphila binaria Mabille, 1891 of Conga chydaea (A. Butler, 1877) (not of Cynea cynea (Hewitson, 1876)), Psoralis concolor Nicolay, 1980 of Ralis immaculatus (Hayward, 1940), Hesperia dido Plötz, 1882 of Cynea (Quinta) cannae (Herrich-Schäffer, 1869) (not of Lerema lochius (Plötz, 1882)), Proteides osembo Möschler, 1883 of Cynea (Cynea) diluta (Herrich-Schäffer, 1869) (not of Cynea (Quinta) cannae (Herrich-Schäffer, 1869)), Cobalopsis brema E. Bell, 1959 of Eutus rastaca (Schaus, 1902), Psoralis panamensis Anderson and Nakamura, 2019 of Rhomba gertschi (Bell, 1937), Cobalus asella Herrich-Schäffer, 1869 of Amblyscirtes alternata (Grote and Robinson, 1867) (not of Amblyscirtes vialis (W. H. Edwards, 1862)), Papias trimacula Nicolay, 1973 of Nastra subsordida (Mabille, 1891), Pamphila bipunctata Mabille, 1889 and Sarega staurus Mabille, 1904 of Lerema pattenii Scudder, 1872 (not of Cymaenes lumina (Herrich-Schäffer, 1869), previously in Lerema Scudder, 1872), Hesperia aethra Plötz, 1886 of Lerema lineosa (Herrich-Schäffer, 1865) (not of Lerema (Morys) compta Butler, 1877), Megistias miaba Schaus, 1902 of Cobalopsis valerius (Möschler, 1879), Phanis sylvia Kaye, 1914 of Lerema etelka (Schaus, 1902) (not of Lerema (Geia) geisa (Möschler, 1879), previously in Morys Godman, 1900), Carystus odilia Burmeister, 1878, Pamphila trebius Mabille, 1891 and Megistias corescene Schaus, 1902 of Cymaenes lumina (Herrich-Schäffer, 1869), Hesperia phocylides Plötz, 1882 of Cymaenes edata (Plötz, 1882) (not of Lerema accius (J. E. Smith, 1797)), Pamphila xenos Mabille, 1898 of Vehilius inca (Scudder, 1872), Mnasilus guianae Lindsey, 1925 of Papias amyrna (Mabille, 1891), Pamphila nubila Mabille, 1891 of Papias integra (Mabille, 1891) (not of Cynea corisana (Plötz, 1882)), Enosis matheri H. Freeman, 1969 of Metiscus atheas Godman, 1900 (previously in Enosis Mabille, 1889), Hesperia infuscata Plötz, 1882 of Mnaseas derasa derasa (Herrich-Schäffer, 1870) (previously Arotis Mabille, 1904), (not of Papias subcostulata (Herrich-Schäffer, 1870)), Pamphila astur Mabille, 1891 of Metiscus angularis (Möschler, 1877) (not of Cymaenes tripunctus theogenis (Capronnier, 1874)), Anthoptus macalpinei H. Freeman, 1969 of Anthoptus inculta (Dyar, 1918), Methionopsis typhon Godman, 1901 of Methionopsis ina (Plötz, 1882), Methionopsis dolor Evans, 1955 of Thargella volasus (Godman, 1901), Hesperia cinica Plötz, 1882 of Dubiella dubius (Stoll, 1781), Cobalus disjuncta Herrich-Schäffer, 1869 of Dubiella dubius (Stoll, 1781) (not of Vettius lafrenaye (Latreille, [1824])), and Saliana vixen Evans, 1955 of Neoxeniades parna (Evans, 1955). The following are new and revised genus-species combinations: Euriphellus cebrenus (Cramer, 1777) (not Salatis Evans, 1952), Gorgopas extensa (Mabille, 1891) (not Polyctor Evans, 1953), Clytius shola (Evans, 1953) (not Staphylus Godman and Salvin, 1896), Perus narycus (Mabille, 1889) (not Ouleus Lindsey, 1925), Perus parvus (Steinhauser and Austin, 1993) (not