Quadrus (Zera) menedemus (Godman and Salvin, 1894)

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 : 13-14

Quadrus (Zera) menedemus (Godman and Salvin, 1894) is a valid species, not a synonym of Quadrus (Zera) tetrastigma (Sepp, [1847])

The genomic tree reveals that Pythonides menedemus Godman and Salvin, 1894 (type locality in Panama: Chiriqui) currently treated as a junior subjective synonym of Quadrus (Zera) tetrastigma (Sepp, [1847]) (type locality in Suriname) is not monophyletic with it and instead is sister to Quadrus (Zera) servius ( Plötz, 1884) (type locality in Brazil) ( Fig. 1 , 2a ). Moreover, genetic differentiation between P. menedemus and Q. tetrastigma is at the level characteristic of distinct species, e.g., their COI barcodes differ by 1.8% (12 bp). Therefore, we propose that Quadrus (Zera) menedemus (Godman and Salvin, 1894) , restored status, is a valid species, not a synonym of Quadrus (Zera) tetrastigma (Sepp, [1847]) . We note that while the name Q. menedemus is defined by its extant syntypes, the identity of Q. tetrastigma could be questioned, because it is identified from the original description and illustrations only; its syntypes likely lost. However, this name has been stably applied to the populations in the Guianas, and we do not have a compelling reason to challenge this interretation. Currently, there is no exceptional need to define Q. tetrastigma by a neotype.

Atarnes Godman and Salvin, 1897 and Eburuncus Grishin, 2012 are subgenera of Milanion Godman and Salvin, 1895

Inspection of the genomic tree reveals close clustering of the three genera: Atarnes Godman and Salvin, 1897 (type species Leucochitonea sallei C. Felder and R. Felder, 1867 ), Eburuncus Grishin, 2012 (type species Leucochitonea unifasciata C. Felder and R. Felder, 1867 ), and Milanion Godman and Salvin, 1895 (type species Papilio hemes Cramer, 1777 ) that taken together are prominently distinct from their relatives ( Fig. 1 ). The tree taxa share a number of phenotypical features and some species in them were misclassified before ( Grishin 2012). Therefore, to simplify the taxonomic classification and reflect all these similarities, we propose to treat Atarnes Godman and Salvin, 1897 , new status, and Eburuncus Grishin, 2012, new status, as subgenera of Milanion Godman and Salvin, 1895 .

Tribe Achlyodini Burmeister, 1878 , subtribe Achlyodina Burmeister, 1878

Figures

Figure 1. The nuclear genome-based phylogeny of Achlyodini (continued in Fig. 3). Only the segment of the tree corresponding to the subtribe Pythonidina is shown. The bottom segment of the tree (subtribe Achlyodinia) is replaced with four dots and is shown in Fig. 3. The tree is constructed from protein-coding regions in autosomes. The tree is rooted with Carcharodus alceae (NVG-18087F02, GenBank barcode OR665734), not displayed. Statistical support values are shown by nodes. For each specimen, its name adopted in this work is given first, and a previously used name is listed in square brackets (if different), supplemented with the DNA sample number, type status (HT holotype, LT lectotype, ST syntype, T type, PT paratype, and PLT paralectotype), general locality and collection year (“old” means collection year known, but likely before 1950, mostly around the turn of 20th century). See Table S1 in the Supplemental file (Zhang et al. 2023d) for additional data about these specimens. Synonyms are given in parentheses preceded by “=”. The type status refers to this synonym if the synonym name is provided. Names of genera (larger font, more prominent tree branches, on the left) and subgenera (smaller font, less prominent tree branches, on the right) are shown by the clades corresponding to these taxa. New genus-group names are highlighted in yellow. Branches leading to the type species (or their closest sequenced relatives) of valid genus-group names are marked with large black dots and refer to the subgenus (in parenthesis) given in the corresponding name, or the genus if the subgenus is not specified. The same notations are used in Fig. 3–6. Names of taxa that before this work were placed in genera Pythonides (red), Gindanes (cyan), Quadrus (green), Ouleus (blue), and Zera (purple) are shown in color to illustrate rampant violations of monophyly in these taxa, corrected here.

Figure 2. Quadrus (Zera) nuclear genome-based phylogeny and type specimens. a) The nuclear genome-based phylogeny of selected species: Q. vivax sp. n. (red), Q. hyacinthinus (purple), Q. hosta (cyan), Q. servius stat. rest. (blue and magenta, with its synonym Achlyodes erisichthon labeled in blue) rooted with the remaining Zera species (not shown; see Fig. 1). Specimens shown below are highlighted in yellow; see Fig. 1 for other notations. b) Lectotype of Quadrus (Zera) servius designated herein, with its labels shown above on the right. c) Holotype of Quadrus (Zera) vivax sp. n. Specimens are in dorsal (left) and ventral (right) views, data in text. The scale bar at the bottom refers to specimens; the scale bar at the top refers to labels reduced by 1/3 compared to specimens.