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, (type genus Aurivillius, 1898); in subfamily Riodininae Grote, 1895: Callistiumini Grishin, (type genus Stichel, 1911); in subfamily Nymphalinae Rafinesque, 1815: Pycinini Grishin, (type genus Doubleday 1849), Rhinopalpini Grishin, (type genus C. & R. Felder 1860), Kallimoidini Grishin, (type genus Shirôzu & Nakanishi 1984), Vanessulini Grishin, (type genus Dewitz 1887), and Doleschalliaini Grishin, (type genus C. & R. Felder 1860); in tribe Mesosemiini Bates, 1859: Grishin, (type genus Westwood, 1851); in tribe Satyrini Boisduval, 1833: Callerebiina Grishin, (type genus Butler, 1867), Gyrocheilina Grishin, (type genus Butler, 1867), and Calistina Grishin, (type genus Hübner, [1823]); in subfamily Euselasiinae Kirby, 1871: Grishin, (type species Hewitson, [1853]), Grishin, (type species Herrich-Schäffer, [1853]), Grishin, (type species Hewitson, 1856), Grishin, (type species Callaghan, 1999), and Grishin, (type species Hewitson, 1856); in subtribe Mesosemiina Bates, 1859: Grishin, (type species Cramer, 1776) and Grishin, (type species Cramer, 1777); in tribe Symmachiini Reuter, 1896: Grishin, (type species Hewitson, 1870) and Grishin, (type species Stichel, 1910); in tribe Riodinini Grote, 1895: Grishin, (type species Bates, 1868), Grishin, (type species Hewitson, 1866), Grishin, (type species Schaus, 1902), and Grishin, (type species Hewitson, 1867); in subtribe Zabuellina Seraphim, Freitas & Kaminski, 2018: Grishin, (type species Bates, 1868); Grishin, (type species Cramer, 1777, parent genus Hübner, [1821]), Grishin, (type species Beutelspacher, 1975, parent genus Scopoli, 1777), Grishin, (type species Cramer, 1775, parent genus Hübner, [1819]), Grishin, (type species Boisduval, 1852, parent genus [Fabricius], 1807), Grishin, (type species [sic] Edwards, 1877, parent genus Billberg, 1820), Grishin, (type species Strecker, 1880, parent genus Dalman, 1816), and in genus Boisduval, 1836: Grishin, (type species Hewitson, 1864), Grishin, (type species Hewitson, 1872), and Grishin, (type species 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: Swainson, 1821 (not in Hübner, [1819]), Hübner, [1819] and Hübner, [1819] (not in Hübner, [1819]), Strand, 1932 (not in Stichel, 1910), Doubleday, 1847 (not in Hübner, [1819]), and Penz & DeVries, 2006 (not in Stichel, 1911). The following taxa are best treated as subgenera: Boisduval, 1836 of Boisduval, Rambur, [Duménil] & Graslin, [1833]; Moore, 1878 and Verity, 1929 of [Fabricius], 1807; Butler, 1870 of Hübner, [1819]; Distant, 1885 of Hübner, 1819; Butler, 1870 of Herrich-Schäffer, 1867; M. Okano & T. Okano, 1985 of Staudinger, 1886; Schaus, 1913 of Staudinger, 1876; Hall, 2005 and Stichel, 1910 of Doubleday, 1847; Hewitson, 1874 of C. & R. Felder, 1865; Strand, 1932 of C. & R. Felder, 1865; Godman & Salvin, 1886 and C. & R. Felder, 1865 of Hübner, [1819]; J. Hall & Willmott, 1996 of Stichel, 1910; Hewitson, 1870 of Hübner, [1819]; J. Hall & Harvey, 2001 of Hall, 2000; Staudinger, 1887 and Rydon, 1971 of Hübner, [1819]; Hübner, [1823] and Dias, 2018 of Hübner, [1819]; Fruhstorfer, 