Genomics-guided refinement of butterfly taxonomy.

Continuing with comparative genomic exploration of worldwide butterfly fauna, we use all protein-coding genes as they are retrieved from the whole genome shotgun sequences for phylogeny construction. Analysis of these genome-scale phylogenies projected onto the taxonomic classification and the knowledge about butterfly phenotypes suggests further refinements of butterfly taxonomy that are presented here. As a general rule, we assign most prominent clades of similar genetic differentiation to the same taxonomic rank, and use criteria based on relative population diversification and the extent of gene exchange for species delimitation. As a result, 7 tribes, 4 subtribes, 14 genera, and 9 subgenera are proposed as new, i.e., in subfamily Pierinae Swainson, 1820: Calopierini Grishin, trib. n. (type genus Calopieris Aurivillius, 1898); in subfamily Riodininae Grote, 1895: Callistiumini Grishin, trib. n. (type genus Callistium Stichel, 1911); in subfamily Nymphalinae Rafinesque, 1815: Pycinini Grishin, trib. n. (type genus Pycina Doubleday 1849), Rhinopalpini Grishin, trib. n. (type genus Rhinopalpa C. & R. Felder 1860), Kallimoidini Grishin, trib. n. (type genus Kallimoides Shirôzu & Nakanishi 1984), Vanessulini Grishin, trib. n. (type genus Vanessula Dewitz 1887), and Doleschalliaini Grishin, trib. n. (type genus Doleschallia C. & R. Felder 1860); in tribe Mesosemiini Bates, 1859: Eunogyrina Grishin, subtrib. n. (type genus Eunogyra Westwood, 1851); in tribe Satyrini Boisduval, 1833: Callerebiina Grishin, subtrib. n. (type genus Callerebia Butler, 1867), Gyrocheilina Grishin, subtrib. n. (type genus Gyrocheilus Butler, 1867), and Calistina Grishin, subtrib. n. (type genus Calisto Hübner, [1823]); in subfamily Euselasiinae Kirby, 1871: Pelolasia Grishin, gen. n. (type species Eurygona pelor Hewitson, [1853]), Myselasia Grishin, gen. n. (type species Eurygona mys Herrich-Schäffer, [1853]), Eurylasia Grishin, gen. n. (type species Eurygona euryone Hewitson, 1856), Maculasia Grishin, gen. n. (type species Euselasia albomaculiga Callaghan, 1999), and Eugelasia Grishin, gen. n. (type species Eurygona eugeon Hewitson, 1856); in subtribe Mesosemiina Bates, 1859: Ectosemia Grishin, gen. n. (type species Papilio eumene Cramer, 1776) and Endosemia Grishin, gen. n. (type species Papilio ulrica Cramer, 1777); in tribe Symmachiini Reuter, 1896: Tigria Grishin, gen. n. (type species Mesene xypete Hewitson, 1870) and Asymma Grishin, gen. n. (type species Symmachia virgatula Stichel, 1910); in tribe Riodinini Grote, 1895: Putridivora Grishin, gen. n. (type species Charis argyrea Bates, 1868), Chadia Grishin, gen. n. (type species Charis cadytis Hewitson, 1866), Inkana Grishin, gen. n. (type species Charis incoides Schaus, 1902), and Oco Grishin, gen. n. (type species Symmachia ocellata Hewitson, 1867); in subtribe Zabuellina Seraphim, Freitas & Kaminski, 2018: Teenie Grishin, gen. n. (type species Calydna tinea Bates, 1868); Boreographium Grishin, subgen. n. (type species Papilio marcellus Cramer, 1777, parent genus Eurytides Hübner, [1821]), Esperourus Grishin, subgen. n. (type species Papilio esperanza Beutelspacher, 1975, parent genus Pterourus Scopoli, 1777), Hyppasonia Grishin, subgen. n. (type species Papilio hyppason Cramer, 1775, parent genus Heraclides Hübner, [1819]), Sisymbria Grishin, subgen. n. (type species Pieris sisymbrii Boisduval, 1852, parent genus Pontia [Fabricius], 1807), Greenie Grishin, subgen. n. (type species Thecla sheridonii [sic] Edwards, 1877, parent genus Callophrys Billberg, 1820), Magda Grishin, subgen. n. (type species Erebia magdalena Strecker, 