Advancing butterfly systematics through genomic analysis.

Within the framework of an ongoing comparative genomic study of global butterfly diversity, we construct phylogenetic trees combining all protein-coding genes assembled from the whole genome shotgun data. When viewed in the context of current taxonomy and phenotypic knowledge, the genome-wide phylogeny points to further advances in butterfly systematics, which are presented here. We assign major clades with comparable levels of genetic divergence to the same taxonomic rank and apply criteria involving relative population divergence and gene flow to define species boundaries. As a result, one genus, 13 subgenera, 62 species, and five subspecies are proposed as new (type species in original combinations or type localities are given in parentheses): a genus Grishin, ( Elwes, 1893) in Deudorigini Doherty, 1886, Lycaenidae [Leach], [1815]; subgenera: in Riodinidae Grote, 1895 (1827): Grishin, ( Hewitson, 1866) and Grishin, ( Westwood, 1851) of Hewitson, 1861, Grishin, ( Godman, 1903) of H. Bates, 1868, Grishin, ( Fabricius, 1793) of J. Hall, 2018, Grishin, ( Hewitson, 1872) of H. Bates, 1868, Grishin, ( Mengel, 1902) of Westwood, 1851, and Grishin, ( Hewitson, 1875) of H. Bates, 1868; in Lycaenidae: Grishin, ( Hopffer, 1855) of Hewitson, 1865, Grishin, ( C. Felder, 1860) of Hewitson, 1863, and Grishin, ( Fabricius, 1793) of F. Moore, 1881; and in Hesperiidae Latreille, 1809: Grishin, ( Bell, 1935), Grishin, ( Bremer & Grey, 1853), and Grishin, ( W. H. Edwards, 1873) of Scudder, 1872; species: in Riodinidae: Grishin, (Mexico: Colima), Grishin, (USA: Texas), Grishin, (Mexico: Sonora), Grishin, (Mexico: Chiapas), () Grishin, (Argentina), () Grishin, (Uruguay), () Grishin, (Panama: Chiriquí), and Grishin, (Peru: San Martin) and in Hesperiidae: Grishin, (Ecuador: Pichincha), Grishin, (Brazil: Amazonas), Grishin, (Guyana), Grishin, (eastern Colombia), Grishin, (Brazil: Pará), Grishin, (Peru: Madre de Dios), Grishin, (Brazil: Amazonas), Grishin, (Brazil: Pará), Grishin, (Mexico: Chiapas), Grishin, (Panama: Darién), Grishin, (Venezuela: Aragua), Grishin, (Ecuador: Napo), Grishin, (Colombia: Bogotá), () Grishin, (Brazil: Santa Catarina), () Grishin, (Venezuela: Merida), () Grishin, (USA: Texas), () Grishin, (Panama: Panamá), () Grishin, (Peru: Piura), () Grishin, (Brazil: Rondônia), () Grishin, (Panama: Darién), () Grishin, (Brazil: Santa Catarina), () Grishin, (likely Southeast or South Brazil), () Grishin, (Peru: Madre de Dios), () Grishin, (Mexico: Veracruz), () Grishin, (Mexico: Chiapas), () Grishin, (Colombia: Cauca), () Grishin, (Costa Rica: Limón), () Grishin, (Brazil: Santa Catarina), () Grishin, (Ecuador: Esmeraldas), () Grishin, (Panama: Panamá Oeste), () Grishin, (Panama: Panamá), () Grishin, (Peru: Cuzco), () Grishin, (Brazil: Rio de Janeiro), () Grishin, (Brazil: Rondônia), Grishin, (Peru: Cuzco), Grishin, (Colombia: Tolima), Grishin, (Argentina: Salta), () Grishin, (Peru: Amazonas), Grishin, (Ghana: Oti), () Grishin, (Argentina: Jujuy), () Grishin, (Peru: Junín), () Grishin, (Peru: Puno), () Grishin, (Honduras: San Pedro Sula), Grishin, (El Salvador: La Libertad), Grishin, (Mexico: Oaxaca), Grishin, (Mexico: Tamaulipas), Grishin, (Argentina: Salta), Grishin, (Peru: Cuzco), () Grishin, (Ecuador: Santo Domingo de los Tsáchilas), Grishin, (Costa Rica: Heredia), Grishin, (Ecuador: Napo), Grishin, (Honduras: San Pedro Sula), Grishin, (Peru: Loreto), and Grishin, (Peru: Madre de Dios); and subspecies: in Nymphalidae Rafinesque, 1815: () Grishin, (Russia: Chukotka) and in Hesperiidae: () Grishin, (Ecuador: Esmeraldas), Grishin, (USA: Texas), Grishin, (Mexico: Hidalgo), and Grishin, (Mexico: Baja California Norte). The following are valid genera: Bates, 1868, (not Hübner, [1819]) and F. Moore, 1881, (not Hewitson, 1863). The following are valid subgenera, not genera or synonyms: F. Moore, 1901, of Hewitson, 1861, J. Hall & Harvey, 2002, of Westwood, 1851, Hübner, [1819], of Hübner, [1807], and Callaghan, 1985, of H. Bates, 1868. The following are valid species, not subspecies or synonyms: () Ménétriés, 1859, (not () (Tauscher, 1806) or () Ménétriés, 1847), () B. Warren, 1936, (not () Ménétriés, 1859), Grishin, 2024, (not Schaus, 1890), (Godman & Salvin, 1886), (not (Hewitson, 1867)), (Linnaeus, 1763), and Evans, 1952, (not (Linnaeus, 1758)), Mabille, 1898, , Mabille & Boullet, 1919, , M. Hering, 1925, , and