Panacea Godman & Salvin, 1883

Zhang, Jing, Cong, Qian, Shen, Jinhui, Opler, Paul A. & Grishin, Nick V., 2021, Genomics-guided refinement of butterfly taxonomy, The Taxonomic Report of the International Lepidoptera Survey 9 (3), pp. 1-55 : 37-40

Panacea Godman & Salvin, 1883 is a junior subjective synonym of Batesia C. Felder & R. Felder, 1862

Introductory comments on Nymphalinae

Five genera from the subfamily Nymphalinae Rafinesque, 1815 were placed as incertae sedis by Wahlberg (2019a): Pycina Doubleday 1849 (type and the only species Pycina zamba Doubleday, [1849] ), Rhinopalpa C. & R. Felder 1860 (type and the only species Rhinopalpa fulva C. & R. Felder, 1860, a junior subjective synonym of Vanessa eudoxia Guérin-Méneville, 1840 , with is a subspecies of Papilio polynice Cramer, 1779 )), Kallimoides Shirôzu & Nakanishi 1984 (type and the only species Kallima rumia Doubleday, 1849 ), Vanessula Dewitz 1887 (type and the only species Vanessula buchneri Dewitz, 1887 , a junior subjective synonym of Liptena milca Hewitson, 1873 ), and Doleschallia C. & R. Felder 1860 (type species Papilio bisaltide Cramer, 1777 ). The reasons behind the incertae sedis placement, such as ancient origin of these taxa forming long branches in the trees, were given by Wahlberg (2019b), who concluded: "Clearly more data are needed for these ancient taxa." To shed light on their classification, we obtained whole genome shotgun for the type species of these five genera and placed them in the phylogenetic context of other Nymphalinae . The tree constructed from protein-coding genes of the Z chromosome is illustrated ( Fig. 42). In this mostly well-supported phylogeny, all currently defined tribes receive 100% statistical support. The five taxa in question are found in deeper radiation near the origins of Nymphalinae and are not closely associated with any of the currently defined tribes. Pycina zamba and Rhinopalpa polynice are confidently placed prior to divergence of other tribes such as Victorinini Scudder, 1893 and Junoniini Reuter, 1896 ( Fig. 42). Doleschallia bisaltide is a confident sister to Melitaeini Herrich-Schäffer, 1843 , but is prominently separated from them, originating well prior to radiation of Melitaeini into subtribes. Notably, Melitaeini are characterized by an elevated evolutionary rate reflected in much longer branches within the tribe compared to those of most other Nymphalinae , including Doleschallia . Finally, although Kallimoides rumia and Vanessula milca are confidently placed in the clade consisting of Victorinini, Junoniini, Melitaeini and Doleschallia , their exact phylogenetic position remains weakly supported because they originated during a rapid radiation event near the last common ancestor of this clade. It is likely that K. rumia is a distant sister of Victorinini. However, the association of V. milca with Junoniini is questionable. It is not likely that the lack of confidence in this phylogenic placement is caused by the shortage of sequence data. It is plausible that these taxa are of hybrid origin, or their evolution involved incomplete lineage sorting resulting in phylogenetic incongruence among different genes. Regardless of their exact phylogenetic origins, and exactly due to this phylogenetic uncertainty, these taxa do not belong to any existing tribes. For all the reasons stated above, each of these five lineages represents a tribe of Nymphalinae , as named below.

Pycinini Grishin, new tribe

http://zoobank.org/ D5C0AE41-2615-4852-9B59-8533C27E4798

Type genus. Pycina Doubleday 1849 .

Definition. Formerly placed within the tribe Coeini Scudder, 1893 , this tribe is not monophyletic with it and instead is sister to all other Nymphalinae Rafinesque, 1815 excluding Coeini ( Fig. 42), which justifies its status as a new tribe. The tribe is diagnosed by the characters given for Pycina on pages 305–306 by Westwood (1850) and on page 330 by Godman & Salvin (1884). In brief, diagnosed by densely hairy eyes, relatively small and thin palpi not extending beyond middle of eyes in lateral view, long slender antennae more than half of forewing length, elongated forewings, more so than in similarly patterned Smyrna Hübner, [1823] , and undulating costal margin of hindwing: i.e., prominently concave near the middle instead of evenly convex throughout as in its relatives.

Genera included. Only the type genus.

Parent Taxon. Subfamily Nymphalinae Rafinesque, 1815 .

Rhinopalpini Grishin, new tribe

http://zoobank.org/ 4A3ACC99-A761-49B9-BA00-00F83FF443DE

Type genus. Rhinopalpa C. & R. Felder 1860 .

Definition. Originates early in the Nymphalinae radiation and is sister to the clade consisting of several tribes, e.g., Kallimini Doherty, 1886 and Melitaeini Herrich-Schäffer, 1843 among others ( Fig. 42), indicating that this new taxon is a tribe. Diagnosed by long and densely scaled stout palpi (as suggested by the name, but not longer than the head as in Libytheinae Boisduval, 1833 ), discal cells short on both wings, about ⅓ of wing length, all forewing R veins run very close together and R 3, R 4 and R 5 stalked for most of their length, forewings produced into apical and tornal lobes, hindwings with stubby tail in the middle of outer margin; below with a row of 6 eyespots along the margin of both wings; caterpillar with long scoli turned anteriad at tips past thoracic segments; pupa unusually shaped somewhat resembling a dead leaf, with 3 pairs of horn-like protuberances, first pair very prominent, directed posteriad.

