Siderone Hübner, [1823]

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 : 29-32

Siderone Hübner, [1823] and Phantos Dias, 2018 are subgenera of Zaretis Hübner, [1819]

Genetic differentiation within the clade of three genera: Siderone Hübner, [1823] (type species Siderone ide Hübner, [1823] , a junior subjective synonym of Papilio nemesis Illiger, 1801 ), Phantos Dias, 2018 (type species Nymphalis callidryas R. Felder, 1869 ) and Zaretis Hübner, [1819] (type species Papilio isidora Cramer, 1779 ) is less than that of Memphis and Anaea (sensu lato, as above) (Fig. 31), suggesting that Siderone and Phantos are subgenera of Zaretis to restore internal consistency of the classification. According to Fig. 3 in Toussaint et al. (2019), all non-monotypic genera of Anaeini as we define them ( Anaea , Memphis , Consul and Zaretis ) diversified around the same time 17-20 Mya, indicating internal consistency of our definition; and monotypic genera Coenophlebia C. & R. Felder, 1862 (type and the only species Siderone archidona Hewitson, 1860 , we have not sequenced this species yet) and Hypna Hübner, [1819] (type species Papilio clytemnestra Cramer, 1777 ) are more distant from others to maintain their distinction.

Maniolina Grote, 1897 and Melanargiina Wheeler, 1903 are junior subjective synonyms of Erebiina Tutt, 1896 and Satyrina Boisduval, 1833 , respectively Paralasa Moore, 1893 belongs to Ypthimina Reuter, 1896

Listed among the "Subtribe uncertain" genera by Wahlberg (2019a), Paralasa Moore, 1893 (type species Erebia kalinda Moore, 1865 ) is a confident sister to Ypthimina Reuter, 1896 (Fig. 32). Therefore, instead of proposing a monotypic subtribe for this genus, we include Paralasa in the subtribe Ypthimina .

Callerebiina Grishin, new subtribe

http://zoobank.org/ 46A68C99-BBC2-4F6C-BB06-4CA2AD2343B8

Type genus. Callerebia Butler, 1867 .

Definition. Several genera of uncertain tribal placement ( Wahlberg 2019a) confidently grouped with others sometimes placed in Ypthimina Reuter, 1896 (Fig. 32 red). Close relationship of Callerebia , Loxerebia and Argestina that are in the red clade has been reported before ( Yang and Zhang 2015). This prominent red clade is indeed a weakly supported sister to Ypthimina (Fig. 32 blue and cyan). However, due to the weak statistical support (therefore possibly erroneous sister relationship with Ypthimina ) and the origin of both clades near the rapid radiation of many Satyrini subtribes, this clade is proposed as a new subtribe. A phenotypically diverse assembly of species, generally characterized by Erebia -like appearance, typically with a row of orange black-centered eyespots on each wing, but these could be reduced to a couple of eyespots, similar to Ypthimina , or wings could be unspotted brown or even mostly white above (with brown borders) in some species; gnathos developed (absent in many Ypthimina ), but arms shorter than uncus, saccus reduced, valva rather stout, flattened and rounded or excavated near the apex, aedeagus rather short, shorter than valva, typically bent and twisted, boomerang-shaped differentiating the subtribe from Erebia Dalman, 1816 . Due to phenotypic diversity, best diagnosed by the DNA characters in the nuclear genome: hm2009277-RA.13:T2076C, hm2009277-RA.13:A130C, hm2015715-RA.2:C2137T, hm2009379-RA.4:T1915C, ahm2002906-RA.2:A128G, where the part before the first dot (e.g. hm2009277-RA) is the protein ID, next number (e.g. 13) is exon of the Heliconius melpomene genome assembly ( Davey et al. 2016), and combination like A130C means base pair C at position 130, changed from A in the ancestor. See <https://osf.io/kj4es/> for the sequences of these exons.

Genera included. The type genus, Proterebia Roos & Arnscheid, 1980 , Argestina Riley, 1923 , Loxerebia Watkins, 1925 , and Physcaeneura Wallengren, 1857 .

Parent Taxon. Tribe Satyrini Boisduval, 1833 .

Gyrocheilina Grishin, new subtribe

http://zoobank.org/ EF7C543A-88B9-4DCC-9847-139CC7763B84

Type genus. Gyrocheilus Butler, 1867 .

Definition. Previously placed in Pronophilina Reuter, 1896 ( Wahlberg 2019a), but is not monophyletic with it. Instead, it originates in early radiation of the clade that is sister to Satyrina Boisduval, 1833 (Figs. 32, 33 orange), likely prior to divergence of subtribes Ypthimina Reuter, 1896 and Erebiina Tutt, 1896, and therefore is a subtribe. Diagnosed by the combination of the following characters: most prominently, female foreleg is much reduced, with 2 tarsal subsegment, not spined; then forewings apically rounded with somewhat undulate outer margin, hindwing with strongly undulate, almost toothed outer margins; forewing discal cell short and broad: slightly longer than half of wing, width ⅓–½ of its length; forewing vein R 1 starts at distal ¼ of discal cell, R 2 very near discal cell apex (only slightly stalked), R 3 and R 4 stalked for ⅓ of their length, R 4 and R 5 stalked for about half of their length, recurrent vein in discal cell from the middle of discocellular vein for ¼ of discal cell length, discocellular vein straight between R 2 and M 1 and between M 2 and M 3 origins, but V-shaped between M 1 and M 2 origins; forewing with 4 (rarely 3) white-centered round eyespots, hindwing without such eyespots but frequently with several cream-colored ovals or crescents; palpi long and porrect, scales on the second segment long, up to 5 times the width of the segment, eyes bare, antennae shorter than half of forewing; pupa suspended, spindleshaped, smooth, head capsule apically extended similar to cremaster, forked at the tip.

Genera included. Only the type genus.

Parent Taxon. Tribe Satyrini Boisduval, 1833 .

Calistina Grishin, new subtribe

http://zoobank.org/ F1D9A7DC-F549-4EC9-8C68-D8032DC4FB95

Type genus. Calisto Hübner, [1823] .

Genera included. Only the type genus.

Parent Taxon. Tribe Satyrini Boisduval, 1833 .

Comments. The phylogenetic tree constructed from all protein-coding regions predicted to be on the Z chromosome (Fig. 33) reveals some affinity of Calisto to Euptychiina Reuter, 1896 . Although this relationship seems possible, it is not obvious from morphology, and only 76% out of 100 selections of positions from the genomic alignment support this placement (0.76 value at the node in Fig. 33). In our experience, this value is too low for confident classification. Therefore, placing Calisto in Euptychiina could be incorrect. Also, it is clearly incorrect to keep Calisto in Pronophilina, because as our tree shows, support for the Pronophilina clade that includes a diverse sample of genera is very strong, at 100% (Fig. 33 green), and Calisto is placed outside of this clade. Therefore, the solution was to propose a new substribe for the Calisto clade. Another curious observation is that the Euptychia Hübner, 1818 (type species Oreas mollina Hübner, [1813] ) clade (i.e., Euptychia sensu lato) is quite removed from the rest of Euptychiina, and their association is only weakly supported (0.56, Fig. 33). This sister to Euptychia sensu lato clade is more prominent than Euptychiina as currently defined, and may deserve subtribal status: a question that could be answered in future studies.