Taxonomic changes suggested by the genomic analysis of Hesperiidae (Lepidoptera).

Our expanded efforts in genomic sequencing to cover additional skipper butterfly (Lepidoptera: Hesperiidae) species and populations, including primary type specimens, call for taxonomic changes to restore monophyly and correct misidentifications by moving taxa between genera and proposing new names. Reconciliation between phenotypic characters and genomic trees suggests three new tribes, two new subtribes, 23 new genera, 17 new subgenera and 10 new species that are proposed here: Psolosini Grishin, (type genus Staudinger, 1889), Ismini Grishin, (type genus Distant, 1886), Eetionini Grishin, (type genus de Nicéville, 1895), Orphina Grishin, (type genus Godman, 1901), Carystoidina Grishin, (type genus Godman, 1901), Grishin, (type species Plötz, 1882), Grishin, (type species Mabille and Boullet, 1912), Grishin, (type species Grishin, ), Grishin, (type species Plötz, 1884), Grishin, (type species Evans, 1937), Grishin, (type species Trimen, 1873), Grishin, (type species Hayward, 1951), Grishin, (type species Mabille, 1889), Grishin, (type species ? Schaus, 1913), Grishin, (type species Schaus, 1902), Grishin, (type species Schaus, 1902), Grishin, (type species Godman, 1900), Grishin, (type species Bell, 1937), Grishin, (type species Bell, 1932), Grishin, (type species Plötz, 1882), Grishin, (type species Bell, 1930), Grishin, (type species Bell, 1930), Grishin, (type species Bell, 1932), Grishin, (type species Lindsey, 1925), Grishin, (type species Hewitson, 1877), Grishin, (type species Godman, 1900), Grishin, (type species Hewitson, 1866), Grishin, (type species Hewitson, 1878), Grishin, (type species Evans, 1952), Grishin, (type species Cramer, 1780), Grishin, (type species Plötz, 1884), Grishin, (type species Plötz, 1884), Grishin, (type species Evans, 1928), Grishin, (type species Herrich-Schäffer, 1869), Grishin, (type species Mabille, 1891), Grishin, (type species [sic] Schaus, 1913), Grishin, (type species Godman, 1900), Grishin, (type species Möschler, 1879), Grishin, (type species Mielke and Casagrande, 2002), Grishin, (type species Godman, 1901), Grishin, (type species Schaus, 1902), Grishin, (type species Evans, 1955), Grishin, (type species Evans, 1955), Grishin, (type species Godman, 1900), Grishin, (type species Evans, 1955), Grishin, (type locality in Brazil: Santa Catarina), Grishin, (type locality in Guyana: Acarai Mts.), Grishin, (type locality in Paraguay: Sapucay), Grishin, (type locality in Brazil: Rondônia), Grishin, (type locality in Ecuador: Riobamba), Grishin, (type locality in Colombia: Bogota), Grishin, (type locality in Panama: Colón), Grishin, (type locality in Nicaragua: Chontales), Grishin, (type locality in Peru: Cuzco), and Grishin, (type locality in Costa Rica). are designated for the following 17 taxa: Mabille, 1888, Plötz, 1882, Mabille, 1888, Mabille, 1888, Herrich-Schäffer, 1869, Mabille, 1891, Plötz, 1884, Mabille, 1891, Plötz, 1882, Herrich-Schäffer, 1869, Mabille, 1891, Godman, 1900, Schaus, 1913, Mabille, 1891, Godman, 1900, Schaus, 1902, and Plötz, 1882. The lectotype of Plötz, 1882 is because it does not agree with the original description and illustration by Plötz, is not from the locality listed in the original description, and therefore is not a syntype. are designated for the following five