The discovery that a skipper butterfly Telegonus fulgerator (Walch, 1775), previously placed in the genus Astraptes Hübner, [1819], is a complex of many similar-looking species-level taxa with different COI barcodes, caterpillar foodplants and body patterns, and subtle differences in adult phenotypes raised a question about which species is the original T. fulgerator. To answer this question, being unable to locate its holotype, we designate the neotype of Papilio fulgerator Walch, 1775, a female specimen from Suriname in the Zoological State Collection, Munich, Germany. This neotype will form the foundation for a comprehensive revision of the T. fulgerator complex based on genomic sequencing and analysis augmented with phenotypic considerations.
The comparative genomics of butterflies yields additional insights into their phylogeny and classification that are compiled here. As a result, 3 genera, 5 subgenera, 5 species, and 3 subspecies are proposed as new, i.e., in Hesperiidae: Antina Grishin, gen. n. (type species Antigonus minor O. Mielke, 1980), Pompe Grishin and Lamas, gen. n. (type species Lerema postpuncta Draudt, 1923), and Curva Grishin, gen. n. (type species Moeris hyagnis Godman, 1900); in Lycaenidae: Fussia Grishin, subgen. n. (type species Polyommatus standfussi Grum-Grshimailo, 1891) and Pava Grishin, subgen. n. (type species Thecla panava Westwood, 1852); in Hesperiidae: Monoca Grishin, subgen. n. (type species Tagiades monophthalma Plötz, 1884), Putuma Grishin, subgen. n. (type species Tisias putumayo Constantino and Salazar, 2013), and Rayia Grishin, subgen. n. (type species Mastor perigenes Godman, 1900); Cissia wahala Grishin, sp. n. (Nymphalidae; type locality in Mexico: Oaxaca); in Hesperiidae: Hedone mira Grishin and Lamas, sp. n. (type locality in Peru: Apurímac), Vidius pompeoides Grishin, sp. n. (type locality in Brazil: Amazonas), Parphorus hermieri Grishin, sp. n. (Hesperiidae; type locality in Brazil: Rondônia), and Zenis par Grishin, sp. n. (Hesperiidae; type locality in Peru: Cuzco); in Pieridae: Glutophrissa drusilla noroesta Grishin, ssp. n. (type locality in USA: Texas, Cameron Co.) and Pieris marginalis siblanca Grishin, ssp. n. (type locality in USA: New Mexico, Lincoln Co.), and Argynnis cybele neomexicana Grishin, ssp. n. (Nymphalidae; type locality in USA: New Mexico, Sandoval Co.). Acidalia leto valesinoides-alba Reuss, [1926] and Acidalia nokomis valesinoides-alba Reuss, [1926] are unavailable names. Neotypes are designated for Mylothris margarita Hübner, [1825] (type locality in Brazil) and Papilio coras Cramer, 1775 (type locality becomes USA: Pennsylvania, Montgomery Co., Flourtown). Mylothris margarita Hübner, [1825] becomes a junior objective synonym of Pieris ilaire Godart, 1819, currently a junior subjective synonym of Glutophrissa drusilla (Cramer, 1777). Lectotypes are designated for Hesperia ceramica Plötz, 1886 (type locality in Indonesia: Seram Island), Pamphila trebius Mabille, 1891 (type locality Colombia: Bogota), Methionopsis modestus Godman, 1901 and Papias microsema Godman, 1900 (type locality in Mexico: Tabasco), Hesperia fusca Grote & Robinson, 1867 (type locality in USA: Georgia), Goniloba corusca Herrich-Schäffer, 1869, and Goniloba devanes Herrich-Schäffer, 1869; the type localities of the last two species, together with Pamphila st
Two new skipper butterfly (Hesperiidae) species are described from the United States: Staphylus floridus Grishin, sp. n. (type locality in Florida, Volusia Co.) and Staphylus ecos Grishin, sp. n. (type locality in Texas, Brewster Co.). They are cryptic and hence escaped recognition. They differ from their sister species by the relative size and morphology of genitalia and by genotype-including and beyond the COI barcode-thus suggesting genetic isolation that argues for their species-level status. A lectotype is designated for Helias ascalaphus Staudinger, 1876. Staphylus opites (Godman & Salvin, 1896), stat. rest. is a species-level taxon and not a synonym of Staphylus vincula (Plötz, 1886), while Pholisora iguala Williams & Bell, 1940, syn. n. is a junior subjective synonym of S. vincula.