1911 of Rothschild & Jordan, 1905; Kirby, 1871 of Fabricius, 1807; Billberg, 1820 and Doubleday, 1849 of Hübner, [1819]; C. Felder, 1861 of Doubleday, 1849; Hübner, [1819], Hübner, [1826], Kirby, 1871, and Smart, 1976 of Hübner, [1819]; Scudder, 1875, Higgins, 1981, Higgins, 1981, Higgins, 1981, and Higgins, 1981 of Boisduval, 1836; and Berg, 1897 of 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; Grote, 1900 of C. Felder, 1862; Ackery, 1975 of [Fabricius], 1807; A. Butler, 1870, Staudinger, 1894, Ureta, 1955, W. D. Field, 1958, W. D. Field, 1958, and W. D. Field, 1958 of Herrich-Schäffer, 1867; Gray, 1856 of Hübner, [1819]; Butler, 1870 of Hübner, 1827; Clench & P. Ehrlich, 1960 and Clench, 1961 of Scudder, 1872; Westwood, 1851 of Doubleday, 1847; Westwood, 1851, Westwood, 1851, Westwood, 1851 and Stichel, 1909 of Hübner, [1819], Hewitson, 1874 of Godman & Salvin, 1886; J. Zikán, 1949 of Hübner, [1819]; Godman & Salvin, 1886 of Doubleday, 1847; Nielsen & Salazar, [2018] of J. Hall & Willmott, 1996; J. Hall, 2002 of Stichel, 1910; Stichel, 1910 of Stichel, 1910; Stichel, 1910 of Hübner, [1819]; Kirby, 1871 of Bates, 1862; Hall & Willmott, 1995 of Bates, 1868; Butler, 1876 and Fereday, 1879 of Doubleday, 1845; Bryk, 1938 of Moore, 1896; Dillon, 1948 of Billberg, 1820; Kirby, 1871 and Staudinger, 1886 of Doubleday, 1849; Dillon, 1948 of Boisduval, 1836; Godman & Salvin, 1883 of C. Felder & R. Felder, 1862; Bates, 1864 of Hübner, [1823]; Higgins, 1959 and Higgins, 1960 of H. Bates, 1864; Higgins, 1981 of Higgins, 1981; Potts, 1943 and Potts, 1943 of Hübner, [1819]; Godman & Salvin, 1879 of Hübner, 1816; and Evans, 1955 of Evans, 1955. The following genera are placed in taxonomic hierarchy: Schaus, 1920 belongs to Euremini Grote, 1898; Callaghan, 1979 belongs to Theopina Clench, 1955; and Moore, 1893 belongs to Ypthimina Reuter, 1896. The following taxa are distinct species rather than subspecies (of species shown in parenthesis): (Boisduval, 1836) (not (Poey, 1832)), Boisduval, 1836 (not (Cramer, 1779)), (Röber, 1913) and (A. Hall, 1935) (not (Hewitson, 1864)), (Lucas, 1857) and (Boddaert, 1783) (not (Cramer, 1779)), (Dyar, 1904) (not (Plötz, 1881)). (Hewitson, 1870) is conspecific with H. Bates, 1862. Additional new and revised combinations are: (E. Doubleday, 1847), (A. Butler, 1871), (Molina, 1782), (Hewitson, [1855]), (Hewitson, 1872), (Godman & Salvin, 1878), (R. Felder, 1869), (Hewitson, 1869), (Godman, 1903), (Staudinger, [1887]), (Stichel, 1919), (Hewitson, 1872), A. Warren & Opler, 1999, (J. Hall, Robbins & Harvey, 2004), (Schaus, 1902), Stichel, 1910, J. Hall & Lamas, 2001, (Boisduval, 1870), (C. & R. Felder, 1865), (Hewitson, [1873]), (R.G. Maza & J. Maza, 2016), (J. Maza & Lamas, 2016), (R.G. Maza & J. Maza, 2018), (R.G. Maza & J. Maza, 2016), (Bates, 1868), (Bates, 1868), (Fabricius, 1793), (Westwood, 1851), (Hopffer, 1874), (Stoll, 1790), and (Stichel, 1924), (Bates, 1868), (Stichel, 1910), (A. Hall, 1928), (Worthington, 1881), and (W. Holland, 1902). Finally, we confirm the combination (Prittwitz, 1865) and find Gorbunov, 2007 as a new subspecies for North America.