taxa: Plötz, 1882, Plötz, 1882, Plötz, 1882, Plötz, 1882, and Plötz, 1882. As a result, the following five taxa are : Möschler, 1883 of Plötz, 1882, Mabille, 1891 of Plötz, 1882, Mabille, 1891 of Plötz, 1882, Schaus, 1902 of Plötz, 1882, and Plötz, 1882 of Hewitson, 1877. Evans, 1955 is fixed as the of Godman, 1900, and Butler, 1877 is reaffirmed as the of Godman, 1900. Furthermore, the following are suggested. Holland, 1896, Evans, 1937, and de Nicéville, 1895 are placed in Aeromachini Tutt, 1906. Larsen and Congdon, 2011 is transferred from Celaenorrhinini Swinhoe, 1912 to Tagiadini Mabille, 1878. Larsen and Collins, 2013 is transferred from Tagiadini Mabille, 1878 to Celaenorrhinini Swinhoe, 1912. The following nine genus-group names are and treated as valid genera: Moore, 1881 (not in Wallengren, 1857), Mabille, 1904 (not in Butler, 1870), Distant, 1886 and Swinhoe, 1913 (not in Butler, 1874), Godman, 1900 (not in Godman, 1900), Hayward, 1942 (not in Godman, 1900), Godman, 1900 (not in Mabille, 1889), Hayward, 1939 (not in Hübner, [1819]), and Godman, 1900 (not in Godman, 1900). The genus-group names Murray, 1875 and Wallengren, 1857 are and treated as valid subgenera of Hübner, [1819]. We confirm Strand, 1911 as a valid genus. The following 24 genera are placed as subgenera, : Austin, 2008 of Watson, 1893; Huang, Chiba, Wang and Fan, 2016 of Evans, 1932; de Nicéville, 1890 of Moore, [1881]; de Nicéville, 1890 of Moore, 1881; Wallengren, 1857 of Hopffer, 1855; Godman and Salvin, 1896 of Godman and Salvin, 1896; Hou, Fan and Chiba, 2021 of Duméril, 1806; de Nicéville, 1898 of Watson, 1893; Moore, [1881] and de Nicéville, 1889 of Butler, 1874; de Jong, 1983 of Godman, 1900; Hayward, 1939 of Scudder, 1872; O. Mielke and Casagrande, 2003 of Mabille, 1904; Evans, 1955 of Evans, 1955; Schaus, 1913 and Hemming, 1939 of Godman, 1901; Evans, 1955 of Godman, 1901; Godman, 1900 of Scudder, 1872; Mabille, 1889 of Hübner, [1831]; Evans, 1955 of Hayward, 1939; Godman, 1900 of Godman, 1901; and Evans, 1955, Evans, 1955, and Mabille, 1904 of Hübner, [1819]. The following 20 genera are treated as junior subjective synonyms: Wallengren, 1857 of Hopffer, 1855; Plötz, 1879 and Mabille, 1903 of Wallengren, 1857; Devyatkin, 1996 of Watson, 1893; Eliot, 1978 of Elwes and Edwards, 1897; Evans, 1937 of Butler, 1870; Mabille, 1904 of Godman, 1901; Evans, 1955, Evans, 1955, and Evans, 1955 of Godman, 1900; O. Mielke and Casagrande, 2002 of Hayward, 1939; Evans, 1955 of Mabille, 1904; Grishin, 2019 of Schaus, 1913; Godman, 1900 of Godman, 1900; Evans, 1955 of Godman, 1900; Mabille, 1904 and Mabille, 1904 of Scudder, 1872; Mabille, 1903 of Godman, 1901; Evans, 1955 of Hübner, [1819]; and Godman, 1901 of Scudder, 1872. Swinhoe, 1919 is a of de Nicéville, 1898 (not of Watson, 1893). The following 27 names are species-level taxa (some in new combinations) : (Mabille, 1888) (not (Cramer, 1777)), (Plötz, 1882) (not (Cramer, 1775)), (Herrich-Schäffer, 1869) (not (Fabricius, 1793)), (Mabille, 1888) (not (Stoll, 1780)), Mabille and Boullet, 1917 (not (Möschler, 1879)), (Mabille, 1889) (not (Cramer, 1775)), (Mabille, 1891) and (Möschler, 1877) (not (Hewitson, 1877)), (Dyar, 1916) (not (Plötz, 1882)), (Mabille, 1889) (not (Mabille, 1889)), (Freeman, 1969) (not (Schaus, 1902)), (Schaus, 1902) (not Godman, 1900), (Mabille, 1891) (not (Mabille, 