Papias integra ( Mabille, 1891 )

Zhang, Jing, Cong, Qian, Shen, Jinhui & Grishin, Nick V., 2022, Taxonomic changes suggested by the genomic analysis of Hesperiidae (Lepidoptera), Insecta Mundi 2022 (921), pp. 1-135 : 75-76

Papias integra ( Mabille, 1891) , reinstated status

Genomic trees reveal a prominent partitioning of specimens identified as Papias subcostulata (Herrich-Schäffer, 1870) (type locality Brazil) into two groups ( Fig. 14 ). Phenotypically, these groups correspond to the two subspecies defined by Evans (1955): the nominotypical subspecies and Papias subcostulata integra Mabille, 1891 (type locality Honduras). Although Mielke and Casagrande (2002), who designated the lectotypes of integra and Hesperia infuscata Plötz, 1882 (type locality “ Brazil ” in the original description, but Suriname on the label of the lectotype), did not advocate the two Evans’ subspecies, our genomic comparison suggests that they are distinct species with H. infuscata being a junior subjective synonym of the nominal P. subcostulata , and P. s. integra being the senior name for the second species. The two species are characterized by Fst/Gmin values of 0.45/ 0.008 in the Z chromosome. According to a recent study ( Cong et al. 2019a), these values indicate strong genetic differentiation and suggest very low gene exchange between these species, thus supporting their distinction from each other. Evans (1955) mentioned that the two “subspecies” overlap in distribution. We sequenced specimens of both from Ecuador (although not from exactly the same localities). For these reasons (genetic and phenotypic distinction, possible sympatry), we reinstate Papias integra ( Mabille, 1891) as a species-level taxon. Due to extensive pale overscaling along the veins below, we place Pamphila allianca Weeks, 1901 (type locality Bolivia) as a junior subjective synonym of P. subcostulata . Interestingly, all the specimens we sequenced have the same COI barcode sequence and thus offer yet another example of distinct species that, like North American Celastrina Tutt, 1906 , do not differ in their barcodes.

Finally, as Mielke and Casagrande (2002) mentioned, type specimens of P. subcostulata are probably lost. They were illustrated by Plötz among his unpublished drawings (no. 286 for P. subcostulata ). These drawings were inspected by Godman (1907), who organized copying many of them, with these copies bound in the library of the Natural History Museum, London. Although the drawing no. 286 was not copied, Godman (1907: 138) made a comment that P. subcostulata may belong to Papias and a specimen from Iquitos in Godman and Salvin collection may be this species. We located this specimen in BMNH, and it has prominent pale overscaling along the veins, in particular on the hindwing, agreeing with Evans’ concept of P. subcostulata . Furthermore, illustrations in Draudt (1921 –1924) frequently resemble Plötz unpublished drawings seen as Godman copies, and more so than they resemble actual type specimens Plötz drawings were made of. Therefore it seems likely that many Draudt illustrations were made not from specimens, but from Plötz drawings. For this reason, Draudt illustrations may be viewed as (frequently inferior) copies of Plötz and probably depict the types of these taxa. The illustration of P. subcostulata (plate 187f in Draudt 1921 –1924) shows ventral side with prominent pale veins and agrees well with the BMNH specimen from Iquitos selected to match Plötz drawing of P. subcostulata . Thus, although the type specimens of P. subcostulata could not be found, several lines of evidence presented here point to the same conclusion that it is the species with paler veins. Therefore, we agree with the Evans’ (1955) identification of P. subcostulata .

Figures

Figure 14. Genomic tree of Moncina, part 2. See Fig. 1 legend for notations.