Staphylus Godman and Salvin, 1896), Pholisora litus (Dyar, 1912) (not Bolla Mabille, 1903), Carrhenes decens (A. Butler, 1874) (not Antigonus Hübner, [1819]), Santa palica (Mabille, 1888) (not Chiothion Grishin, 2019), Bralus nadia (Nicolay, 1980) (not Anisochoria Mabille, 1876), Acerbas sarala (de Nicéville, 1889) (not Lotongus Distant, 1886), Caenides sophia (Evans, 1937) (not Hypoleucis Mabille, 1891), Hypoleucis dacena (Hewitson, 1876) (not Caenides Holland, 1896), Dotta tura (Evans, 1951) (not Astictopterus C. Felder and R. Felder, 1860), Nervia wallengrenii (Trimen, 1883) (not Kedestes Watson, 1893), Testia mammaea (Hewitson, 1876) (not Decinea Evans, 1955), Oxynthes trinka (Evans, 1955) (not Orthos Evans, 1955), Metrocles argentea (Weeks, 1901) (not Paratrytone Godman, 1900), Metrocles scitula (Hayward, 1951) (not Mucia Godman, 1900), Metrocles schrottkyi (Giacomelli, 1911) (not Metron Godman, 1900), Niconiades derisor (Mabille, 1891) (not Decinea Evans, 1955), Paratrytone samenta (Dyar, 1914) (not Ochlodes Scudder, 1872), Oligoria (Cobaloides) locutia (Hewitson, 1876) (not Quinta Evans, 1955), Psoralis (Saniba) laska (Evans, 1955) (not Vidius Evans, 1955), Psoralis (Saniba) arva (Evans, 1955) and Psoralis (Saniba) umbrata (Erschoff, 1876) (not Vettius Godman, 1901), Psoralis (Saniba) calcarea (Schaus, 1902) and Psoralis (Saniba) visendus (E. Bell, 1942) (not Molo Godman, 1900), Alychna gota (Evans, 1955) (not Psoralis Mabille, 1904), Adlerodea asema (Mabille, 1891) and Adlerodea subpunctata (Hayward, 1940) (not Eutychide Godman, 1900), Ralis immaculatus (Hayward, 1940) (not Mucia Godman, 1900), Rhinthon braesia (Hewitson, 1867) and Rhinthon bajula (Schaus, 1902) (not Neoxeniades Hayward, 1938), Cymaenes lochius Plötz, 1882 (not Lerema Scudder, 1872), Paracarystus ranka (Evans, 1955) (not Thoon Godman, 1900), Tricrista aethus (Hayward, 1951), Tricrista canta (Evans, 1955), Tricrista slopa (Evans, 1955), Tricrista circellata (Plötz, 1882), and Tricrista taxes (Godman, 1900) (not Thoon Godman, 1900), Gallio madius (E. Bell, 1941) and Gallio seriatus (Mabille, 1891) (not Vehilius Godman, 1900), Gallio garima (Schaus, 1902) (not Tigasis Godman, 1900), Tigasis corope (Herrich-Schäffer, 1869) (not Cynea Evans, 1955), Tigasis perloides (Plötz, 1882) (not Cymaenes Scudder, 1872), Amblyscirtes (Flor) florus (Godman, 1900) (not Repens Evans, 1955), Vidius fraus (Godman, 1900) (not Cymaenes Scudder, 1872), Nastra celeus (Mabille, 1891) (not Vehilius Godman, 1900), Nastra nappa (Evans, 1955) (not Vidius Evans, 1955), Vehilius warreni (Weeks, 1901) and Vehilius limae (Lindsey, 1925) (not Cymaenes Scudder, 1872), Cymaenes lumina (Herrich-Schäffer, 1869) (not Lerema Scudder, 1872), Cobalopsis valerius (Möschler, 1879) (not Cobalopsis Godman, 1900), Cobalopsis dictys (Godman, 1900) (not Papias Godman, 1900), Lerema (Morys) venias (Bell, 1942) (not Cobalopsis Godman, 1900), Papias latonia (Schaus, 1913) (not Cobalopsis Godman, 1900), Dion iccius (Evans, 1955) and Dion uza (Hewitson, 1877) (not Enosis Mabille, 1889), Vistigma (Vistigma) opus (Steinhauser, 2008) (not Thoon Godman, 1900), Saturnus fartuga (Schaus, 1902) (not Parphorus Godman, 1900), Phlebodes fuldai (E. Bell, 1930) (not Vettius Godman, 1901), Mnasitheus padus (Evans, 1955) (not Moeris Godman, 1900), Naevolus brunnescens (Hayward, 1939) (not Psoralis Mabille, 1904), Lamponia ploetzii (Capronnier, 1874) (not Vettius Godman, 1901), Mnestheus silvaticus Hayward, 1940 (not Ludens Evans, 1955), Rigga spangla (Evans, 1955) (not Sodalia Evans, 1955), Corticea vicinus (Plötz, 1884) (not Lento Evans, 1955), Mnasalcas thymoetes (Hayward, 1942) (not Mnasicles Godman, 1901), Mnasalcas boyaca (Nicolay, 1973) (not Pamba Evans, 1955), Vertica brasta (Evans, 1955) (not Lychnuchus Hübner, [1831]), Carystina discors Plötz, 1882 (not Cobalus Hübner, [1819]), Zetka irena (Evans, 1955) (not Neoxeniades Hayward, 1938), and Neoxeniades parna (Evans, 1955) (not Niconiades Hübner, [1821]). The following are new or revised species-subspecies combinations: Tagiades neira moti Evans, 1934, Tagiades neira canonicus Fruhstorfer, 1910, Tagiades sheba vella Evans, 1934, Tagiades sheba lola Evans, 1945, Tagiades korela biakana Evans, 1934, Tagiades korela mefora Evans, 1934, Tagiades korela suffusus Rothschild, 1915, Tagiades korela brunta Evans, 1949, Tagiades ravi ravina Fruhstorfer, 1910, Tagiades atticus carnica Evans, 1934, Tagiades atticus nankowra Evans, 1934, Tagiades atticus helferi C. Felder, 1862, Tagiades atticus balana Fruhstorfer, 1910, Tagiades inconspicua mathias Evans, 1934, Tagiades hovia kazana Evans, 1934, Tagiades elegans fuscata de Jong and Treadaway, 2007, Tagiades elegans semperi Fruhstorfer, 1910, Metron hypochlora tomba Evans, 1955, Decinea denta pruda Evans, 1955, and Choranthus orientis eleutherae (Bates, 1934) (previously in Pyrrhocalles Mabille, 1904). In addition to the abovementioned changes, the following new combinations involve newly proposed genus group names: Fulvatis fulvius (Plötz, 1882) and Fulvatis scyrus (E. Bell, 1934) (not Salatis Evans, 1952); Adina adrastor (Mabille and Boullet, 1912) (not Bungalotis Watson, 1893); Nascus (Praxa) prax Evans, 1952, Nascus (Bron) broteas (Cramer, 1780), and Nascus (Bron) solon (Plötz, 1882) (not Pseudonascus Austin, 2008); Chirgus (Turis) veturius (Plötz, 1884); Paches (Tiges) liborius (Plötz, 1884), and Paches (Tiges) mutilatus (Hopffer, 1874) (not Antigonus Hübner, [1819]); Paches (Tiges) exosa (A. Butler, 1877); Tolius tolimus (Plötz, 1884) and Tolius luctuosus (Godman & Salvin, 1894) (not Echelatus Godman and Salvin, 1894); Ancistroides (Ocrypta) caerulea (Evans, 1928), Ancistroides (Ocrypta) renardi (Oberthür, 1878), Ancistroides (Ocrypta) waigensis (Plötz, 1882), Ancistroides (Ocrypta) aluensis (Swinhoe, 1907), Ancistroides (Ocrypta) flavipes (Janson, 1886), and Ancistroides (Ocrypta) maria (Evans, 1949) (not Notocrypta de Nicéville, 1889); Lennia lena (Evans, 1937), Lennia