A genomic perspective on the taxonomy of the subtribe Carcharodina (Lepidoptera: Hesperiidae: Carcharodini)
We obtained whole genome shotgun sequences and phylogenetically analyzed protein-codingregions of representative skipper butterflies from the genus Carcharodus Hubner, [1819] and its closerelatives. Type species of all available genus-group names were sequenced. We find that species attributed tofour exclusively Old World genera (Spialia Swinhoe, 1912, Gomalia Moore, 1879, Carcharodus Hubner, [1819]and Muschampia Tutt, 1906) form a monophyletic group that we call a subtribe Carcharodina Verity, 1940. Inthe phylogenetic trees built from various genomic regions, these species form 7 (not 4) groups that we treat asgenera. We find that Muschampia Tutt, 1906 is not monophyletic, and the 5th group is formed by currentlymonotypic genus Favria Tutt, 1906 new status (type species Hesperia cribrellum Eversmann, 1841), which issister to Gomalia. The 6th and 7th groups are composed of mostly African species presently placed in Spialia.These groups do not have names and are described here as Ernsta Grishin, gen. n. (type species Pyrgus colotesDruce, 1875) and Agyllia Grishin, gen. n. (type species Pyrgus agylla Trimen, 1889). Two subgroups arerecognized in Ernsta: the nominal subgenus and a new one: Delaga Grishin, subgen. n. (type species Pyrgusdelagoae Trimen, 1898). Next, we observe that Carcharodus is not monophyletic, and species formerly placedin subgenera Reverdinus Ragusa, 1919 and Lavatheria Verity, 1940 are here transferred to Muschampia.Furthermore, due to differences in male genitalia or DNA sequences, we reinstate Gomalia albofasciataMoore, 1879 and Gomalia jeanneli (Picard, 1949) as species, not subspecies or synonyms of Gomalia elma(Trimen, 1862), and Spialia bifida (Higgins, 1924) as a species, not subspecies of Spialia zebra (Butler, 1888).Sequencing of the type specimens reveals 2.2-3.2% difference in COI barcodes, the evidence that combinedwith wing pattern differences suggests a new status of a species for Spialia lugens (Staudinger, 1886) andSpialia carnea (Reverdin, 1927), formerly subspecies of Spialia orbifer (Hubner, [1823]).