Genomics-guided refinement of butterfly taxonomy.

Abstract

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, <b>trib. n.</b> (type genus <i>Calopieris</i> Aurivillius, 1898); in subfamily Riodininae Grote, 1895: Callistiumini Grishin, <b>trib. n.</b> (type genus <i>Callistium</i> Stichel, 1911); in subfamily Nymphalinae Rafinesque, 1815: Pycinini Grishin, <b>trib. n.</b> (type genus <i>Pycina</i> Doubleday 1849), Rhinopalpini Grishin, <b>trib. n.</b> (type genus <i>Rhinopalpa</i> C. & R. Felder 1860), Kallimoidini Grishin, <b>trib. n.</b> (type genus <i>Kallimoides</i> Shirôzu & Nakanishi 1984), Vanessulini Grishin, <b>trib. n.</b> (type genus <i>Vanessula</i> Dewitz 1887), and Doleschalliaini Grishin, <b>trib. n.</b> (type genus <i>Doleschallia</i> C. & R. Felder 1860); in tribe Mesosemiini Bates, 1859: <i>Eunogyrina</i> Grishin, <b>subtrib. n.</b> (type genus <i>Eunogyra</i> Westwood, 1851); in tribe Satyrini Boisduval, 1833: Callerebiina Grishin, <b>subtrib. n.</b> (type genus <i>Callerebia</i> Butler, 1867), Gyrocheilina Grishin, <b>subtrib. n.</b> (type genus <i>Gyrocheilus</i> Butler, 1867), and Calistina Grishin, <b>subtrib. n.</b> (type genus <i>Calisto</i> Hübner, [1823]); in subfamily Euselasiinae Kirby, 1871: <i>Pelolasia</i> Grishin, <b>gen. n.</b> (type species <i>Eurygona pelor</i> Hewitson, [1853]), <i>Myselasia</i> Grishin, <b>gen. n.</b> (type species <i>Eurygona mys</i> Herrich-Schäffer, [1853]), <i>Eurylasia</i> Grishin, <b>gen. n.</b> (type species <i>Eurygona euryone</i> Hewitson, 1856), <i>Maculasia</i> Grishin, <b>gen. n.</b> (type species <i>Euselasia albomaculiga</i> Callaghan, 1999), and <i>Eugelasia</i> Grishin, <b>gen. n.</b> (type species <i>Eurygona eugeon</i> Hewitson, 1856); in subtribe Mesosemiina Bates, 1859: <i>Ectosemia</i> Grishin, <b>gen. n.</b> (type species <i>Papilio eumene</i> Cramer, 1776) and <i>Endosemia</i> Grishin, <b>gen. n.</b> (type species <i>Papilio ulrica</i> Cramer, 1777); in tribe Symmachiini Reuter, 1896: <i>Tigria</i> Grishin, <b>gen. n.</b> (type species <i>Mesene xypete</i> Hewitson, 1870) and <i>Asymma</i> Grishin, <b>gen. n.</b> (type species <i>Symmachia virgatula</i> Stichel, 1910); in tribe Riodinini Grote, 1895: <i>Putridivora</i> Grishin, <b>gen. n.</b> (type species <i>Charis argyrea</i> Bates, 1868), <i>Chadia</i> Grishin, <b>gen. n.</b> (type species <i>Charis cadytis</i> Hewitson, 1866), <i>Inkana</i> Grishin, <b>gen. n.</b> (type species <i>Charis incoides</i> Schaus, 1902), and <i>Oco</i> Grishin, <b>gen. n.</b> (type species <i>Symmachia ocellata</i> Hewitson, 1867); in subtribe Zabuellina Seraphim, Freitas & Kaminski, 2018: <i>Teenie</i> Grishin, <b>gen. n.</b> (type species <i>Calydna tinea</i> Bates, 1868); <i>Boreographium</i> Grishin, <b>subgen. n.</b> (type species <i>Papilio marcellus</i> Cramer, 1777, parent genus <i>Eurytides</i> Hübner, [1821]), <i>Esperourus</i> Grishin, <b>subgen. n.