1880, parent genus Erebia Dalman, 1816), and in genus Eresia Boisduval, 1836: Notilia Grishin, subgen. n. (type species Eresia orthia Hewitson, 1864), Levinata Grishin, subgen. n. (type species Eresia levina Hewitson, 1872), and Ithra Grishin, subgen. n. (type species Phyciodes ithra Kirby, 1900). Furthermore, we resurrect 6 genera, change the rank of 36 currently used genera to subgenus, synonymize 3 subtribes, 42 genera or subgenera, assign 3 genera to tribes and subtribes, and transfer 34 additional species to genera different from those these taxa are presently assigned to, present evidence to support 7 taxa as species instead of subspecies, and 1 taxon as a subspecies instead of species. Namely, the following taxa are valid genera: Terias Swainson, 1821 (not in Eurema Hübner, [1819]), Erythia Hübner, [1819] and Marmessus Hübner, [1819] (not in Euselasia Hübner, [1819]), Eucorna Strand, 1932 (not in Voltinia Stichel, 1910), Cremna Doubleday, 1847 (not in Napaea Hübner, [1819]), and Hallonympha Penz & DeVries, 2006 (not in Zabuella Stichel, 1911). The following taxa are best treated as subgenera: Zegris Boisduval, 1836 of Anthocharis Boisduval, Rambur, [Duménil] & Graslin, [1833]; Baltia Moore, 1878 and Pontieuchloia Verity, 1929 of Pontia [Fabricius], 1807; Phrissura Butler, 1870 of Appias Hübner, [1819]; Saletara Distant, 1885 of Catophaga Hübner, 1819; Leodonta Butler, 1870 of Pereute Herrich-Schäffer, 1867; Takashia M. Okano & T. Okano, 1985 of Polycaena Staudinger, 1886; Corrachia Schaus, 1913 of Styx Staudinger, 1876; Ionotus Hall, 2005 and Voltinia Stichel, 1910 of Cremna Doubleday, 1847; Hermathena Hewitson, 1874 of Ithomiola C. & R. Felder, 1865; Lucillella Strand, 1932 of Esthemopsis C. & R. Felder, 1865; Mesenopsis Godman & Salvin, 1886 and Xenandra C. & R. Felder, 1865 of Symmachia Hübner, [1819]; Pirascca J. Hall & Willmott, 1996 of Pterographium Stichel, 1910; Imelda Hewitson, 1870 of Echenais Hübner, [1819]; Calicosama J. Hall & Harvey, 2001 of Behemothia Hall, 2000; Polygrapha Staudinger, 1887 and Fountainea Rydon, 1971 of Anaea Hübner, [1819]; Siderone Hübner, [1823] and Phantos Dias, 2018 of Zaretis Hübner, [1819]; Harsiesis Fruhstorfer, 1911 of Platypthima Rothschild & Jordan, 1905; Vila Kirby, 1871 of Biblis Fabricius, 1807; Diaethria Billberg, 1820 and Perisama Doubleday, 1849 of Callicore Hübner, [1819]; Antigonis C. Felder, 1861 of Haematera Doubleday, 1849; Asterope Hübner, [1819], Nica Hübner, [1826], Peria Kirby, 1871, and Callicorina Smart, 1976 of Temenis Hübner, [1819]; Anthanassa Scudder, 1875, Castilia Higgins, 1981, Telenassa Higgins, 1981, Dagon Higgins, 1981, and Janatella Higgins, 1981 of Eresia Boisduval, 1836; and Wallengrenia Berg, 1897 of Polites Scudder, 1872. The following taxa are junior subjective synonyms: Maniolina Grote, 1897 of Erebiina Tutt, 1896; Melanargiina Wheeler, 1903 of Satyrina Boisduval, 1833; Phyciodina Higgins, 1981 of Melitaeina Herrich-Schäffer, 1843; Cunizza Grote, 1900 of Hesperocharis C. Felder, 1862; Reliquia Ackery, 1975 of Pontia [Fabricius], 1807; Tatochila A. Butler, 1870, Piercolias Staudinger, 1894, Hypsochila Ureta, 1955, Theochila W. D. Field, 1958, Pierphulia W. D. Field, 1958, and Infraphulia W. D. Field, 1958 of Phulia Herrich-Schäffer, 1867; Mesapia Gray, 1856 of Aporia Hübner, [1819]; Catasticta Butler, 1870 of Archonias Hübner, 1827; Sandia Clench & P. Ehrlich, 1960 andXamia Clench, 1961 of Incisalia Scudder, 1872; Hades Westwood, 1851 of Methone Doubleday, 1847; Semomesia Westwood, 1851, Mesophthalma Westwood, 1851, Perophthalma Westwood, 1851 and Leucochimona Stichel, 1909 of Mesosemia Hübner, [1819], Xynias Hewitson, 1874 