Habrodais Scudder, 1876

Zhang, Jing, Cong, Qian, Shen, Jinhui, Opler, Paul A. & Grishin, Nick V., 2020, Genomic evidence suggests further changes of butterfly names, The Taxonomic Report of the International Lepidoptera Survey 8 (7), pp. 1-41 : 13-14

Habrodais Scudder, 1876 , Favonius Sibatani & Ito, 1942 , Neozephyrus Sibatani & Ito, 1942 , Quercusia Verity, 1943 , Chrysozephyrus Shirôzu & Yamamoto, 1956 , and Sibataniozephyrus Inomata, 1986 are junior subjective synonyms of Hypaurotis Scudder, 1876

Inspecting genomic phylogenetic trees of US butterfly species ( Zhang et al. 2019d), we noticed a close relationship between the only two New World genera from the tribe Theclini Swainson, 1830 : Hypaurotis Scudder, 1876 (type species Thecla crysalus W. H. Edwards, 1873 ) and Habrodais Scudder, 1876 (type species Thecla grunus Boisduval, 1852 ). Despite dissimilar wing patterns and colors, divergence between the type species of these genera is indeed comparable to that of congeners (Fig. 6 top) and is even lower than the divergence in some compact genera, such as Chlorostrymon Clench, 1961 , Ministrymon Clench, 1961 or Electrostrymon Clench, 1961 (Fig. 6 top), and particularly in more diverse genera such as Strymon Hübner, 1818 , Callophrys Billberg, 1820 , or Satyrium Scudder, 1876 ( Zhang et al. 2019d). COI barcodes of Hy. crysalus and Ha. grunus are only 4.3% (28 bp) different: divergence similar to that at times reported for different individuals of the same species ( Zakharov et al. 2009; Kodandaramaiah et al. 2013), strongly suggesting that these two species are congeneric. The combination Hypaurotis grunus has been used previously in publications ( Garth 1934), however, we failed to find the combination Habrodais crysalus published. Also, H. crysalus has a slightly broader distribution and is a more familiar butterfly. Therefore, among Fig. 6. Habrodais and others. Nuclear Hypaurotis , genome tree above, the two names published in the same work COI barcode dendrogram below. Specimens without locality given are ( Scudder 1876), we use Hypaurotis as from GenBank and their accession numbers are indicated. valid, and Habrodais as its junior subjective synonym, resulting in Hypaurotis grunus (Boisduval, 1852) and Hypaurotis poodiae J. Brown & Faulkner, 1982 , revised and new combinations.

Next, when a genomic dataset of an Old World species Favonius quercus (Linnaeus, 1758) was included in the Theclinae tree, it clustered closely with Hypaurotis (Fig. 6 top). The COI barcodes of H. crysalus and F. quercus differ by 4.7% (31 bp). Because we lack genomic data for other genus-group names from the Thecla section of Eliot ( Eliot 1973), we downloaded available COI barcode data from GenBank ( Sayers et al. 2020). We find that while for most species pairs, e. g., H. crysalus and Favonius orientalis (Murray, 1875) , the barcodes are more similar (6%), for others, e. g., H. crysalus and Thecla betulae (Linnaeus, 1758) (the type species of Thecla Fabricius, 1807 ) the difference is larger (8.2%). The COI barcode distance dendrogram computed using BioNJ ( Gascuel 1997) as implemented by the phylogeny.fr server ( Dereeper et al. 2008) reveals close clustering of species we propose to place in Hypaurotis (Fig. 6 bottom, red) and separation of T. betulae (green) from this cluster. In the absence of genomic data, this COI barcode analysis of their type species suggest that in addition to Habrodais Scudder, 1876 and Quercusia Verity, 1943 (type species Papilio quercus Linnaeus, 1758 ), the following four genus-group names Sibataniozephyrus Inomata, 1986 (type species Zephyrus fujisanus Matsumura, 1910 ), Neozephyrus Sibatani & Ito, 1942 (type species given as Thecla taxila Bremer, 1861 , which was, however, a misidentified Dipsas japonica Murray, 1875 ; according to the Art 70.3.2. of the ICZN Code the actual taxonomic identity of this species is chosen and japonica is fixed as type species), Chrysozephyrus Shirôzu & Yamamoto, 1956 (type species Thecla smaragdina Bremer, 1861 ) and Favonius Sibatani & Ito, 1942 (type species Dipsas orientalis Murray, 1875 ) are junior subjective synonyms of Hypaurotis Scudder, 1876 , which is a genus distinct from Thecla Fabricius, 1807 . In accord with genetic similarities, all these species are similar in appearance ( Eliot 1973). We expect that future studies will reveal additional synonyms and possibly a subgeneric structure of Hypaurotis .

Finally, even from a practical standpoint of American butterfly knowledge, it seems more instructive to treat H. crysalus and H. grunus as congeneric emphasizing on their close kinship (despite apparent phenotypic dissimilarity), instead of placing them in two monotypic (or nearly monotypic) genera that accentuate their tenuous (but superficial) uniqueness. Finding their close relatives in the Old World places Hypaurotis among other Holarctic Theclinae genera such as Callophrys and Satyrium and emphasizes somewhat unusual but recurrent pattern revealing the connection between the Old and the New Worlds.