Tisona Higgins, 1981

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 : 43-44

Tisona Higgins, 1981 is a junior subjective synonym of Ortilia Higgins, 1981

The monotypic genus Tisona Higgins, 1981 (type and the only species Phyciodes saladillensis Giacomelli, 1911 ) was proposed on the basis of genitalic differences, most significantly "penis apex with crossing ostium-folds" ( Higgins 1981), i.e., with two twisted finger-like processes at the tip. Higgins likened Tisona to Tegosa Higgins,

1981 (type species Acraea claudina Eschscholtz, 1821 ). To better understand its phylogenetic affinities, we sequenced the only available although century-old specimen of Tisona saladillensis in the National Museum of Natural History collection (USNM) (Fig. 46). It lacks the right hindwing, and one of its labels, probably written by William Schaus, reads ||

Argentina | Giacomelli ||, suggesting that this specimen may have been part of the type series. In the genomic tree, it clusters closely and confidently with Ortilia Higgins, 1981 (type species

Papilio liriope Cramer, 1775 ), away from Tegosa (Fig. 45).

COI barcodes of O. liriope and T. saladillensis differ by 5.5%

(36 bp). This small difference strongly suggests that the two species are congeneric, and we place Tisona Higgins, 1981 as a junior subjective synonym of Ortilia Higgins, 1981 . Although the unusual genitalia of this species revealed its uniqueness despite the common Phyciodes -like wing patterns, DNA analysis was necessary to understand its origins and classification. Anthanassa Scudder, 1875 , Castilia Higgins, 1981 , Telenassa Higgins, 1981 , Dagon Higgins, 1981 , and Janatella Higgins, 1981 are subgenera of Eresia Boisduval, 1836

The most inclusive prominent clade that contains Phyciodes Hübner, [1819] (type species Papilio cocyta Cramer, 1777 ) and is consistent in genetic diversification (corrected for its higher evolutionary rate) with how other Melitaeina genera are defined is labeled " Phyciodes sensu lato " in Fig. 45. It includes the majority of Melitaeina genera and species as they are presently defined. This current classification that remains largely unchanged since the Higgins revision (1981), represents the other extreme and appears to be oversplit, because many genera defined by Higgins are too close genetically and do not stand out as prominent clades in the tree (Fig. 45, first word in species names). Most notably, Anthanassa Scudder, 1875 (type species Melitaea texana Edwards, 1863 ), Castilia Higgins, 1981 (type species Eresia castilla C. & R. Felder, 1862), Telenassa Higgins, 1981 (type species Argynnis teletusa Godart, [1824] ), Dagon Higgins, 1981 (type species Eresia catula Hopffer, 1874 ), Janatella Higgins, 1981 (type species Eresia leucodesma C. & R. Felder, 1861), and Eresia Boisduval, 1836 (type species Nereis eunice Hübner, [1807] ) cluster closely together, and the internal branches separating them are short and indistinct. While most of Higgins genera are indeed monophyletic, with some exceptions noted in the tree by colors (Fig. 45) and discussed below, the lack of their distinctiveness and low divergence is more consistent with that of subgenera. E.g., COI barcodes of the type species of Eresia and Anthanassa differ by only 6.5% (43 bp), without any correction for the generally elevated evolutionary rate of these lineages. Moreover, even Phyciodes is not particularly removed from this cluster of closely related "genera": COI barcodes of Phyciodes tharos (Drury, 1773) and Anthanassa texana (W. H. Edwards, 1863) differ by only 7.3% (48 bp). Therefore, we would generally favor broader defined genera and could propose " Phyciodes sensu lato " (Fig. 45) as a genus to combine most of these Higgins genera. However, looking for a compromise to meaningfully classify this species-rich lineage, we opt for a centrist solution. We note that Phyciodes sensu stricto and Tegosa Higgins, 1981 (type species Acraea claudina Eschscholtz, 1821 ) do form somewhat prominent clades in the tree, and we keep them as genera. As a result, clades split prior to them also keep their genus rank: Phystis Higgins, 1981 (type and the only species Eresia simois Hewitson, 1864 ), Mazia Higgins, 1981 (type and the only species Melitaea amazonica Bates, 1864 ) and Ortilia Higgins, 1981 (type species Papilio liriope Cramer, 1775 ). All others: Anthanassa Scudder, 1875 , Castilia Higgins, 1981 , Telenassa Higgins, 1981 , Dagon Higgins, 1981 , and Janatella Higgins, 1981 are placed as subgenera of Eresia Boisduval, 1836 . Finally, we note several clades in the tree that disrupt monophyly of the genera as defined by Higgins (Fig. 45: red and magenta " Ortilia " and orange " Eresia "). These clades also appear in the phylogenetic studies based on gene markers, although their exact positions relative to the other clades varied ( Wahlberg and Freitas 2007; Long et al. 2014). To remove ambiguity about their status, these clades are named as subgenera of Eresia below.