Two new species of Hermeuptychia Forster, 1964 are described. Hermeuptychia sinuosa Grishin, sp. n. (type locality Guatemala: El Progreso, Morazán) is an isolated member of the genus that does not readily fit into known species groups, as suggested by its distinct male and female genitalia and COI DNA barcode sequences. It is distinguished from its congeners by prominently wavy submarginal lines, rounder wings and distinctive genitalia, and can typically be identified by a white dot, instead of an eyespot, near the ventral hindwing apex. Hermeuptychia occidentalis Grishin, sp. n. (type locality Mexico: Guerrero, Acapulco) belongs to the Hermeuptychia sosybius group as indicated by the presence of androconia on the dorsal surface of the wings, genitalia and COI DNA barcodes, and in addition to DNA characters, differs from its relatives in the shape of the uncus and female genitalia. Neotypes of Oreas strigata canthe Hübner, [1811] (type locality Suriname: Gelderland, Suriname River), Megisto acmenis Hübner, 1823 (type locality Argentina: Buenos Aires), and Satyrus cantheus Godart, [1824] (type locality USA: Florida, Pinellas Co., St. Petersburg) and lectotype of Euptychia celmis var. bonaërensis [sic] Burmeister, 1878 (type locality Argentina: Buenos Aires) are designated. These designations establish Hermeuptychia canthe as a valid species widely distributed in South America from Colombia to Bolivia and Southeast Brazil, Euptychia celmis var. bonaërensis [sic] Burmeister, 1878 as a junior objective synonym of Yphthimoides acmenis, and S. cantheus as a junior subjective synonym of Hermeuptychia sosybius (Fabricius, 1793). Papilio camerta Cramer, 1780 is treated as nomen dubium requiring further studies to determine an identity that is consistent with the original description, as it may be conspecific with Paryphthimoides poltys (Prittwitz, 1865) instead of being a Hermeuptychia species as currently assumed.
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, trib. n. (type genus Calopieris Aurivillius, 1898); in subfamily Riodininae Grote, 1895: Callistiumini Grishin, trib. n. (type genus Callistium Stichel, 1911); in subfamily Nymphalinae Rafinesque, 1815: Pycinini Grishin, trib. n. (type genus Pycina Doubleday 1849), Rhinopalpini Grishin, trib. n. (type genus Rhinopalpa C. & R. Felder 1860), Kallimoidini Grishin, trib. n. (type genus Kallimoides Shirôzu & Nakanishi 1984), Vanessulini Grishin, trib. n. (type genus Vanessula Dewitz 1887), and Doleschalliaini Grishin, trib. n. (type genus Doleschallia C. & R. Felder 1860); in tribe Mesosemiini Bates, 1859: Eunogyrina Grishin, subtrib. n. (type genus Eunogyra Westwood, 1851); in tribe Satyrini Boisduval, 1833: Callerebiina Grishin, subtrib. n. (type genus Callerebia Butler, 1867), Gyrocheilina Grishin, subtrib. n. (type genus Gyrocheilus Butler, 1867), and Calistina Grishin, subtrib. n. (type genus Calisto Hübner, [1823]); in subfamily Euselasiinae Kirby, 1871: Pelolasia Grishin, gen. n. (type species Eurygona pelor Hewitson, [1853]), Myselasia Grishin, gen. n. (type species Eurygona mys Herrich-Schäffer, [1853]), Eurylasia Grishin, gen. n. (type species Eurygona euryone Hewitson, 1856), Maculasia Grishin, gen. n. (type species Euselasia albomaculiga Callaghan, 1999), and Eugelasia Grishin, gen. n. (type species Eurygona eugeon Hewitson, 1856); in subtribe Mesosemiina Bates, 1859: Ectosemia Grishin, gen. n. (type species Papilio eumene Cramer, 1776) and Endosemia Grishin, gen. n. (type species Papilio ulrica Cramer, 1777); in tribe Symmachiini Reuter, 1896: Tigria Grishin, gen. n. (type species Mesene xypete Hewitson, 1870) and Asymma Grishin, gen. n. (type species Symmachia virgatula Stichel, 1910); in tribe Riodinini Grote, 1895: Putridivora Grishin, gen. n. (type species Charis argyrea Bates, 1868), C
Further genomic sequencing of butterflies by our research group expanding the coverage of species and specimens from different localities, coupled with genome-scale phylogenetic analysis and complemented by phenotypic considerations, suggests a number of changes to the names of butterflies, mostly those recorded from the United States and Canada. Here, we present evidence to support these changes. The changes are intended to make butterfly classification more internally consistent at the genus, subgenus and species levels. I.e., considering all available evidence, we attempt to assign similar taxonomic ranks to the clades of comparable genetic differentiation, which on average is correlated with the age of phylogenetic groups estimated from trees. For species, we use criteria