Grishin, Zhang & Cong & Shen & Song & Grishin, 2023

Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina & Grishin, Nick V., 2023, Genomics-based taxonomic rearrangement of Achlyodini and Carcharodini (Lepidoptera: Hesperiidae: Pyrginae), Insecta Mundi 2023 (16), pp. 1-33 : 29-30

Capilla Grishin , new subgenus

http://zoobank.org/ 942D19CE-7B9C-47E7-9650-1721B9A18AB4

Type species. Helias aurocapilla Staudinger, 1876 , currently a junior subjective synonym of Hesperia musculus Burmeister, 1875 .

Definition. The nuclear genomic tree reveals that Staphylus Godman and Salvin, 1896 (type species Helias ascalaphus Staudinger, 1876 ) splits into five clades at the subgenus level ( Fig. 5 ). The subgenera corresponding to two of these clades, which are sister to each other, do not have available names and are new. One of these new subgenera is phenotypically diverse, keying to E.32.3, 8, 10a, 11a, 13d, 19, 21, or 22 in Evans (1953) and is distinguished from its relatives by the following characters: wings typically narrower than in relatives, several species with more extensive white/hyaline forewing spots; the only subgenus that includes species with divided, rounded (except the very tip), or bulbous at the tip uncus, but uncus undivided and narrow, nearly needle-like in some species, and not strongly concave in lateral view, tegumen usually broader than uncus, vinculum relatively straight in lateral view, harpe mostly broad, with rounded or truncate distal margin, overlays ampulla from the inside (at least in species with lanceolate uncus), ampulla frequently expanded and projects to the end of harper in some species, aedeagus frequently shorter and broader than in relatives. Confident identification is provided by DNA, and a combination of the following base pairs is diagnostic: in the nuclear genome: aly1405.20.35:T489C, aly1405.20.35:T444C, aly1405.20.35:G452C, aly 1370.7.2:A2191G, aly 1370.7.2:T804G, and the COI barcode T29(mostly T), A130Y, A241(mostly T), A256T, T412(mostly A), T578(mostly T), A580(mostly A).

Etymology. The name is a feminine noun in the nominative singular, formed from the type species name: [auro] Capilla.

Species included. Pholisora lizeri Hayward, 1938 , Pholisora caribbea R. Williams and E. Bell, 1940 , Pholisora buena R. Williams and E. Bell, 1940 , Staphylus melius Steinhauser, 1989 , Hesperia musculus Burmeister, 1875 , Pholisora imperspicua Hayward, 1940 , Pholisora corumba R. Williams and E. Bell, 1940 , Helias ascalon Staudinger, 1876 , Staphylus eryx Evans, 1953 , Hesperia melangon Mabille, 1883 , Nisoniades tucumanus Plötz, 1884 , Staphylus perna Evans, 1953 , Staphylus insignis O. Mielke, 1980 , Helias tyro Mabille, 1878 , and Pholisora azteca Scudder, 1872 .

Parent taxon. Genus Staphylus Godman and Salvin, 1896 .

Composition of Carcharodina Verity, 1940 with comments on nomenclature

The subtribe Carcharodina Verity, 1940 consists of the following genera: Noctuana E. Bell, 1937 , Windia H. Freeman, 1969 , Carcharodus Hübner, [1819] , Muschampia Tutt, 1906 (the senior name for this genus is Sloperia Tutt, 1906 , see below), Favria Tutt, 1906 , Gomalia Moore, 1879 , Agyllia Grishin, 2019, Ernsta Grishin, 2019, and Spialia Swinhoe, 1912 ( Fig. 6 ). We phylogenetically arrange genera in the subtribe to start with the New World representatives, because they would follow all the rest of Carcharodini that are from the New World, thus not disrupting the geographic arrangement. The ordering ends with the strongly checkered in wing pattern members of the subtribe, such as Spialia , because the next tribe to follow in the taxonomic list would be Pyrginae Burmeister, 1878. To link the two tribes by phenotypic similarity of their constituent species, Pyrginae could start from the Old World checkered skippers in the genus Pyrgus Hübner, [1819] ), which are superficially similar to Spialia and many Muschampia , and are the only representatives of Pyrginae in the Old Word. Furthermore, genera in the two pairs ( Favria , Muschampia ) and ( Carcharodus , Muschampia ) best be next to each other due to phenotypic similarity between some species in each pair, and the phylogeny additionally constrains the pairing ( Favria , Gomalia ). We choose to place Carcharodus first due to some wing pattern similarity with Windia , and because this order results in a continuous placement of African members (in Gomalia , Agyllia , and Ernsta ), ending with mostly Palearctic Spialia to follow with Palearctic and similar-looking Pyrgus .

In agreement with the ICZN Code, the name Sloperia Tutt, 1906 takes precedence over Muschampia Tutt, 1906 , as selected by the first reviser ( Warren 1926) and should be the correct name to use instead of Muschampia , as pointed out by Hemming (1967). However, judging by the literature ( Wiemers et al. 2018; Wiemers et al. 2020) and discussions with colleagues, the prevailing usage of Muschampia is preferred by most of them. We do not have a solution to this problem, and it joins the list of similar problems when there is a disagreement between the articles of the ICZN Code and the community opinion (e.g., gender agreement ICZN Code articles vs. the community of Lepidopterists is an example of a more significant, but similar in spirit, issue). We see the advantages of both treatments in this specific case. Indeed, Muschampia has been a widely used name familiar to many people, and it seems that the community opinion should outweigh the priority of a semi-random decision made by the first reviser. Conversely, the rules are universal, clearly spelled out, and long-term, but the community opinions are subject to change, especially with generational changes. Furthermore, the name Sloperia is shorter and may be easier for newcomers to learn, fitting the array of other names in the subtribe, such as Spialia , Gomalia , and Favria . Whatever the decision, it is not a question to be solved by scientific tools. It is possible that after some period of deliberation, the community might embrace Sloperia (it is not a bad option at all), or the community would be more ready to make a stronger case for ICZN to issue an opinion. However, whichever path is taken, we strongly oppose solving this (and other) nomenclatural problem by fiddling with taxonomy just so that the name Muschampia is still in use (e.g., with the sole purpose of accommodating the use of Muschampia , treating both Muschampia and Sloperia as valid genera).

Figures

Figure 5. The nuclear genome-based phylogeny of Carcharodini, middle segment (continues in 6, see Fig. 4 for the first segment) showing Pholisorina subtrib. n. Species formerly placed in Staphylus are labeled in red. Asterisk marks a species with a dedicated study (Lemes et al. 2023) that already assigned it to Bolla. See Fig. 1 for notations.

Figure 6. The nuclear genome-based phylogeny of Carcharodini, the last segment (continues from Fig. 4 and 5) showing Carcharodina. Different genera are shown in different colors. See Fig. 1 for notations. The red asterisk on a tree branch denotes the arrival of Carcharodina in the Old World from the New World. See the discussion about Muschampia (a junior name) vs. Sloperia (should have priority) in the text.