1891), previously in Godman, 1900), Scudder, 1872 (not (J. E. Smith, 1797)), (Möschler, 1879) (not (Capronnier, 1874)), (Bell, 1942) (not (Herrich-Schäffer, 1869)), (Schaus, 1902) (not (Möschler, 1879), previously in Godman, 1900), (Godman, 1900) (not (Mabille, 1891)), (Godman, 1900) (not (Scudder, 1872)), (Mabille, 1891) (not (Butler, 1877), previously in Godman, 1900), (Mabille, 1891) (not (Herrich-Schäffer, 1870)), Godman, 1900 (not Plötz, 1882), (Mabille, 1891) (not (Hewitson, 1877), previously in Mabille, 1889), (Hayward, 1942) (not (Godman, 1900), previously in Godman, 1900), (Draudt, 1923) (not (Capronnier, 1874)), Godman, 1901 (not (Plötz, 1882)), and (Godman, 1901) (not (Plötz, 1884)). The following 57 taxa are elevated from subspecies to species, (some in ): (Hewitson, 1867) (not (C. Felder and R. Felder, 1862), (A. Butler, 1872) (not (Westwood, 1852)), Evans, 1932 (not Möschler, 1878), Fruhstorfer, 1910 (not Elwes and Edwards, 1897), Evans, 1934, Evans, 1934, Plötz, 1884, Mabille, 1883, and Plötz, 1885 (not (Hopffer, 1874)), Mabille, 1891 and Butler, 1882 (not (C. Felder, 1860)), Mabille, 1876, (Moore, [1866]), (Fabricius, 1793), Plötz, 1884, Butler, 1870, Rothschild, 1915, and Swinhoe, 1904 (not (Stoll, [1781])), Evans, 1934 and Mabille, 1877 (not (Moore, [1866])), Evans, 1941 and Elwes and Edwards, 1897 (not Moore, [1881]), (Distant, 1886) (not (Hewitson, 1868)), (Evans, 1953) (not (Schaus, 1902)), (Röber, 1925) (not (R. Felder, 1869)), Evans, 1953 (not (Mabille, 1891)), Hayward, 1933 and Mabille, 1876 (not (Butler, 1870)), Evans, 1953 (not Mabille, 1898), (Evans, 1953) (not (Mabille, 1888)), (Möschler, 1879) (not (Herrich-Schäffer, 1865)), Evans, 1949 (not (Plötz, 1882)), (Reakirt, 1866) (not (Boisduval, 1852)), Bell, 1930 (not (Mabille, 1889)), (Draudt, 1923) (not (Giacomelli, 1911), previously in Godman, 1900), (H. Freeman, 1969), Evans, 1955, and (Mabille, 1895) (not (Hewitson, 1876)), Godman, 1900 (not (Hewitson, 1866)), Draudt, 1923 (not (Cramer, 1777)), (Hewitson, 1870) (not (Herrich-Schäffer, 1869)), (Evans, 1955) (not (Schaus, 1913), previously in Evans, 1955), Evans, 1955, (not (Schaus, 1913)), (Evans, 1955) (not (Schaus, 1902), previously in Hayward, 1938), (Bell, 1941) (not (Mabille, 1891)), (E. Bell, 1940) (not (Schaus, 1902) previously in Godman, 1900), (Plötz, 1882), (Dyar, 1913) and (Hayward, 1940) (not (Burmeister, 1878)), (Evans, 1955) (not (Hewitson, 1868), previously in Mabille, 1889), Evans, 1955 (not (Mabille, 1883)), (Mabille, 1891), (Godman, 1900), and (Weeks, 1901) (not (Capronnier, 1874)), (E. Bell, 1941) and (Hayward, 1942) (not (Mabille, 1898)), and (Skinner, 1920) (not (Herrich-Schäffer, 1863), previously in Mabille, 1904). (Elwes and J. Edwards, 1897) is a valid subspecies, not a synonym of (Mabille, 1891), here placed in synonymy with (Hewitson, 1876), . We confirm the species status of (Evans, 1952) and Evans, 1955. Freeman, 1969 is placed as a subspecies of (Reakirt, 1866), . The following 41 taxa are , either newly proposed or transferred from synonymy with other species or subspecies: Plötz, 1882 of (Hewitson, 1876), Mabille, 1888 of (Plötz, 1882), Butler, 1870 of (Herrich-Schäffer, 1869), Evans, 1953 of (Mabille, 1888), Plötz, 1884 of (Plötz, 1884) (not of (Möschler, 1877)), Strand, 1921 of (not of ), Plötz, 1884 of (Mabille, 1878) (not of (Latreille, [1824])), Mabille, 1889 of (Mabille, 1889), Schaus, 1902 of (Mabille, 1891), Weeks, 1906 of (Möschler, 1877), Evans, 1955 of (Mabille, 1891), Mabille, 1891 of (A. Butler, 1877) (not of (Hewitson, 1876)), Nicolay, 1980 of (Hayward, 1940), Plötz, 1882 of (Herrich-Schäffer, 1869) (not of (Plötz, 1882)), Möschler, 1883 of (Herrich-Schäffer, 1869) (not of (Herrich-Schäffer, 1869)), E. Bell, 1959 of (Schaus, 1902), Anderson and Nakamura, 2019 of (Bell, 1937), Herrich-Schäffer, 1869 of (Grote and Robinson, 1867) (not of (W. H. Edwards, 1862)), Nicolay, 1973 of (Mabille, 1891), Mabille, 1889 and Mabille, 1904 of Scudder, 1872 (not of (Herrich-Schäffer, 1869), previously in Scudder, 1872), Plötz, 1886 of (Herrich-Schäffer, 1865) (not of Butler, 1877), Schaus, 1902 of (Möschler, 1879), Kaye, 1914 of (Schaus, 1902) (not of (Möschler, 1879), previously in Godman, 1900), Burmeister, 1878, Mabille, 1891 and Schaus, 1902 of (Herrich-Schäffer, 1869), Plötz, 1882 of (Plötz, 1882) (not of (J. E. Smith, 1797)), Mabille, 1898 of (Scudder, 1872), Lindsey, 1925 of (Mabille, 1891), Mabille, 1891 of (Mabille, 1891) (not of (Plötz, 1882)), H. Freeman, 1969 of Godman, 1900 (previously in Mabille, 1889), Plötz, 1882 of (Herrich-Schäffer, 1870) (previously Mabille, 1904), (not of (Herrich-Schäffer, 1870)), Mabille, 1891 of (Möschler, 1877) (not of (Capronnier, 1874)), H. Freeman, 1969 of (Dyar, 1918), Godman, 1901 of (Plötz, 1882), Evans, 1955 of (Godman, 1901), Plötz, 1882 of (Stoll, 1781), Herrich-Schäffer, 1869 of (Stoll, 1781) (not of (Latreille, [1824])), and Evans, 1955 of (Evans, 1955). The following are : (Cramer, 1777) (not Evans, 1952), (Mabille, 1891) (not Evans, 1953), (Evans, 1953) (not Godman and Salvin, 1896), (Mabille, 1889) (not Lindsey, 1925), (Steinhauser and Austin, 1993) (not Godman and Salvin, 1896), (Dyar, 1912) (not Mabille, 1903), (A. Butler, 1874) (not Hübner, [1819]), (Mabille, 1888) (not Grishin, 2019), (Nicolay, 1980) (not Mabille, 1876), (de Nicéville, 1889) (not Distant, 1886), (Evans, 1937) (not Mabille, 1891), (Hewitson, 1876) (not Holland, 1896), (Evans, 1951) (not C. Felder and R. Felder, 1860), (Trimen, 1883) (not Watson, 1893), (Hewitson, 1876) (not Evans, 1955), (Evans, 1955) (not Evans, 1955), (Weeks, 1901) (not Godman, 1900), (Hayward, 1951) (not Godman, 1900), (Giacomelli, 1911) (not Godman, 1900), (Mabille, 1891) (not Evans, 1955), (Dyar, 1914) (not Scudder, 1872), (Hewitson, 1876) (not Evans, 1955), (Evans, 1955) (not Evans, 1955), (Evans, 1955) and (Erschoff, 1876) (not Godman, 1901), (Schaus, 1902) and (E. Bell, 1942) (not Godman, 1900), (Evans, 1955) (not Mabille, 1904), (Mabille, 1891) and (Hayward, 1940) (not Godman, 1900), (Hayward, 1940) (not Godman, 1900), (Hewitson, 1867) and (Schaus, 1902) (not Hayward, 1938), Plötz, 1882 (not Scudder, 1872), (Evans, 1955) (not Godman, 1900), (Hayward, 1951), (Evans, 1955), (Evans, 1955), (Plötz, 1882), and (Godman, 1900) (not Godman, 1900), (E. Bell, 1941) and (Mabille, 1891) (not Godman, 1900), (Schaus, 1902) (not Godman, 1900), (Herrich-Schäffer, 1869) (not Evans, 1955), (Plötz, 1882) (not Scudder, 1872), (Godman, 1900) (not Evans, 1955), (Godman, 1900) (not Scudder, 1872), (Mabille, 1891) (not Godman, 1900), (Evans, 1955) (not Evans, 1955), (Weeks, 1901) and (Lindsey, 1925) (not Scudder, 1872), (Herrich-Schäffer, 1869) (not Scudder, 1872), (Möschler, 1879) (not Godman, 1900), (Godman, 