devised by genomic analysis of the genetic differentiation across suture zones and comparison of sympatric populations of closely related species. As a result, we resurrect 4 genera and 1 subgenus from subgeneric status or synonymy, change the rank of 8 currently used genera to subgenus, synonymize 7 genus-group names, summarize evidence to support 19 taxa as species instead of subspecies and 1 taxon as subspecies instead of species, along with a number of additional changes. One new genus and one new subspecies are described. Namely, the following taxa are treated as genera Tharsalea Scudder, 1876, Helleia Verity, 1943, Apangea Zhdanko, 1995, and Boldenaria Zhdanko, 1995. Tetracharis Grote, 1898 is a valid subgenus (not a synonym of Anthocharis Boisduval, Rambur, [Duménil] & Graslin, [1833]) that consists of Anthocharis cethura C. Felder & R. Felder, 1865 (Müller, 1764), Anthocharis midea (Hübner, [1809]), and Anthocharis limonea (A. Butler, 1871). The following are subgenera: Speyeria Scudder, 1872 of Argynnis Fabricius, 1807; Aglais Dalman, 1816 and Polygonia Hübner, [1819] of Nymphalis Kluk, 1780; Palaeonympha Butler, 1871 of Megisto Hübner, [1819]; Hyponephele Muschamp, 1915 of Cercyonis Scudder, 1875; Pyronia Hübner, [1819] and Aphantopus Wallengren, 1853 of Maniola Schrank, 1801 and Pseudonymphidia Callaghan, 1985 of Pachythone. Lafron Grishin, gen. n. (type species Papilio orus Stoll, [1780], parent subfamily Lycaeninae [Leach], [1815]) is described. Dipsas japonica Murray, 1875 is fixed as the type species of Neozephyrus Sibatani & Ito, 1942. The following taxa are junior subjective synonyms: Falcapica Klots, 1930 of Tetracharis Grote, 1898; Habrodais Scudder, 1876, Favonius Sibatani & Ito, 1942, Neozephyrus Sibatani & Ito, 1942, Quercusia Verity, 1943, Chrysozephyrus Shirôzu & Yamamoto, 1956, and Sibataniozephyrus Inomata, 1986 of Hypaurotis Scudder, 1876; Plesioarida Trujano & García, 2018 of R
We obtained and analyzed whole genome shotgun sequences of all 845 species of butterflies recorded from Canada and the United States. Genome-scale phylogenetic trees constructed from the data reveal several non-monophyletic genera and suggest improved classification of species included in these genera. Here, these changes are formalized and 2 subgenera are described: Amblyteria Grishin, subgen. n. (type species Goniloba exoteria Herrich-Schäffer, 1869, parent genus Amblyscirtes Scudder, 1872), and Coa Grishin, subgen. n. (type species Hesperia baracoa Lucas, 1857, parent genus Polites Scudder, 1872). Furthermore, we resurrect 3 genera and 2 subgenera from synonymy, change the rank of 6 currently used genera to subgenus, synonymize 2 genera, transfer 3 (2 resurrected) subgenera and 11 additional species to different genera than those these taxa were assigned to, and raise one name from synonym to species rank. Namely, Hedone Scudder, 1872 and Limochores Scudder, 1872 are valid genera and not synonyms of Polites Scudder, 1872; Pendantus K. Johnson & Kroenlein, 1993 is a valid genus and not a synonym of Electrostrymon Clench, 1961; and Sphaenogona Butler, 1870 and Lucidia Lacordaire, 1833 are valid subgenera of Abaeis Hübner, [1819] (new placement) and not synonyms of Eurema Hübner, [1819]. The following taxa are best treated as subgenera: Mimoides Brown, 1991 of Eurytides Hübner, [1821] (sensu lato); Philotiella Mattoni, [1978] of Euphilotes Mattoni, [1978]; Neominois Scudder, 1875 of Oeneis Hübner, [1819]; Agraulis Boisduval & Le Conte, [1835] of Dione Hübner, [1819]; Copaeodes Speyer, 1877 of Oarisma Scudder, 1872; and Problema Skinner & R. Williams, 1924 of Atrytone Scudder, 1872. Phaeostrymon Clench, 1961 and Saliana Evans, 1955 are junior subjective synonyms of Satyrium Scudder, 1876 and Calpodes Hübner, [1819], respectively. The entire subgenus Erynnides Burns, 1964 is transferred from Erynnis Schrank, 1801 to Gesta Evans, 1953. New genus-species combinations resulting from transfer of species between genera are: Nastra perigenes (Godman, 1900) (not Vidius Evans, 1955); Troyus fantasos (Cramer, 1780), Troyus onaca (Evans, 1955), Troyus aurelius (Plötz, 1882), Troyus marcus (Fabricius, 1787), Troyus diversa (Herrich-Schäffer, 1869), and Troyus drova (Evans, 1955) (not Vettius Godman, 1901); Oligoria percosius (Godman, 1900), Oligoria rindgei (H. Freeman, 1969), Oligoria lucifer (Hübner, [1831]), and Oligoria mustea (H. Freeman, 1979) (not Decinea Evans, 1955). Urbanus alva Evans, 1952 is a valid species and not a synonym of Urbanus belli (Hayward, 1935), new status.
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