</b> (type species <i>Papilio esperanza</i> Beutelspacher, 1975, parent genus <i>Pterourus</i> Scopoli, 1777), <i>Hyppasonia</i> Grishin, <b>subgen. n.</b> (type species <i>Papilio hyppason</i> Cramer, 1775, parent genus <i>Heraclides</i> Hübner, [1819]), <i>Sisymbria</i> Grishin, <b>subgen. n.</b> (type species <i>Pieris sisymbrii</i> Boisduval, 1852, parent genus <i>Pontia</i> [Fabricius], 1807), <i>Greenie</i> Grishin, <b>subgen. n.</b> (type species <i>Thecla sheridonii</i> [sic] Edwards, 1877, parent genus <i>Callophrys</i> Billberg, 1820), <i>Magda</i> Grishin, <b>subgen. n.</b> (type species <i>Erebia magdalena</i> Strecker, 1880, parent genus <i>Erebia</i> Dalman, 1816), and in genus <i>Eresia</i> Boisduval, 1836: <i>Notilia</i> Grishin, <b>subgen. n.</b> (type species <i>Eresia orthia</i> Hewitson, 1864), <i>Levinata</i> Grishin, <b>subgen. n.</b> (type species <i>Eresia levina</i> Hewitson, 1872), and <i>Ithra</i> Grishin, <b>subgen. n.</b> (type species <i>Phyciodes ithra</i> 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: <i>Terias</i> Swainson, 1821 (not in <i>Eurema</i> Hübner, [1819]), <i>Erythia</i> Hübner, [1819] and <i>Marmessus</i> Hübner, [1819] (not in <i>Euselasia</i> Hübner, [1819]), <i>Eucorna</i> Strand, 1932 (not in <i>Voltinia</i> Stichel, 1910), <i>Cremna</i> Doubleday, 1847 (not in <i>Napaea</i> Hübner, [1819]), and <i>Hallonympha</i> Penz & DeVries, 2006 (not in <i>Zabuella</i> Stichel, 1911). The following taxa are best treated as subgenera: <i>Zegris</i> Boisduval, 1836 of <i>Anthocharis</i> Boisduval, Rambur, [Duménil] & Graslin, [1833]; <i>Baltia</i> Moore, 1878 and <i>Pontieuchloia</i> Verity, 1929 of <i>Pontia</i> [Fabricius], 1807; <i>Phrissura</i> Butler, 1870 of <i>Appias</i> Hübner, [1819]; <i>Saletara</i> Distant, 1885 of <i>Catophaga</i> Hübner, 1819; <i>Leodonta</i> Butler, 1870 of <i>Pereute</i> Herrich-Schäffer, 1867; <i>Takashia</i> M. Okano & T. Okano, 1985 of <i>Polycaena</i> Staudinger, 1886; <i>Corrachia</i> Schaus, 1913 of <i>Styx</i> Staudinger, 1876; <i>Ionotus</i> Hall, 2005 and <i>Voltinia</i> Stichel, 1910 of <i>Cremna</i> Doubleday, 1847; <i>Hermathena</i> Hewitson, 1874 of <i>Ithomiola</i> C. & R. Felder, 1865; <i>Lucillella</i> Strand, 1932 of <i>Esthemopsis</i> C. & R. Felder, 1865; <i>Mesenopsis</i> Godman & Salvin, 1886 and <i>Xenandra</i> C. & R. Felder, 1865 of <i>Symmachia</i> Hübner, [1819]; <i>Pirascca</i> J. Hall & Willmott, 1996 of <i>Pterographium</i> Stichel, 1910; <i>Imelda</i> Hewitson, 1870 of <i>Echenais</i> Hübner, [1819]; <i>Calicosama</i> J. Hall & Harvey, 2001 of <i>Behemothia</i> Hall, 2000; <i>Polygrapha</i> Staudinger, 1887 and <i>Fountainea</i> Rydon, 1971 of <i>Anaea</i> Hübner, [1819]; <i>Siderone</i> Hübner, [1823] and <i>Phantos</i> Dias, 2018 of <i>Zaretis</i> Hübner, [1819]; <i>Harsiesis</i> Fruhstorfer, 1911 of <i>Platypthima</i> Rothschild & Jordan, 