Genomics-based taxonomic rearrangement of Achlyodini and Carcharodini (Lepidoptera: Hesperiidae: Pyrginae)

Abstract

Genomic analysis of Pyrginae Burmeister, 1878 (Lepidoptera: Hesperiidae Latreille, 1809) with an emphasis on the tribes Achlyodini Burmeister, 1878 and Carcharodini Verity, 1940 reveals many inconsistencies between the resulting phylogeny and the current classification. These problems are corrected by proposing new taxa, changing the ranks of others, or synonymizing them, and transferring species between genera. As a result, five subtribes, one genus, 20 subgenera, and one species are proposed as new: Cyclosemiina Grishin, new subtribe (type genus Cyclosemia Mabille, 1878), Ilianina Grishin, new subtribe (type genus Iliana E. Bell, 1937), Nisoniadina Grishin, new subtribe (type genus Nisoniades Hübner, [1819]), Burcina Grishin, new subtribe (type genus Burca E. Bell and W. Comstock, 1948), and Pholisorina Grishin, new subtribe (type genus Pholisora Scudder, 1872), all in Carcharodini; Lirra Grishin, new genus (type species Leucochitonea limaea Hewitson, 1868) in Pythonidina Grishin, 2019; Trifa Grishin, new subgenus (type species Tagiades jacobus Plötz, 1884), Tuberna Grishin, new subgenus (type species Pythonides contubernalis Mabille, 1883), Ebona Grishin, new subgenus (type species Quadrus eboneus E. Bell, 1947), Noctis Grishin, new subgenus (type species Achlyodes accedens Mabille, 1895), and Cyrna Grishin, new subgenus (type species Achlyodes cyrna Mabille, 1895) of Quadrus Lindsey, 1925; Liddia Grishin, new subgenus (type species Helias pallida R. Felder, 1869), Minna Grishin, new subgenus (type species Achlyodes minna Evans,1953), and Thilla Grishin, new subgenus (type species Eurypterus later Mabille, 1891) of Eantis Boisduval, 1836; Torgus Grishin, new subgenus (type species Ouleus gorgus E. Bell, 1937) of Iliana E. Bell, 1937; Fenops Grishin, new subgenus (type species Cabares enops Godman and Salvin, 1894) of Polyctor Evans, 1953; Bezus Grishin, new subgenus (type species Pellicia bessus Möschler, 1877) and Macarius Grishin, new subgenus (type species Pellicia macarius Herrich-Schäffer, 1870) of Nisoniades Hübner, [1819]; Quadralis Grishin, new subgenus (type species Pterygospidea extensa Mabille, 1891) of Gorgopas Godman and Salvin, 1894; Menuda Grishin, new subgenus (type species Nisoniades menuda Weeks, 1902) and Narycus Grishin, new subgenus (type species Pythonides narycus Mabille, 1889) of Perus Grishin, 2019; Bovaria Grishin, new subgenus (type species Achlyodes cyclops Mabille, 1876), Sebia Grishin, new subgenus (type species Nisoniades eusebius Plötz, 1884), and Stolla Grishin, new subgenus (type species Pholisora balsa E. Bell, 1937) of Bolla Mabille, 1903; Vulga Grishin, new subgenus (type species Achlyodes vulgata Möschler, 1879) and Capilla Grishin, new subgenus (type species Helias aurocapilla Staudinger, 1876, currently a junior subjective synonym of Hesperia musculus Burmeister, 1875) of Staphylus Godman and Salvin, 1896; and Quadrus (Zera) vivax Grishin, new species (type locality in Brazil: Rio de Janeiro). The following 10 are subgenera, not genera or synonyms: Ouleus Lindsey, 1925 and Zera Evans, 1953 of Quadrus Lindsey, 1925; Atarnes Godman and Salvin, 1897 and Eburuncus Grishin, 2012 of Milanion Godman and Salvin, 1895; Pachyneuria Mabille, 1888 and Austinus O. Mielke and Casagrande, 2016 of Sophista Plötz, 1879; Hemipteris Mabille, 1889 and Mictris Evans, 1955 of Pellicia Herrich-Schäffer, 1870; and Hesperopsis Dyar, 1905 and Scantilla Godman and Salvin, 1896 of Staphylus Godman and Salvin, 1896. The following 7 