Genera included. Only the type genus.

Parent Taxon. Subfamily Nymphalinae Rafinesque, 1815 .

Comments. The name Rhinopalpini as published by Teshirogi (2016) appears to be a nomen nudum: Referred to Wahlberg et al. (2005), a work that does not mention the name, and the name is not "explicitly indicated as intentionally new" in Teshirogi (2016), thus failing Art. 16.1. of the ICZN Code ( ICZN 1999).

Kallimoidini Grishin, new tribe

http://zoobank.org/ 22C74A4F-6B24-4502-99BB-B8E3C66AFABE

Type genus. Kallimoides Shirôzu & Nakanishi 1984 .

Definition. This tribe is a distant sister of Victorinini Scudder, 1893 with moderate support ( Fig. 42). Being an ancient lineage as indicated by its phylogenetic placement and only moderate statistical support for association with Victorinini, it is assigned the status of a tribe. Diagnostic characters for this tribe are those given in detail for the genus Kallimoides on pages 107-108 and wing venation and genitalia are illustrated in Figs. 5 and 40-45, respectively, by Shirôzu & Nakanishi (1984). Most notably, the new tribe is diagnosed by S-shaped vein R 1 on forewing, partially coalescing with Sc and then with R 2; prominently reduced tegumen and large S-shaped aedeagus terminally thinning into a hook in male genitalia; and the absence of genital plate in females.

Genera included. Only the type genus.

Parent Taxon. Subfamily Nymphalinae Rafinesque, 1815 .

Vanessulini Grishin, new tribe

http://zoobank.org/ D584B74C-8DC8-479A-AD25-9197A03A843B

Type genus. Vanessula Dewitz 1887 .

Definition. Belongs to the same clade with Victorinini, Junoniini, Melitaeini and Doleschallia , but not closely allied to any of them as judged by its ancient origin and poor statistical support for its placement within this clade, justifying its status of a tribe ( Fig. 42). The tribe is diagnosed by the characters given for Vanessula on pages 145–146 by Dewitz (1887). In brief, forewing vein R 1 near its origin partly coalescent with Sc, vein R 2 stalked with R 3, R 3 with R 4 and R 4 with R 5 for at least half of their lengths each, discal cell closed, short, about 1/3 of forewing; eyes bare, palpi long and thin, slightly longer than head, antennae half of forewing length; wings moderately elongated, wing shape similar to Melitaeini .

Genera included. Only the type genus.

Parent Taxon. Subfamily Nymphalinae Rafinesque, 1815 .

Doleschalliaini Grishin, new tribe

http://zoobank.org/ 88D32044-DB8E-486F-8C09-0D188E5CDC93

Type genus. Doleschallia C. & R. Felder 1860 .

Definition. Formerly included in Kallimini Doherty, 1886 due to extensive superficial similarity. In the genomic tree, confidently placed as sister to Melitaeini Newman, [1870] , but more distant from them than they are from each other, and is of an ancient origin estimated nearly 70 Mya ( Su et al. 2017), which justifies its status of a tribe ( Fig. 42). Similar to Kallima Doubleday 1849 in wing shape and coloration, apparently due to convergence. Diagnosed by its produced forewing apex and hindwing tornus extended in a short tail, below colored as a dead brown leaf with mid-rib (as in Kallimini ), with several eyespots along the wing margins; discal cells open on both wings (closed in Kallimini ), all forewing R veins close together, R 4 and R 5 stalked for 2/3 of their length, hindwing humeral vein simple (branches into 2 in Kallimini ); eyes bare, pupa smooth and rounded, abdomen segments without protuberances that are present in Kallimini .

Genera included. Only the type genus.

Parent Taxon. Subfamily Nymphalinae Rafinesque, 1815 .

Comments. Doleschalla Walker 1861 ( Diptera : Tachinidae ) is the type genus of Doleschallini . According to Art. 55.4. of the ICZN Code ( ICZN 1999), one letter difference, which in this case would be an extra "i" from the suffix "-ina", avoids homonymy. However, it does not avoid confusion, including internet search engines that correct possible spelling errors, especially in this case, because it is a duplication of a letter in the suffix formation from the correctly determined stem. The Code gives a choice to alleviate the problem: Art. 29.4. (original stem formation to be maintained) and Recommendation 29A are followed here, and the entire name of the type genus is taken as a stem with hopes to diminish confusions. The choice of being ridiculed for grammatically incorrect stem formation is a "lesser evil" compared to increased confusion of those who use the name.

Additional genomic support to partition Nymphalis genus group into 4 genera: Hypanartia Hübner, [1821] , Vanessa [Fabricius], 1807, Antanartia Rothschild & Jordan, 1903, and Nymphalis Kluk, 1780

To achieve a more objective, internally consistent and eventually stable classification, we have placed Aglais Dalman, 1816 (type species Papilio urticae Linnaeus, 1758 ) and Polygonia Hübner, [1819] (type species Papilio c-aureum Linnaeus, 1758 ) as subgenera of Nymphalis Kluk, 1780 (type species Papilio polychloros Linnaeus, 1758 ) on the basis of genomic evidence ( Zhang et al. 2020). Here, we obtained and