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: Rapis Grishin, subgen. n. (type species Papilio rapae Linnaeus, 1758, genus Pieris Schrank, 1801) in Pieridae Swainson, 1820 and Callitera Grishin, subgen. n. (type species Eurygona? pulcherrima Herrich-Schäffer, [1853], genus Isapis E. Doubleday, 1847), Matizada Grishin, subgen. n. (type species Themone poecila H. Bates, 1868, genus Isapis E. Doubleday, 1847), and Parapanara Grishin, subgen. n. (type species Lyropteryx diadocis Stichel, 1910, genus Paraphthonia Stichel, 1910) in Riodinidae Grote, 1895 (1827). New species are (type localities in parenthesis): Chlosyne pardelina Grishin, sp. n. (USA: Texas, Duval Co.) in Nymphalidae Rafinesque, 1815; Erythia paracheles Grishin, sp. n. (Panama: Canal Zone), Erythia borrosa Grishin, sp. n. (Panama: Panama), and Cremna telarania Grishin, sp. n. (Bolivia: La Paz) in Riodinidae; and Euriphellus colombiensis Grishin, sp. n. (Colombia: Río Dagua), Euriphellus ecuadoricus Grishin, sp. n. (Ecuador: Canelos), Urbanus (Urbanus) cubanus Grishin, sp. n. (Cuba: Havana), Gorgythion guyanus Grishin, sp. n. (Guyana: Essequibo), Lon co Grishin, sp. n. (Mexico: Guerrero), Lon ma Grishin, sp. n. (Panama: Chiriquí), and Lon chia Grishin, sp. n. (Mexico: Chiapas) in Hesperiidae Latreille, 1809. New subspecies are (type localities in parenthesis): Chlosyne definita dolosa Grishin, ssp. n. (Mexico: Chihuahua) in Nymphalidae and Limochores mystic nino Grishin, ssp. n. (USA: Arizona, Coconino Co.), Hesperia pahaska hannawackeri Grishin, ssp. n. (USA: Utah, San Juan Co.), Pseudocopaeodes eunus ash Grishin, ssp. n. (USA: Nevada, Nye Co.), Ochlodes napa kaibab Grishin, ssp. n. (USA: Arizona, Coconino Co.), and Lon melane sur Grishin, ssp. n. (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 Perpheres Hirowatari, 1992 as a valid genus, not a junior subjective synonym of Danis [Fabricius], 1807 (Lycaenidae); treat Sinopieris H. Huang, 1995 and Artogeia Verity, 1947 as subgenera (not genera or synonyms) of Pieris Schrank, 1801 (Pieridae); regard these genera as subgenera: Serradinga G. Henning & S Henning, 1996 of Dingana van Son, 1955 (Nymphalidae), Necyria Westwood, 1851 of Lyropteryx Westwood, 1851 (Riodinidae), and Nothodanis Hirowatari, 1992 of Danis [Fabricius], 1807 (Lycaenidae); and propose that the following are junior subjective synonyms, not genera, species, or subspecies: Eugrumia Della Bruna, Gallo, Lucarelli & Sbordoni, 2000 of Sinerebia Nakatani, 2017 in Nymphalidae: Satyrini: Ypthimina Reuter, 1896, new placement (not Erebiina Tutt, 1896), Pistoria Hemming, 1964 of Caleta Fruhstorfer, 1922 and Upolampes Bethune-Baker, 1908 of Thaumaina Bethune-Baker, 1908 in Lycaenidae, and Synargis orestessa Hübner, [1819] of Synargis soranus (Stoll, 1781) and Mesosemia eumene furia Stichel, 1910 of Ectosemia erinnya (Stichel, 1910) in Riodinidae. The following taxa are species, not subspecies or synonyms: Pontia edusa (Fabricius, 1777) (not Pontia daplidice (Linnaeus, 1758)) and Pontia johnstonii (Crowley, 1887) (not Pontia helice (Linnaeus, 1764)) in Pieridae; Chlosyne anastasia (Hemming, 1934) (not Chlosyne definita (E. Aaron, [1885])) and Chlosyne bollii (W. H. Edwards, 1878) (not Chlosyne theona (Ménétriés, 1855)) in Nymphalidae; Ectosemia attavus (J. Zikán, 1952) (not Ectosemia eumene (Cramer, 1776)), Cremna dentata (Stichel, 1910) (not Cremna radiata (Godman & Salvin, 1886)), Cremna pupillata Stichel, 1915 (not Cremna alector (Geyer, 1837)), Lasaia peninsularis Clench, 1972 (not Lasaia sula Staudinger, 1888), and Synargis arche (Hewitson, 1865) (not Synargis orestessa Hübner, [1819]) in Riodinidae; and