of Mesenopsis Godman & Salvin, 1886; Stichelia J. Zikán, 1949 of Symmachia Hübner, [1819]; Chimastrum Godman & Salvin, 1886 of Mesene Doubleday, 1847; Alethea Nielsen & Salazar, [2018] of Pirascca J. Hall & Willmott, 1996; Panaropsis J. Hall, 2002 of Pterographium Stichel, 1910; Comphotis Stichel, 1910 of Phaenochitonia Stichel, 1910; Colaciticus Stichel, 1910 of Baeotis Hübner, [1819]; Nahida Kirby, 1871 of Ithomeis Bates, 1862; Machaya Hall & Willmott, 1995 of Pachythone Bates, 1868; Percnodaimon Butler, 1876 and Erebiola Fereday, 1879 of Argyrophenga Doubleday, 1845; Hestinalis Bryk, 1938 of Mimathyma Moore, 1896; Catacore Dillon, 1948 of Diaethria Billberg, 1820; Mesotaenia Kirby, 1871 and Orophila Staudinger, 1886 of Perisama Doubleday, 1849; Paulogramma Dillon, 1948 of Catagramma Boisduval, 1836; Panacea Godman & Salvin, 1883 of Batesia C. Felder & R. Felder, 1862; Napeocles Bates, 1864 of Siproeta Hübner, [1823]; Texola Higgins, 1959 and Dymasia Higgins, 1960 of Microtia H. Bates, 1864; Tisona Higgins, 1981 of Ortilia Higgins, 1981; Abananote Potts, 1943 and Altinote Potts, 1943 of Actinote Hübner, [1819]; Episcada Godman & Salvin, 1879 of Ceratinia Hübner, 1816; and Appia Evans, 1955 of Pompeius Evans, 1955. The following genera are placed in taxonomic hierarchy: Prestonia Schaus, 1920 belongs to Euremini Grote, 1898; Petrocerus Callaghan, 1979 belongs to Theopina Clench, 1955; and Paralasa Moore, 1893 belongs to Ypthimina Reuter, 1896. The following taxa are distinct species rather than subspecies (of species shown in parenthesis): Pyrisitia westwoodii (Boisduval, 1836) (not Pyrisitia dina (Poey, 1832)), Biblis aganisa Boisduval, 1836 (not Biblis hyperia (Cramer, 1779)), Phystis variegata (Röber, 1913) and Phystis pratti (A. Hall, 1935) (not Phystis simois (Hewitson, 1864)), Phocides batabano (Lucas, 1857) and Phocides bicolora (Boddaert, 1783) (not Phocides pigmalion (Cramer, 1779)), Lobotractus mysie (Dyar, 1904) (not Lobotractus valeriana (Plötz, 1881)). Nahida coenoides (Hewitson, 1870) is conspecific with Ithomeis aurantiaca H. Bates, 1862. Additional new and revised combinations are: Teriocolias deva (E. Doubleday, 1847), Teriocolias reticulata (A. Butler, 1871), Hesperocharis leucothea (Molina, 1782), Methone euploea (Hewitson, [1855]), Methone eucerus (Hewitson, 1872), Methone hypophaea (Godman & Salvin, 1878), Methone eubule (R. Felder, 1869), Methone onorata (Hewitson, 1869), Methone authe (Godman, 1903), Methone dolichos (Staudinger, [1887]), Methone baucis (Stichel, 1919), Methone eucrates (Hewitson, 1872), Napaea danforthi A. Warren & Opler, 1999, Napaea dramba (J. Hall, Robbins & Harvey, 2004), Napaea sanarita (Schaus, 1902), Napaea agroeca Stichel, 1910, Napaea tumbesia J. Hall & Lamas, 2001, Napaea umbra (Boisduval, 1870), Napaea phryxe (C. & R. Felder, 1865), Napaea cebrenia (Hewitson, [1873]), Napaea loxicha (R.G. Maza & J. Maza, 2016), Napaea maya (J. Maza & Lamas, 2016), Napaea necaxa (R.G. Maza & J. Maza, 2018), Napaea totonaca (R.G. Maza & J. Maza, 2016), Mesene aeolia (Bates, 1868), Pterographium hypochloris (Bates, 1868), Phaenochitonia florus (Fabricius, 1793), Ourocnemis carausius (Westwood, 1851), Ourocnemis principalis (Hopffer, 1874), Ourocnemis renaldus (Stoll, 1790), and Ourocnemis aerosus (Stichel, 1924), Hallonympha maculosa (Bates, 1868), Exoplisia aphanis (Stichel, 1910), Phystis fontus (A. Hall, 1928), Phocides batabano okeechobee (Worthington, 1881), and Phocides batabano batabanoides (W. Holland, 1902). Finally, we confirm the combination Zabuella castanea (Prittwitz, 1865) and find Pyrgus centaureae dzekh Gorbunov, 2007 as a new subspecies for North America.