Agyllia Grishin

Zhang, Jing, Brockmann, Ernst, Cong, Qian, Shen, Jinhui & Grishin, Nick V., 2020, A genomic perspective on the taxonomy of the subtribe Carcharodina (Lepidoptera: Hesperiidae: Carcharodini), Zootaxa 4748 (1), pp. 182-194 : 186-187

Agyllia Grishin , gen. n.

http://zoobank.org/ 095B9432-5CCE-4CBF-8EB6-B9711FDABA25

Type species: Pyrgus agylla Trimen, 1889 ( Fig. 2c ).

Diagnosis. Keys to 2 in de Jong (1978: 28), constituting his asterodia species group. Morphologically differs from close relatives by the following characters. Out of three spots in forewing discal cell, rectangular middle spot (the largest) closer to streak-like spot at distal end of cell than to well-developed and rounded basal spot; no dorsal white spots at base of CuA 2 -1A+2A cell (space 1B). Ventral hindwing with a straight median white band, i.e., a white spot in cell Rs-M 1 (space 6) joins the central spot (discal cell) to the outer (and not inner) spot in cell Sc+R 1 -Rs (space 7). In male genitalia, uncus deeply incised; valva with large costal process and harpe (=cucullus) lacks a fold covering the costal process. In DNA COI barcode region, a combination of the following base pairs is diagnostic: A307T, A352T, T364C, C401T, T403A, T500C, and A502T.

Derivation of the name. The name is a feminine noun in the nominative singular derived from the name of the type species.

Species included: Encompasses asterodia species groups as it was defined by de Jong (1978). Full species list is given below.

Second, we observe that Carcharodus is not monophyletic. Only one species, Carcharodus tripolina (Verity, 1925) groups with the type species of the genus Carcharodus alceae (Esper, 1780) . These results are consistent with the recent treatment by Coutsis (2016), who placed all other Carcharodus species in Reverdinus Ragusa, 1919 . In our trees, Reverdinus is in the same cluster with Muschampia Tutt, 1906 and the branch length separating Reverdinus from other Muschampia is not significantly larger than the branch lengths separating Muschampia species from each other. Thus, we consider Reverdinus Ragusa, 1919 to be a subgenus of Muschampia . Additionally, we see that genus names previously given to various groups currently placed in Muschampia indeed denote monophyletic groups within the genus and we suggest to treat these groups as subgenera: Warrenohesperia Strand, 1928 , Sloperia Tutt, 1906 and Tuttia Warren, 1926 ( Fig. 1 ).

Third, we find that “ Muschampia ” cribrellum (Eversmann, 1841) , the type species of the genus Favria Tutt, 1906 is not monophyletic with Muschampia . Instead, it is a confident sister of Gomalia in nuclear genome trees ( Fig. 1 ab). Its phylogenetic position is not very strongly supported in the mitogenome tree (88% bootstrap, Fig. 1c ), but it is well-separated from Muschampia . This species has been a puzzle and is uniquely characterized by spined mid-tibiae. Therefore we reinstate Favria as a valid genus, currently monotypic.

Fourth, we see that the holotype of Tavetana jeanneli Picard, 1949 ( Fig. 3 ) is not a dark form of Gomalia elma (Trimen, 1862) as currently considered, but a Gomalia species well removed from it. COI barcodes of the two species differ by nearly 7% (45 base pairs). Moreover, the differences in genitalia of the Indian Gomalia elma albofasciata Moore, 1879 (see plate 23, D 2 in Evans, 1949) and the African nominal subspecies (plate 13 in Evans, 1937) argue for the species status of the Indian taxon. Most notably, ampulla of male genitalic valva is expanded in C. albofasciata compared to C. elma , in which costa smoothly transitions to a tooth-like ending of harpe.

Furthermore, we elevate to species Spialia lugens (Staudinger, 1886) and Spialia carnea (Reverdin, 1927) formerly considered subspecies of Spialia orbifer (Hübner, [1823]) . Sequencing of S. lugens and S. carnea type specimens in the Berlin Museum für Naturkunde reveals 2.2%-3.2% difference in COI barcode from nominotypical populations of S. orbifer . Distinct barcodes combined with the differences in facies suggest species-level status for these taxa. Spialia lugens differs from the two other species by the larger size, darker wing above with faint or absent submarginal sports, rarely, and mostly in females, better developed ( de Jong, 1978). S. carnea is characterized by warm reddish to brown-yellow color of hindwing below and reduced submarginal spots on hindwing below in cells M 1 -M 2 and M 2 -M 3.

Finally, difference in male genitalia, notably the shape of uncus ( de Jong, 1978; Evans, 1937), suggest that Ernsta bifida (Higgins, 1924) , a species distinct from Ernsta zebra (Butler, 1888) and not its subspecies. Taken together, the data we obtained suggest the following taxonomic arrangement of the subtribe Carcharodina.

Taxonomic arrangement of the subtribe Carcharodina. Based on our analysis, the list of species arranged into genera and subgenera is given below. Synonymic names are included for genera and subgenera. Names treated as synonyms (genera and names of type species that are considered to be synonyms) are preceded by “=”: not followed by daggers are subjective junior synonyms; † objective junior synonyms; ‡ unavailable names (such as homonyms and nomina nuda); “preocc.” indicates preoccupied, the taxonomic order (all insects) of the senior name is shown in brackets. Synonyms are attributed to subgenera. Type species (TS) for genera and subgenera are listed and underlined. For type species that are considered to be synonyms, valid names are shown in parenthesis. For valid genera and subgenera (not their synonyms), names of the type species or names which type species are considered to be synonyms of, are underlined in the list. Subspecies names are not listed pending further studies.

Figures

FIGURE 1. Phylogenetic trees of Carcharodina. The trees are constructed from nucleotide sequences of protein-coding regions from: a. nuclear genome; b. Z-chromosome; c. mitochondrial genome. Statistical significance values are shown by each node. d. COI barcode distance diagram is shown to emphasize on close relationships within Carcharodina. In panels a) and b), vertical green (yellow-shaded) line defines genera: each clade crossed by the line is a genus in our treatment; vertical thin gray (blue-shaded) line defines subgenera. Branches in Carcharodina are colored by genus.

FIGURE 2. Specimens of Ernsta and Agyllia. Dorsal and ventral views are on the left and right, respectively. DNA sample IDs are given, other data are in the Tab. S1: a. Ernsta colotes the type species of the genus Ernsta gen. n., NVG-18054F06; b. Ernsta delagoae, the type species of the subgenus Delaga subgen. n., NVG-18054F08; c. Agyllia agylla, the type species of the genus Agyllia gen. n., NVG-19039F03.

FIGURE 3. Gomalia jeanneli holotype. The specimen is in the Muséum National d’Histoire Naturelle, Paris, France. Dorsal and ventral views are on the left and right, respectively. The largest scale bar refers to the specimen, labels are shown at 1/3 of specimen size and genitalia enlarged twice compared to the specimen. DNA sample ID NVG-18079B11.