binoevatus (Mabille, 1891), Lennia maracanda (Hewitson, 1876), and Lennia lota (Evans, 1937) (not Leona Evans, 1937); Trida barberae (Trimen, 1873) and Trida sarahae (Henning and Henning, 1998) (not Kedestes Watson, 1893); Noxys viricuculla (Hayward, 1951) (not Oxynthes Godman, 1900); Xeniades (Tixe) quadrata (Herrich-Schäffer, 1869), Xeniades (Tixe) rinda (Evans, 1955), Xeniades (Tixe) putumayo (Constantino and Salazar, 2013) (not Tisias Godman, 1901); Gracilata quadrinotata (Mabille, 1889) (not Styriodes Schaus, 1913); Hermio hermione (Schaus, 1913) (not Lento Evans, 1955); Cynea (Nycea) hycsos (Mabille, 1891), Cynea (Nycea) corisana (Plötz, 1882), Cynea (Nycea) popla Evans, 1955, Cynea (Nycea) iquita (E. Bell, 1941), Cynea (Nycea) robba Evans, 1955, Cynea (Nycea) melius (Geyer, 1832), and Cynea (Nycea) irma (Möschler, 1879); Eutus rastaca (Schaus, 1902) (not Eutychide Godman, 1900); Eutus yesta (Evans, 1955) (not Thoon Godman, 1900); Eutus mubevensis (E. Bell, 1932) (not Tigasis Godman, 1900); Gufa gulala (Schaus, 1902) (not Mucia Godman, 1900); Gufa fusca (Hayward, 1940) (not Tigasis Godman, 1900); Godmia chlorocephala (Godman, 1900) (not Onophas Godman, 1900); Rhomba gertschi (E. Bell, 1937) (not Justinia Evans, 1955); Mnasicles (Nausia) nausiphanes (Schaus, 1913) (not Tigasis Godman, 1900); Amblyscirtes (Flor) florus (Godman, 1900) (not Repens Evans, 1955); Rectava ignarus (E. Bell, 1932) (not Papias Godman, 1900); Rectava vorgia (Schaus, 1902) (not Cobalopsis Godman, 1900); Rectava nostra (Evans, 1955) (not not Vidius Evans, 1955); Lerema (Geia) geisa (Möschler, 1879) and Lerema (Geia) lyde (Godman, 1900) (not Morys Godman, 1900); Contrastia distigma (Plötz, 1882) (not Cymaenes Scudder, 1872); Mit (Mit) badius (E. Bell, 1930) (not Styriodes Schaus, 1913); Mit (Mit) gemignanii (Hayward, 1940), (not Mnasitheus Godman, 1900); Mit (Rotundia) schausi (Mielke and Casagrande, 2002), (not Enosis Mabille, 1889); Picova steinbachi (E. Bell, 1930) (not Saturnus Evans, 1955); Lattus arabupuana (E. Bell, 1932) (not Eutocus Godman, 1901); Gubrus lugubris (Lindsey, 1925) (not Vehilius Godman, 1900); Thargella (Pseudopapias) tristissimus (Schaus, 1902) (not Papias Godman, 1900); Koria kora (Hewitson, 1877) (not Justinia Evans, 1955); Justinia (Septia) septa Evans, 1955; Corta lycortas (Godman, 1900) (not Orthos Evans, 1955); Vertica (Brasta) brasta (Evans, 1955) (not Lychnuchus Hübner, [1831]); Calvetta calvina (Hewitson, 1866) (not Cobalus Hübner, [1819]); Neoxeniades (Bina) gabina (Godman, 1900) (not Orthos Evans, 1955); Oz ozias (Hewitson, 1878) and Oz sebastiani Salazar and Constantino, 2013 (not Lychnuchoides Godman, 1901); and Carystoides (Balma) balza Evans, 1955 and Carystoides (Balma) maroma (Möschler, 1877). Finally, unless stated otherwise, all subgenera, species, subspecies and synonyms of mentioned genera and species are transferred together with their parent taxa, and taxa not mentioned in this work remain as previously classified.