1905; <i>Vila</i> Kirby, 1871 of <i>Biblis</i> Fabricius, 1807; <i>Diaethria</i> Billberg, 1820 and <i>Perisama</i> Doubleday, 1849 of <i>Callicore</i> Hübner, [1819]; <i>Antigonis</i> C. Felder, 1861 of <i>Haematera</i> Doubleday, 1849; <i>Asterope</i> Hübner, [1819], <i>Nica</i> Hübner, [1826], <i>Peria</i> Kirby, 1871, and <i>Callicorina</i> Smart, 1976 of <i>Temenis</i> Hübner, [1819]; <i>Anthanassa</i> Scudder, 1875, <i>Castilia</i> Higgins, 1981, <i>Telenassa</i> Higgins, 1981, <i>Dagon</i> Higgins, 1981, and <i>Janatella</i> Higgins, 1981 of <i>Eresia</i> Boisduval, 1836; and <i>Wallengrenia</i> Berg, 1897 of <i>Polites</i> 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; <i>Cunizza</i> Grote, 1900 of <i>Hesperocharis</i> C. Felder, 1862; <i>Reliquia</i> Ackery, 1975 of <i>Pontia</i> [Fabricius], 1807; <i>Tatochila</i> A. Butler, 1870, <i>Piercolias</i> Staudinger, 1894, <i>Hypsochila</i> Ureta, 1955, <i>Theochila</i> W. D. Field, 1958, <i>Pierphulia</i> W. D. Field, 1958, and <i>Infraphulia</i> W. D. Field, 1958 of <i>Phulia</i> Herrich-Schäffer, 1867; <i>Mesapia</i> Gray, 1856 of <i>Aporia</i> Hübner, [1819]; <i>Catasticta</i> Butler, 1870 of <i>Archonias</i> Hübner, 1827; <i>Sandia</i> Clench & P. Ehrlich, 1960 and<i>Xamia</i> Clench, 1961 of <i>Incisalia</i> Scudder, 1872; <i>Hades</i> Westwood, 1851 of <i>Methone</i> Doubleday, 1847; <i>Semomesia</i> Westwood, 1851, <i>Mesophthalma</i> Westwood, 1851, <i>Perophthalma</i> Westwood, 1851 and <i>Leucochimona</i> Stichel, 1909 of <i>Mesosemia</i> Hübner, [1819], <i>Xynias</i> Hewitson, 1874 of <i>Mesenopsis</i> Godman & Salvin, 1886; <i>Stichelia</i> J. Zikán, 1949 of <i>Symmachia</i> Hübner, [1819]; <i>Chimastrum</i> Godman & Salvin, 1886 of <i>Mesene</i> Doubleday, 1847; <i>Alethea</i> Nielsen & Salazar, [2018] of <i>Pirascca</i> J. Hall & Willmott, 1996; <i>Panaropsis</i> J. Hall, 2002 of <i>Pterographium</i> Stichel, 1910; <i>Comphotis</i> Stichel, 1910 of <i>Phaenochitonia</i> Stichel, 1910; <i>Colaciticus</i> Stichel, 1910 of <i>Baeotis</i> Hübner, [1819]; <i>Nahida</i> Kirby, 1871 of <i>Ithomeis</i> Bates, 1862; <i>Machaya</i> Hall & Willmott, 1995 of <i>Pachythone</i> Bates, 1868; <i>Percnodaimon</i> Butler, 1876 and <i>Erebiola</i> Fereday, 1879 of <i>Argyrophenga</i> Doubleday, 1845; <i>Hestinalis</i> Bryk, 1938 of <i>Mimathyma</i> Moore, 1896; <i>Catacore</i> Dillon, 1948 of <i>Diaethria</i> Billberg, 1820; <i>Mesotaenia</i> Kirby, 1871 and <i>Orophila</i> Staudinger, 1886 of <i>Perisama</i> Doubleday, 1849; <i>Paulogramma</i> Dillon, 1948 of <i>Catagramma</i> Boisduval, 1836; <i>Panacea</i> Godman & Salvin, 1883 of <i>Batesia</i> C. Felder & R. Felder, 1862; <i>Napeocles</i> Bates, 1864 of <i>Siproeta</i> Hübner, [1823]; <i>Texola</i> Higgins, 1959 and <i>Dymasia</i> Higgins, 1960 of <i>Microtia</i> H. Bates, 1864; <i>Tisona</i> Higgins, 1981 of <i>Ortilia</i> Higgins, 1981; <i>Abananote</i