Genomic analysis reveals new species and subspecies of butterflies.

Large-scale genomic sequencing of butterfly taxa reveals new findings that are presented here. While we focus on detecting species by comparative genomics and define subspecies as groups of populations genetically differentiated from each other but not as strongly as species (i.e., subspecies as "species in the making"), we report other adjustments to butterfly classification. As a result, 4 subgenera, 11 species, and 6 subspecies are proposed as new. New subgenera are: Grishin, (type species Linnaeus, 1758, genus Schrank, 1801) in Pieridae Swainson, 1820 and Grishin, (type species ? Herrich-Schäffer, [1853], genus E. Doubleday, 1847), Grishin, (type species H. Bates, 1868, genus E. Doubleday, 1847), and Grishin, (type species Stichel, 1910, genus Stichel, 1910) in Riodinidae Grote, 1895 (1827). New species are (type localities in parenthesis): Grishin, (USA: Texas, Duval Co.) in Nymphalidae Rafinesque, 1815; Grishin, (Panama: Canal Zone), Grishin, (Panama: Panama), and Grishin, (Bolivia: La Paz) in Riodinidae; and Grishin, (Colombia: Río Dagua), Grishin, (Ecuador: Canelos), () Grishin, (Cuba: Havana), Grishin, (Guyana: Essequibo), Grishin, (Mexico: Guerrero), Grishin, (Panama: Chiriquí), and Grishin, (Mexico: Chiapas) in Hesperiidae Latreille, 1809. New subspecies are (type localities in parenthesis): Grishin, (Mexico: Chihuahua) in Nymphalidae and Grishin, (USA: Arizona, Coconino Co.), Grishin, (USA: Utah, San Juan Co.), Grishin, (USA: Nevada, Nye Co.), Grishin, (USA: Arizona, Coconino Co.), and Grishin, (Mexico: Baja California Sur) in Hesperiidae. Furthermore, we confirm 1 genus, resurrect 2 subgenera, change the rank of 3 genera to subgenera, synonymize 3 genera, 1 species, and 1 subspecies, and present evidence to support 17 taxa (1 confirmed) as species instead of subspecies or synonyms. Namely, we confirm Hirowatari, 1992 as a valid genus, not a junior subjective synonym of [Fabricius], 1807 (Lycaenidae); treat H. Huang, 1995 and Verity, 1947 as subgenera (not genera or synonyms) of Schrank, 1801 (Pieridae); regard these genera as subgenera: G. Henning & S Henning, 1996 of van Son, 1955 (Nymphalidae), Westwood, 1851 of Westwood, 1851 (Riodinidae), and Hirowatari, 1992 of [Fabricius], 1807 (Lycaenidae); and propose that the following are junior subjective synonyms, not genera, species, or subspecies: Della Bruna, Gallo, Lucarelli & Sbordoni, 2000 of Nakatani, 2017 in Nymphalidae: Satyrini: Ypthimina Reuter, 1896, new placement (not Erebiina Tutt, 1896), Hemming, 1964 of Fruhstorfer, 1922 and Bethune-Baker, 1908 of Bethune-Baker, 1908 in Lycaenidae, and Hübner, [1819] of (Stoll, 1781) and Stichel, 1910 of (Stichel, 1910) in Riodinidae. The following taxa are species, not subspecies or synonyms: (Fabricius, 1777) (not (Linnaeus, 1758)) and (Crowley, 1887) (not (Linnaeus, 1764)) in Pieridae; (Hemming, 1934) (not (E. Aaron, [1885])) and (W. H. Edwards, 1878) (not (Ménétriés, 1855)) in Nymphalidae; (J. Zikán, 1952) (not (Cramer, 1776)), (Stichel, 1910) (not (Godman & Salvin, 1886)), Stichel, 1915 (not (Geyer, 1837)), Clench, 1972 (not Staudinger, 1888), and (Hewitson, 1865) (not Hübner, [1819]) in Riodinidae; and () (Weeks, 1901) (confirmed as not () (A. Butler, 1870)), Schaus, 1902 (not (Prittwitz, 1868)), (Plötz, 1883) (not (Snellen, 1872)), Mabille, 1877 (not (Westwood, 1852)), (W. H. Edwards, 1865) and J. Scott, 1981 (not (Boisduval, 1852)), and (Herrich-Schäffer, 1869) and (Evans, 1955) (not (W. H. Edwards, 1869)) in Hesperiidae. In addition, we propose new genus-species combinations: (Hewitson, 1858) (not Westwood, 1851), (Hewitson, 1873) and (Stichel, 1915) (not Hübner, [1819]), (Lathy, 1926), (H. Bates, 1868) and (Schaus, 1913) (not Hübner, 1819), (Stichel, 1930) (not Hübner, [1819]), (Herrich-Schäffer, [1853]) and (H. Bates, 1868) (not Westwood, 1851), (Stichel, 1910) (not Westwood, 1851), W. Holland, 1891 (not Hübner, [1819]), and (de Jong, 1983) (not Godman, 1900); new species-subspecies combination J. Emmel & T. Emmel, 1998 (not (Boisduval, 1852)); and transfer junior subjective synonyms between taxa: var. W. G. Wright, 1905 of E. Doubleday, 1847 (not of W. H. Edwards, 1870), Plötz, 1881 of (Scudder, 1872) (not of (Linnaeus, 1758)), Strand, 1921 of Mabille, 1903 (not of (Plötz, 1884)), Schaus, 1902 of (Prittwitz, 1868) (not of (Möschler, 1877)), and W. H. Edwards, 1883 of (W. H. Edwards, 1881) (not of (Boisduval, 1852)). Furthermore, by finding an additional specimen, we confirm Grishin, 2023 as a species-level taxon using genomic analysis; we conclude that populations of Leussler, 1938 in most of New Mexico and the White Mountains, Arizona are the nominal subspecies, not Lindsey, 1940; and report a natural hybrid between (W. H. Edwards, [1878]) and (Tinkham, 1944). are designated for 6 names: Bang-Haas, 1927 (type locality in China: Gansu) in Nymphalidae; Stichel, 1910 (type locality in Peru: Pozuzo) and Stichel, 1910 (type locality Bolivia: La Paz, Farinas) in Riodinidae; and Plötz, 1881 (type locality in the Antilles), Plötz, 1883 (type locality likely in the USA, not Guatemala, as deduced by genomic comparison), confirming this name as a junior subjective synonym of (Boisduval & Le Conte, [1837]), and ? Schaus, 1913 (type locality Mexico: Jalisco, Guadalajara, not Costa Rica: Guápiles), not a , but a subspecies of W. Barnes & McDunnough, 1916, in Hesperiidae. A is designated for Herrich-Schäffer, 1869 (type locality becomes Mexico: Oaxaca). Plötz, 1883 is regarded as a , not a junior subjective synonym of (W. H. Edwards, 1865).