Salazar & Vargas, [2017], (not Godman & Salvin, 1879), () (Williams, 1931), (not () (Butler, 1870)), () (Röber, 1925), (not () (Mabille, 1877)), () (Plötz, 1881), (not () (C. Felder & R. Felder, 1867)), () (H. Freeman, 1969), (not () (Plötz, 1881), ), () (Herrich-Schäffer, 1869), and () (Evans, 1952), (not () (Herrich-Schäffer, 1869)), () (Stoll, 1781), , () (Evans, 1952), , and () (Evans, 1952), (not () (Cramer, 1780)), () (Evans, 1952), and () (Hewitson, 1876), (not () Staudinger, 1875), () (Mabille, 1888), (not () (Herrich-Schäffer, 1869)), () (Mabille, 1889), and () (Plötz, 1882), (not () Herrich-Schäffer, 1870), () Mabille, 1889, , () Williams & Bell, 1939, , and () Evans, 1953, (not () Plötz, 1882), () Steinhauser, 1989, (not Schaus, 1902, ), Swinhoe, 1885, (not F. Moore, 1879), () (Weymer, 1890), (not () (Erschoff, 1876)), () (Mabille, 1875), (not () (Hewitson, 1874)), () Evans, 1953, (not () Hayward, 1942), (Draudt, 1923), (not (Hewitson, 1870)), and (Herrich-Schäffer, 1869), , (Möschler, 1877), , and (Plötz, 1886), (not (Herrich-Schäffer, 1869). The following are valid subspecies, not species or synonyms: () Staudinger, 1894 (not () Ménétriés, 1859), () R. Williams & E. Bell, 1939, (not a synonym of () (Mabille, 1889), ), () Ureta, 1956, , () Draudt, 1923, , and (Mabille, 1891), (not a synonym of (Herrich-Schäffer, 1869)). Plötz, 1883, is a junior objective synonym of Linnaeus, 1763. The following are junior subjective synonyms, new or transferred between taxa: Röber, 1903, of Bates, 1868, , (not of Hübner, [1819]), Hewitson, 1876, of (Hewitson, 1867), Plötz, 1882, of () Staudinger, 1875, Ehrmann, 1918, of () (Herrich-Schäffer, 1869) (not () (Stoll, 1790), Plötz, 1882, and Schaus, 1902, of () Plötz, 1882, Capronnier, 1874, of (Herrich-Schäffer, 1869) (not of (Herrich-Schäffer, 1869)), Shuey, 2024, of (Herrich-Schäffer, 1869), , and Godman, 1901, and Draudt, 1923, of (Mabille, 1891), (not of (Herrich-Schäffer, 1869)). (W. Holland, 1920) (not Hewitson, 1863) is a genus-species and the following are new species-subspecies combinations: () Mabille, 1903 (not () (Cramer, 1780)), () Evans, 1952 (not () (Stoll, 1790), and Benyamini, 1990, (not (Trimen, 1862)). are designated for 24 taxa: Plötz, 1883 (the Amazonian region), Swainson, 1831 (South America), Godman & Salvin, 1879 (Nicaragua), Plötz, 1881 (Mexico, likely south-central or southern), (Herrich-Schäffer, 1869) (Brazil), Staudinger, 1875 (Panama: Chiriquí), Plötz, 1882 (likely Panama: Chiriquí), Plötz, 1882 (likely Venezuela), Plötz, 1882 (Brazil: likely São Paulo), Plötz, 1882 (Brazil: likely Rio de Janeiro), Mabille, 1891 (Brazil: likely Rio de Janeiro), Schaus, 1902 (Brazil: Rio de Janeiro), Mabille, 1889 (Panama: Chiriquí), Godman & Salvin, 1897 (Mexico: Nayarit), Dyar, 1924 (Mexico: Colima), Weymer, 1890 (Bolivia), Godman & Salvin, 1896 (Mexico: Guerrero), Herrich-Schäffer, 1869 (Southeast and South Brazil), Herrich-Schäffer, 1869 (the Amazonian region, likely Suriname), Hewitson, 1876 (Brazil: Bahia), Möschler, 1877 (Suriname), Mabille, 1891 (Panama: Chiriquí), Godman, 1901 (Panama: Chiriquí), and Capronnier, 1874 (Southeast or South Brazil). are designated for six taxa: Linnaeus, 1758 (Suriname), Linnaeus, 1763 (the Amazonian region), Linnaeus, 1763 (French Guiana), Swainson, 1831 (French Guiana), Plötz, 1881 (Southeast or South Brazil), and Plötz, 1886 (Panama: Panama). The type locality of Draudt, 1923 is not in South Brazil, but likely in Panama: Chiriquí, as deduced by genomic comparison. Pelham, 2008 and (W. H. Edwards, 1870) may be sympatric in British Columbia, Canada, and Grishin, 2023 is sympatric with Grishin, 2023 in Monteverde, Costa Rica. Additional specimens of () Grishin, 2023 confirm it as a species-level taxon, and Plötz, 1882 is confirmed as a junior subjective synonym of () (Mabille, 1877) by further genomic sequencing. The holotype of () Grishin, 2023 is illustrated after being spread. Curiously, (Godman, 1900) and Austin & A. Warren, 2009 lack overall genetic differentiation typical of species-level taxa. Furthermore, preliminary taxonomic lists of Hübner, [1819] and () Scudder, 1889 (from the clade analyzed in this work) are given. Finally, unless stated otherwise, all subgenera, species, subspecies, and synonyms of mentioned genera, subgenera, and species are transferred together with their parent taxa, and taxa not mentioned in this work remain as previously classified.