are species, not subspecies: Quadrus (Ebona) cristatus (Steinhauser, 1989) (not Quadrus (Ebona) negrus (Nicolay, 1980)), Quadrus (Quadrus) ophia (A. Butler, 1870) (not Quadrus (Quadrus) lugubris (R. Felder, 1869)), Quadrus (Zera) gellius (Mabille,1903) and Quadrus (Zera) servius (Plötz, 1884) (not Quadrus (Zera) hyacinthinus (Mabille, 1877)), Mimia pazana Evans,1953 (not Mimia phidyle (Godman and Salvin, 1894)), Polyctor (Polyctor) dagua Evans, 1953 (not Polyctor (Polyctor) polyctor (Prittwitz, 1868)), and Staphylus (Vulga) satrap Evans, 1953 (not Staphylus (Vulga) saxos Evans, 1953); and these 8 are species, not synonyms: Quadrus (Zera) menedemus (Godman and Salvin, 1894) (not Quadrus (Zera) tetrastigma (Sepp, [1847])), Pellicia (Pellicia) bilinea Mabille, 1889 (not Pellicia (Pellicia) dimidiata Herrich-Schäffer, 1870), Pellicia (Hemipteris) nema Williams and Bell, 1939 (not Pellicia (Pellicia) theon Plötz, 1882), Bolla (Bovaria) sodalis Schaus, 1913 (not Bolla (Bolla) imbras (Godman and Salvin, 1896)), Bolla (Bovaria) aplica (E. Bell, 1937) (not Bolla (Sebia) eusebius (Plötz, 1884)), Bolla (Sebia) chilpancingo (E. Bell, 1937) (not Bolla (Bolla) subapicatus (Schaus, 1902)), and Bolla (Stolla) madrea (R. Williams and E. Bell, 1940) and Bolla (Stolla) hazelae (Hayward, 1940) (not Bolla (Stolla) zorilla (Plötz,1886)). The following 2 are junior subjective synonyms: Achlyodes erisichthon Plötz, 1884 of Quadrus (Zera) servius (Plötz, 1884) (not a subspecies of Quadrus (Zera) tetrastigma (Sepp, [1847]) and Staphylus subapicatus Schaus, 1902 of Bolla (Bolla) imbras (Godman and Salvin, 1896). Furthermore, we propose the following additional new genus-species combination: Gindanes homer (Evans, 1953), Gindanes nides (O. Mielke and Casagrande, 2002), Gindanes maraca (O. Mielke and Casagrande, 1992), Gindanes jenmorrisae (Shuey and Ramírez. 2022), Gindanes tullia (Evans, 1953), Gindanes herennius (Geyer, [1838]), Gindanes proxenus (Godman and Salvin, 1895), Gindanes parallelus (Mabille, 1898), Gindanes braga (Evans, 1953), Gindanes hampa (Evans, 1953), Gindanes rosa (Steinhauser, 1989), Gindanes neivai (Hayward, 1940), Gindanes mundo (H.Freeman, 1979), Gindanes eminus (E. Bell, 1934), Quadrus (Trifa) francesius Freeman, 1969, Quadrus (Trifa) ineptus (Draudt, 1922), Quadrus (Trifa) jacobus (Plötz, 1884), Quadrus (Tuberna) lancea (Hewitson, 1868), Quadrus (Ebona) pescada (E. Bell, 1956), Lirra pteras (Godman and Salvin, 1895), and Lirra limaea (Hewitson, 1868) (not Pythonides Hübner, 1819); Quadrus (Cyrna) zora (Evans, 1953) (not Bolla Mabille, 1903); Eantis later (Mabille, 1891) and Eantis haber (Mabille, 1891) (not Aethilla Hewitson, 1868); Iliana (Torgus) gorgus (E. Bell, 1937) and Iliana (Torgus) taurus (Evans, 1953) (not Eantis Boisduval, 1836); Bolla (Stolla) evemerus (Godman and Salvin, 1896), Bolla (Stolla) chlora (Evans, 1953), Bolla (Stolla) astra (R. Williams and E. Bell, 1940), Bolla (Stolla) balsa (E. Bell, 1937), Bolla (Stolla) tridentis (Steinhauser, 1989), Bolla (Stolla) esmeraldus (L. Miller, 1966), Bolla (Stolla) chlorocephala (Latreille, [1824]), and Bolla (Stolla) incanus (E.Bell, 1932) (not Staphylus Godman and Salvin, 1896). Finally, lectotypes are designated for Achlyodes servius Plötz, 1884 (type locality in Brazil: Rio de Janeiro), Pellicia theon Plötz, 1882 (type locality in South America), and Nisoniades eusebius Plötz, 1884 (type locality in Central America). Unless stated otherwise, all subgenera, species, subspecies, and synonyms of mentioned genera and species are transferred with their parent taxa, and others remain as previously classified.