Butterfly classification and species discovery using genomics.

Genomic sequencing of worldwide butterfly fauna followed by phylogenetic analysis of protein-coding genes informs butterfly classification throughout the taxonomic hierarchy, from families to species. As a rule, we attribute the same taxonomic rank to more prominent clades of comparable divergence (i.e., at the same level in the tree). For species delimitation, we use criteria based on relative genetic differentiation and the extent of gene exchange between populations. We analyze the current taxonomic classification of butterflies in the light of genomic phylogenies and encounter clades that correspond to yet unnamed taxa. As a result, 11 tribes, 33 subtribes, 2 genera, 11 subgenera, and 12 species are proposed as new. New tribes are: in family Nymphalidae Rafinesque, 1815: in subfamily Heliconiinae Swainson, 1822: Vindulini Grishin, (type genus Hemming, 1934) and Algiini Grishin, (type genus Herrich-Schäffer, 1864), Lebadeini Grishin, (type genus C. Felder, 1861, in subfamily Limenitidinae Behr, 1864), and Amnosiini Grishin, (type genus E. Doubleday, 1849, in subfamily Pseudergolinae Jordan, 1898) and in family Lycaenidae [Leach], [1815]: in subfamily Aphnaeinae Distant, 1884: Cigaritini Grishin, (type genus Donzel, 1848) and Axiocersini Grishin, (type genus Hübner, [1819]) and in subfamily Theclinae: Drinini Grishin, (type genus Nicéville, 1890), Hypochrysopini Grishin, (type genus C. Felder & R. Felder, 1860), Jalmenini Grishin, (type genus Hübner, 1818), Pseudalmenini Grishin, (type genus H. H. Druce, 1902), and Rapalini Grishin, (type genus F. Moore, 1881). New subtribes are: in Papilionidae Latreille, [1802], Meandrusina Grishin, (type genus F. Moore, 1888, in Papilionini Latreille, [1802]); in Pieridae Swainson, 1820: Gandacina Grishin, (type genus F. Moore, 1906, in Coliadini Swainson, 1821), Hebomoiina Grishin, (type genus Hübner, [1819], in Anthocharidini Scudder, 1889), and Pseudopierina Grishin, (type genus Godman & Salvin, 1890, in Dismorphiini Schatz, 1886); in Nymphalidae: Lachnopterina Grishin, (type genus E. Doubleday, 1847, in Algiini Grishin, trib. n.), in Vagrantini Pinratana & Eliot, 1996: Terinosina Grishin, (type genus Boisduval, 1836) and Smerinina Grishin, (type genus Hewitson, 1874), in Adoliadini Doubleday, 1845: Evenaina Grishin, (type genus Westwood, 1850) and Pseudathymina Grishin, (type genus Staudinger, 1891), and Kumothalina Grishin, (type genus Overlaet, 1940, in Cymothoini Dhungel & Wahlberg, 2018); in Riodinidae Grote, 1895 (1827): Teratophthalmina Grishin, (type genus Stichel, 1909, in Mesosemiini Bates, 1859), Argyrogrammanina Grishin, (type genus Strand, 1932, in Symmachiini Reuter, 1896), in the tribe Calydnini Seraphim, Freitas & Kaminski, 2018: Echenaidina Grishin, (type genus Hübner, [1819]) and Echydnina Grishin, (type genus J. Hall, 2002), and Cariina Grishin, (type genus Hübner, 1823, in Riodinini Grote, 1895 (1827); in Lycaenidae: Megalopalpina Grishin, (type genus Röber, 1886, in Miletini Reuter, 1896), Pseudaletidina Grishin, (type genus H. H. Druce, 1888, in Cigaritini Grishin, trib. n.), in Aphnaeini Distant, 1884: Aloeidina Grishin, (type genus Hübner, [1819]) and Phasisina Grishin, (type genus Hübner, [1819]), Pilodeudorigina Grishin, (type genus H. H. Druce, 1891, in Rapalini Grishin, trib. n.), Hemiolaina Grishin, (type genus Aurivillius, 1922, in Oxylidini Eliot, 1973), Cupidopsina Grishin, (type genus Karsch, 1895, in Hypotheclini Eliot, 1973), and in tribe Polyommatini Swainson, 1827: Theclinesthina Grishin, (type genus Röber, 1891), Azanina Grishin, (type genus F. Moore, 1881), Unina Grishin, (type genus Nicéville, 1890), Ionolycina Grishin, (type genus Toxopeus, 1929), Pithecopina Grishin, (type genus Horsfield, 1828), Zizulina Grishin, (type genus Chapman, 1910), Jamidina Grishin, (type genus Hübner, [1819]), Fameganina Grishin, (type genus Eliot, 1973), Oboroniina Grishin, (type genus Karsch, 1893), and Uranothaumatina Grishin, (type genus Butler, 1895); and in Hesperiidae Latreille, 1809, Cupithina Grishin, (type genus F. Moore, 1884, in Astictopterini Swinhoe, 1912). The new genera are Grishin, (type species Holla nd, 1891, in Gretnini Grishin, 2019) and Grishin, (type species Nicéville, 1884, in Baorini Doherty, 1886). New subgenera are: in Pieridae, Grishin, (type species H. Bates, 1861, in genus A. Butler, 1870); in Nymphalidae: Grishin, (type species Latreille, [1813], in genus Hübner, [1821]) and Grishin, (type species Hübner, [1823], in genus [Fabricius], 1807); in Lycaenidae, Grishin, (type species Linnaeus, 1764, in genus Butler, 1898); and in Hesperiidae: Grishin, (type species (?) Trimen, 1868, in genus Mabille, 1885), Grishin, (type species Holland, 1894, in genus Evans, 1937), Grishin, (type species Mabille, 1878, in genus Eliot, 1978), Grishin, (type species Evans, 1937, in genus Evans, 1937), Grishin, (type species Hewitson, 1876, in genus Evans, 1937), and in genus Hübner, 1819: Grishin, (type species H. H. Druce, 1912), and Grishin, (type species Boisduval, 1833). New species are: in Nymphalidae: Grishin, (type locality in the USA: AZ, Pima/Santa Cruz Cos.) and (type locality in Mexico: Sonora, Yécora), in Riodinidae, Grishin, (type locality in Peru: Madre de Dios), in Hesperiidae: () Grishin, (type locality in USA: CO, Jefferson Co.), () Grishin, (type locality in USA: FL, Volusia Co.), () Grishin, (type locality in Mexico: Oaxaca), () Grishin, (type locality in Brazil: Amapá), Grishin, (type locality in Guatemala), Grishin, (type locality in USA: AZ, Pima Co.), Grishin, (type locality in Togo), Grishin, (type locality in Central African Republic), and Grishin, (type locality in Malawi). Furthermore, we elevate a tribe to subfamily, resurrect 3 tribes, 3 subtribes, 5 genera (and confirm 1), 3 subgenera, change the rank of 5 currently recognized tribes to subtribes (and confirm 1), 10 genera to subgenera, synonymize 4 genera, and present evidence to support 21 taxa as species instead of subspecies and 3 taxa as subspecies instead of synonyms. Namely, we reinstate Liphyrinae Doherty, 1889 as a subfamily (was a tribe of Miletinae Reuter, 1896), treat the following as tribes: Leptidiini Grote, 1897 (in Dismorphiinae Schatz, 1886) and in Theclinae Swainson, 1830: Surendrini Koçak & Seven, 1997 (not a subtribe of Arhopalini Bingham, 1907) and Myrinini Toxopeus, 1929 (not a synonym of Amblypodiini Doherty, 1886), and subtribes: Callidryina Kirby, 1896 and Gonepterygina Verity, 1920 (in Coliadini Swainson, 1821), Abrotina Hemming, 1960 and Bebeariina Hemming, 1960 (in Adoliadini Doubleday, 1845), and Sarotina Bridges, 1988 (in Helicopini Stichel, 1928). We confirm Michener, 1943 as a valid genus, not a junior subjective synonym of Scudder, 1889 and resurrect from synonymy the following genera: Bernardi, 1953 (not Hübner, [1819]), Nicéville, 1894 (not Hübner, [1819]), Mabille, 1891 (not F. Moore, [1881]), Evans, 1912 (not Swinhoe, 1893) and subgenera: Özdikmen, 2008 and Hübner, [1821] of [Fabricius], 1807 and Wallengren, 1857 of Donzel, 1848. We change the status of the