Three new subfamilies of skipper butterflies (Lepidoptera, Hesperiidae)

Abstract

We obtained and analyzed whole genome data for more than 160 representatives of skipper butterflies (family Hesperiidae) from all known subfamilies, tribes and most distinctive genera. We found that two genera, Katreus Watson, 1893 and Ortholexis Karsch, 1895, which are sisters, are well-separated from all other major phylogenetic lineages and originate near the base of the Hesperiidae tree, prior to the origin of some subfamilies. Due to this ancient origin compared to other subfamilies, this group is described as Katreinae Grishin, subfam. n. DNA sequencing of primary type specimens reveals that Ortholexismelichroptera Karsch, 1895 is not a female of Ortholexisholocausta Mabille, 1891, but instead a female of Ortholexisdimidia Holland, 1896. This finding establishes O.dimidia as a junior subjective synonym of O.melichroptera. Furthermore, we see that Chamunda Evans, 1949 does not originate within Pyrginae Burmeister, 1878, but, unexpectedly, forms an ancient lineage of its own at the subfamily rank: Chamundinae Grishin, subfam. n. Finally, a group of two sister genera, Barca de Nicéville, 1902 and Apostictopterus Leech, [1893], originates around the time Hesperiinae Latreille, 1809 have split from their sister clade. A new subfamily Barcinae Grishin, subfam. n. sets them apart from all other Hesperiidae.

The mitogenome of a Malagasy butterfly Malaza fastuosus (Mabille, 1884) recovered from the holotype collected over 140 years ago adds support for a new subfamily of Hesperiidae (Lepidoptera)

“…The resulting tree topology is consistent with previous phylogenetic studies that did not include Malaza (Sahoo et al 2016;Warren et al 2008Warren et al , 2009Zhang et al 2017aZhang et al , 2019. Coeliadinae are the sister to all other Hesperiidae; Euschemon is a sister to the rest of Hesperiidae except Coeliadinae; the relationship between Eudaminae and Pyrginae is unresolved (0.79 posterior probability) but with Pyrginae sister to Tagiadinae, and Heteropterinae are in the same clade with Hesperiinae (Zhang et al 2019) (the last two subfamilies are monocot feeders as caterpillars, with secondary colonization of dicots by some Hesperiinae).…”

“…examined specimens and preparations of Malaza, including genitalia dissections at NHMUK (H. 828, H. 901, H. 902, BMNH(E) 526064 -dry genitalia on card) and wing venation for M. carmides and M. empyreus, for characters of potential relevance in the systematics of Hesperiidae. Reinforced by the mitogenomic tree, and its morphological distinctness from the other three subfamilies, we conclude that the Malaza lineage should have the same taxonomic rank as Heteropterinae, Trapezitinae, Barcinae, and Hesperiinae (see Zhang et al 2019). To reflect the ancient origin of Malaza and its distinctness from other Hesperiidae, a new subfamily for it is proposed.…”

“…By DNA, a combination of the following base pairs in COI barcode is diagnostic: 38G, 319A, 451T, 475T, 479C, 481T, 512G, and 604A. To derive these DNA characters, we used a barcode alignment of 160 representative Hesperii-dae specimens from all major phylogenetic groups included in Zhang et al (2019). These base pairs are conserved in all three species of Malaza and uniquely place Malaza within the monocot-feeding Hesperiidae to diagnose Malazinae as distinct from the three other subfamilies, i.e., base pairs that are (i) the same as in Heteropterinae but different from most Hesperiinae and Trapezitinae (604); or (ii) the same as in most Hesperiinae but different from most Heteropterinae and Trapezitinae (319, 481); or (iii) the same as in Trapezitinae, but different from most Hesperiinae and Heteropterinae (475); or (iv) the same as most Trapezitinae and Heteropterinae but mostly different from Hesperiinae (38, 512); or (v) the same as in few or most Hesperiinae and Trapezitinae, but different from Heteropterinae (451,475,479).…”

“…Malaza and its junior subjective synonym Manarina Mabille, 1904. The Malazinae is the third monotypic subfamily of Hesperiidae after the Indian Chamundinae (Zhang et al 2019) and the Euschemoninae. The latter consists of a single species, Euschemon rafflesia (MacLeay, [1826]).…”

Phylogenetic position and taxonomic rearrangement ofDavidina(Lepidoptera, Nymphalidae), an enigmatic butterfly genus new for Europe and America

“…In recent years, significant progress has been made in the study of the phylogenetic relationships and taxonomic structure of butterflies (Lepidoptera, Papilionoidea) in general (Wahlberg et al, 2005(Wahlberg et al, , 2014Espeland et al, 2015Espeland et al, , 2018Sahoo et al, 2016;Mitter et al, 2017;Seraphim et al, 2018;Toussaint et al, 2018;Zhang et al 2019;Allio et al, 2020;Wiemers et al, 2020) and the species-rich family Nymphalidae in particular (Wahlberg et al, 2003;2009;Leneveu et al, 2009;Peña et al, 2015;De Moya et al, 2017;Kodandaramaiah et al, 2018;Zhou et al, 2020).…”