Quadrus (Zera) difficilis (Weeks, 1901) (confirmed as not Quadrus (Zera) zera (A. Butler, 1870)), Gorgythion marginata Schaus, 1902 (not Gorgythion begga (Prittwitz, 1868)), Pardaleodes murcia (Plötz, 1883) (not Pardaleodes incerta (Snellen, 1872)), Pardaleodes pusiella Mabille, 1877 (not Pardaleodes sator (Westwood, 1852)), Ochlodes napa (W. H. Edwards, 1865) and Ochlodes santacruza J. Scott, 1981 (not Ochlodes sylvanoides (Boisduval, 1852)), and Lon vitellina (Herrich-Schäffer, 1869) and Lon poa (Evans, 1955) (not Lon melane (W. H. Edwards, 1869)) in Hesperiidae. In addition, we propose new genus-species combinations: Ectosemia nesti (Hewitson, 1858) (not Semomesia Westwood, 1851), Ectosemia acuta (Hewitson, 1873) and Ectosemia eurythmia (Stichel, 1915) (not Mesosemia Hübner, [1819]), Eugelasia satyroides (Lathy, 1926), Eugelasia modesta (H. Bates, 1868) and Pelolasia leucophryna (Schaus, 1913) (not Euselasia Hübner, 1819), Lyropteryx melaniae (Stichel, 1930) (not Melanis Hübner, [1819]), Isapis pulcherrima (Herrich-Schäffer, [1853]) and Isapis poecila (H. Bates, 1868) (not Themone Westwood, 1851), Paraphthonia diadocis (Stichel, 1910) (not Lyropteryx Westwood, 1851), Tarucus clathratus W. Holland, 1891 (not Castalius Hübner, [1819]), and Vidius tanna (de Jong, 1983) (not Cobalopsis Godman, 1900); new species-subspecies combination Ochlodes santacruza catalina J. Emmel & T. Emmel, 1998 (not Ochlodes sylvanoides (Boisduval, 1852)); and transfer junior subjective synonyms between taxa: Parnassius smintheus var. niger W. G. Wright, 1905 of Parnassius smintheus smintheus E. Doubleday, 1847 (not of Parnassius smintheus behrii W. H. Edwards, 1870), Goniurus proteoides Plötz, 1881 of Urbanus proteus domingo (Scudder, 1872) (not of Urbanus proteus proteus (Linnaeus, 1758)), Bolla subgisela Strand, 1921 of Staphylus melangon epicaste Mabille, 1903 (not of Bolla eusebius (Plötz, 1884)), Gorgythion beggoides Schaus, 1902 of Gorgythion begga begga (Prittwitz, 1868) (not of Gorgythion plautia (Möschler, 1877)), and Pamphila milo W. H. Edwards, 1883 of Ochlodes agricola verus (W. H. Edwards, 1881) (not of Ochlodes agricola nemorum (Boisduval, 1852)). Furthermore, by finding an additional specimen, we confirm Semalea malawi Grishin, 2023 as a species-level taxon using genomic analysis; we conclude that populations of Hesperia pahaska Leussler, 1938 in most of New Mexico and the White Mountains, Arizona are the nominal subspecies, not H. pahaska williamsi Lindsey, 1940; and report a natural hybrid between Chlosyne bollii (W. H. Edwards, [1878]) and Chlosyne chinatiensis (Tinkham, 1944). Lectotypes are designated for 6 names: Erebia atramentaria Bang-Haas, 1927 (type locality in China: Gansu) in Nymphalidae; Mesosemia eumene erinnya Stichel, 1910 (type locality in Peru: Pozuzo) and Mesosemia eumene furia Stichel, 1910 (type locality Bolivia: La Paz, Farinas) in Riodinidae; and Goniurus proteoides Plötz, 1881 (type locality in the Antilles), Hesperia erratica Plötz, 1883 (type locality likely in the USA, not Guatemala, as deduced by genomic comparison), confirming this name as a junior subjective synonym of Lon zabulon (Boisduval & Le Conte, [1837]), and Lerodea ? rupilius Schaus, 1913 (type locality Mexico: Jalisco, Guadalajara, not Costa Rica: Guápiles), not a nomen dubium, but a subspecies of Atrytonopsis edwardsi W. Barnes & McDunnough, 1916, in Hesperiidae. A neotype is designated for Cobalus vitellina Herrich-Schäffer, 1869 (type locality becomes Mexico: Oaxaca). Hesperia amanda Plötz, 1883 is regarded as a nomen dubium, not a junior subjective synonym of Ochlodes sylvanoides napa (W. H. Edwards, 1865).