following taxa from tribes to subtribes: Nathalina Bálint, 2022 and Kricogonina Bálint, 2022 of Euremini Grote, 1898, Tarakina Eliot, 1973 of Spalgini, 1929, Horagina Swinhoe, 1910 and Loxurina Swinhoe, 1910 of Cheritrini Swinhoe, 1910, and Niphandina Sibatani & Ito, 1942 of Polyommatini; from genus to subgenus: E. Doubleday, 1847 of Hübner, [1819], Seven, 1997 of Wallengren, 1875, Nabokov, 1945 of Draudt, 1921, Balletto, 1993 of Hemming, 1960, Mabille, 1904 of Hübner, 1818, Berger, 1978 of Mabille, 1885, Evans, 1937 of Evans, 1937, Jong & Coutsis, 2017 and Kemal & Koçak, 2020 of Hübner, 1819, and Fan & Chiba, 2016 of Evans, 1935; and from genus to junior subjective synonym: Heath, 1997 of Butler, 1886, Larsen, 2013 of Aurivillius, 1925, Aurivillius, 1925 of Holland, 1896, Oberthür, 1916 of Mabille, 1898. The following taxa are species, not subspecies or synonyms: (M. Bates, 1939), and (A. Butler, 1871), (not (Poey, 1832)), (E. Doubleday, 1847), and (Wallengren, 1860), (not (Geyer, 1832)), (Röber, 1909), (not (E. Doubleday, 1847)), (Aurivillius, 1899), (not (Hübner, 1823)), () Skinner, 1911, (not () (W. H. Edwards, 1863)), () (H. Freeman, 1943), and () (Austin & J. Emmel, 1998), (not () (Scudder, 1870)), Evans, 1953, (not (W. H. Edwards, 1882)), (W. H. Edwards, 1882), (not (Herrich-Schäffer, 1869)), Glassberg 2023, (not (Fabricius, 1793)), () Mabille, 1889, and () (Plötz, 1884), (not () (Trimen, 1868)), () (Lindsey & L. Miller, 1965), , (not (Holland, 1894)), (L. Miller, 1971), , (not (Holland, 1894)), (Strand, 1912), , and (Larsen, 2013), , (not (Mabille, 1891)), (Mabille, 1890), (not (Plötz, 1886)), (Fruhstorfer, 1911), (not (Hewitson, 1876)), Evans, 1937, (not Evans, 1937), and Godman, 1900, (not a synonym of (W. H. Edwards, 1882)), while (H. Freeman, 1942), (not a synonym of W. H. Edwards, 1862) and (Lintner, 1878), (not a synonym of (Boisduval, 1852)) are subspecies. In addition, we propose new genus-species combinations: (Poey, [1852]), (H. Bates, 1861), (H. Bates, 1861), (Kollar, 1850), (C. Felder & R. Felder, 1861), (Lichy, 1976), and (Lathy, 1898) (not Hübner, [1819]), (Alphéraky, 1889) (not Tutt, 1908), () (Neave, 1910), () (Holland, 1894), () (Holland, 1894), (Evans, 1937), (Holland, 1894), (Evans, 1937), (Collins & Larsen, 2008), (Mabille, 1891), and (Holland, 1893) (not Hemming, 1939), (Holland, 1896) (not Butler, 1870), (Mabille, 1878) (not Aurivillius, 1925), (Plötz, 1879), (Evans, 1937), (Trimen, 1864), and (A. Butler, 1895) (not C. Felder & R. Felder, 1860), (Mabille, 1884) (not Kemal & Koçak, 2020), and () (Larsen, 2013), () (Gaede, 1916), () (Evans, 1937), () (Holland, 1896), and () (Evans, 1937) (not Evans, 1949); and a new species-subspecies combination Evans, 1953 (not ). Strand, 1921 is a junior subjective synonym of () (Plötz, 1884), not of () (Trimen, 1868). We transferred the tribes Oxylidini Eliot, 1973, Remelanini Eliot, 1973, and Hypolycaenini Swinhoe, 1910 from Theclinae Swainson, 1831 to Polyommatinae Swainson, 1827. We conclude that [No genus] Draudt, 1924 is an unavailable name. The is designated for Herrich-Schäffer, 1869 (type locality likely in Mexico: Oaxaca, as deduced by sequence comparison), and the is designated for Plötz, 1882 (type locality becomes Brazil: Bahia). The type localities of Holland, 1896 and Holland, 1896 are Sierra Leone: Freetown and Cameroon: Efoulan, respectively, as determined from the locality labels of primary type specimens. Finally, we provide taxonomic lists for Euremini (to worldwide subgenera and American species) and Lycaenidae (to subtribes).