1900) (not Godman, 1900), (Bell, 1942) (not Godman, 1900), (Schaus, 1913) (not Godman, 1900), (Evans, 1955) and (Hewitson, 1877) (not Mabille, 1889), (Steinhauser, 2008) (not Godman, 1900), (Schaus, 1902) (not Godman, 1900), (E. Bell, 1930) (not Godman, 1901), (Evans, 1955) (not Godman, 1900), (Hayward, 1939) (not Mabille, 1904), (Capronnier, 1874) (not Godman, 1901), Hayward, 1940 (not Evans, 1955), (Evans, 1955) (not Evans, 1955), (Plötz, 1884) (not Evans, 1955), (Hayward, 1942) (not Godman, 1901), (Nicolay, 1973) (not Evans, 1955), (Evans, 1955) (not Hübner, [1831]), Plötz, 1882 (not Hübner, [1819]), (Evans, 1955) (not Hayward, 1938), and (Evans, 1955) (not Hübner, [1821]). The following are : Evans, 1934, Fruhstorfer, 1910, Evans, 1934, Evans, 1945, Evans, 1934, Evans, 1934, Rothschild, 1915, Evans, 1949, Fruhstorfer, 1910, Evans, 1934, Evans, 1934, C. Felder, 1862, Fruhstorfer, 1910, Evans, 1934, Evans, 1934, de Jong and Treadaway, 2007, Fruhstorfer, 1910, Evans, 1955, Evans, 1955, and (Bates, 1934) (previously in Mabille, 1904). In addition to the abovementioned changes, the following involve newly proposed genus group names: (Plötz, 1882) and (E. Bell, 1934) (not Evans, 1952); (Mabille and Boullet, 1912) (not Watson, 1893); Evans, 1952 (Cramer, 1780), and (Plötz, 1882) (not Austin, 2008); (Plötz, 1884); (Plötz, 1884), and (Hopffer, 1874) (not Hübner, [1819]); (A. Butler, 1877); (Plötz, 1884) and (Godman & Salvin, 1894) (not Godman and Salvin, 1894); (Evans, 1928), (Oberthür, 1878), (Plötz, 1882) (Swinhoe, 1907) (Janson, 1886), and (Evans, 1949) (not de Nicéville, 1889); (Evans, 1937) (Mabille, 1891) (Hewitson, 1876), and (Evans, 1937) (not Evans, 1937); (Trimen, 1873) and (Henning and Henning, 1998) (not Watson, 1893); (Hayward, 1951) (not Godman, 1900); (Herrich-Schäffer, 1869) (Evans, 1955) (Constantino and Salazar, 2013) (not Godman, 1901); (Mabille, 1889) (not Schaus, 1913); (Schaus, 1913) (not Evans, 1955); (Mabille, 1891) (Plötz, 1882) Evans, 1955 (E. Bell, 1941) Evans, 1955 (Geyer, 1832), and (Möschler, 1879); (Schaus, 1902) (not Godman, 1900); (Evans, 1955) (not Godman, 1900); (E. Bell, 1932) (not Godman, 1900); (Schaus, 1902) (not Godman, 1900); (Hayward, 1940) (not Godman, 1900); (Godman, 1900) (not Godman, 1900); (E. Bell, 1937) (not Evans, 1955); (Schaus, 1913) (not Godman, 1900); (Godman, 1900) (not Evans, 1955); (E. Bell, 1932) (not Godman, 1900); (Schaus, 1902) (not Godman, 1900); (Evans, 1955) (not not Evans, 1955); (Möschler, 1879) and (Godman, 1900) (not Godman, 1900); (Plötz, 1882) (not Scudder, 1872); (E. Bell, 1930) (not Schaus, 1913); (Hayward, 1940), (not Godman, 1900); (Mielke and Casagrande, 2002), (not Mabille, 1889); (E. Bell, 1930) (not Evans, 1955); (E. Bell, 1932) (not Godman, 1901); (Lindsey, 1925) (not Godman, 1900); (Schaus, 1902) (not Godman, 1900); (Hewitson, 1877) (not Evans, 1955); Evans, 1955; (Godman, 1900) (not Evans, 1955); (Evans, 1955) (not Hübner, [1831]); (Hewitson, 1866) (not Hübner, [1819]); (Godman, 1900) (not Evans, 1955); (Hewitson, 1878) and Salazar and Constantino, 2013 (not Godman, 1901); and Evans, 1955 and (Möschler, 1877). Finally, unless stated otherwise, all subgenera, species, subspecies and synonyms of mentioned genera and species are transferred together with their parent taxa, and taxa not mentioned in this work remain as previously classified.