taxonID	type	description	language	source
03DC6105FFDF6969FE16A3D5FD72CB79.taxon	type_taxon	Type species. Papilio marcellus Cramer, 1777. Definition. As revealed by genome-scale phylogenetic trees (Zhang et al. 2019 d; Zhang et al. 2019 c), the type species of this new subgenus is in the same clade with the subgenus Mimoides K. Brown, 1991 (type species Papilio Fig. 2. Genus Eurytides with subgenera Boreographium ariarathes Esper, 1788) (Fig. 2). However, its subgen. n. (red), Mimoides (blue), Neographium (cyan), wing patterns and shapes are more similar to Protesilaus (purple) and Eurytides (black).	en	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): 1-55
03DC6105FFDF6969FE16A3D5FD72CB79.taxon	description	subgenus Neographium Möhn, 2002 (type species Papilio philolaus Boisduval, 1836). This phenotypic distinction and early divergence from the common ancestor with Mimoides suggest that the marcellus clade should be defined as a subgenus of its own. This new subgenus is diagnosed by male genitalia: a unique broadly rounded ear-shaped harpe with a dorsal narrow keel projecting ventrad as a tooth for half of its length, and with two prominent narrow teeth: dorsal tooth directed anteriad, ventral tooth directed caudad, as illustrated on plate 66 in Tyler et al (1994). In other subgenera, the harpe is either narrower, or the keel is broader or not projecting beyond harpe, or the teeth are smaller or directed differently.	en	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): 1-55
03DC6105FFDF6969FE16A3D5FD72CB79.taxon	etymology	Etymology. The name is a masculine noun in the nominative singular, formed from Boreo [tis] (Latin for northern) + Graphium in reference to the northernmost representatives of the " Graphium " complex of taxa in America distributed up to northern Minnesota (Lotts and Naberhaus 2021). Species included. Only the type species. Parent taxon. Genus Eurytides Hübner, [1821].	en	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): 1-55
03DC6105FFDF696AFDF7A7A3FD61CF97.taxon	type_taxon	Type species. Papilio esperanza Beutelspacher, 1975. lobe near the distal end of harpe in male genitalia, as illustrated on plate 100 in Tyler et al. (1994); and may be distinguished from other Pterourus species by its ventral forewing discal cell with 4 yellow longitudinal rays in the basal half, and from the three species in the subgenus Pterourus by its basal edge of the postdiscal band on the ventral hindwing that are closer to the outer margin than to the discal cell.	en	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): 1-55
03DC6105FFDF696AFDF7A7A3FD61CF97.taxon	etymology	Etymology. The name is a masculine noun in the nominative singular, formed as a fusion of the type species name with its genus name: Esper [anza] + [Pter] ourus. Species included. Only the type species. Parent taxon. Genus Pterourus Scopoli, 1777.	en	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): 1-55
03DC6105FFDC696AFDF0A3C6FC80CB91.taxon	type_taxon	Type species. Papilio hyppason Cramer, 1775. Definition. Previously, we noted this unnamed subgenus in the COI barcode dendrogram (Shiraiwa et al. 2014). Here, its definition is formalized. While the COI barcodes demonstrate its distinction from other subgenera of Heraclides Hübner, [1819] (type species Papilio thoas Linnaeus, 1771), its unexpected sister Fig. 4. Genus Heraclides with subgenera Hyppasonia subgen. n. (red), Heraclides (blue), Calaides (cyan) and Priamides (purple). relationship with the nominal subgenus	en	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): 1-55
03DC6105FFDC696AFDF0A3C6FC80CB91.taxon	description	Heraclides — instead of with Priamides Hübner, [1819] (type species Priamides hipponous Hübner, [1819], which is a junior objective synonym of Heraclides anchisiades (Esper, 1788 )) as hinted by similarities in wing shape and patterns — was discovered by Lewis et. al. (2015). Our genomic tree confirms this placement, and at the same time reveals prominent genetic differentiation from the nominal subgenus (Fig. 4). Curiously, the long branch in the tree that defines this subgenus suggests accelerated evolution that may explain its phenotypic differences from the nominal subgenus and likely mimetic wing patterns. This new subgenus is distinguished from others by a bilobed, crab-claw harpe with strongly unequal lobes: one lobe broad, rounded and serrated distad, and the other lobe shorter, narrow, tooth-like (for illustration see plate 83 in Tyler et al. (1994 )). In other species with a bilobed harpe, the two lobes are nearly equal or at least the larger lobe is narrower and prominently constricted before the serrated end.	en	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): 1-55
03DC6105FFDC696AFDF0A3C6FC80CB91.taxon	etymology	Etymology. The name is a feminine noun in the nominative singular, formed from the type species name. Species included. Only the type species. Parent taxon. Genus Heraclides Hübner, [1819].	en	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): 1-55
03DC6105FFDD696BFE44A64BFA17C86E.taxon	description	Compared to Pyrisitia dina helios (M. Bates, 1934) (type locality Bahamas) the Fst / Gmin statistics are 0.46 / 0.03 and their COI barcodes are 2.7 % (18 bp) different. Therefore, we propose that Pyrisitia Fig. 6. Pyrisitia westwoodii (red), and dina (blue) and parvumbra (Jamaica). westwoodii (Boisduval, 1836), reinstated status is a species-level taxon and employ Pyrisitia westwoodii gabriela Le Crom & Llorente, 2004 comb. nov. We also confirm Pyrisitia parvumbra (Kaye, 1925) as a species (Fig. 6; 2.6 %, 17 bp barcode difference).	en	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): 1-55
03DC6105FFD26965FE81A14AFA2ECCBE.taxon	description	Reliquia Ackery, 1975 is a junior subjective synonym of Pontia [Fabricius], 1807 Baltia Moore, 1878 and Pontieuchloia Verity, 1929 are subgenera of Pontia [Fabricius], 1807 A small genus Baltia Moore, 1878 (type species Mesapia shawii Bates, 1873) originates within Pontia [Fabricius], 1807 (type species Papilio daplidice Linnaeus, 1758), rendering Pontia paraphyletic (Fig. 9). Baltia is a confident but comparatively distant sister to the clade of two closely related species Pontia chloridice Hübner, [1813] (the type species of Pontieuchloia Verity, 1929, currently a junior subjective synonym of Pontia) and Pontia beckerii (W. H. Edwards, 1871). E. g., COI barcodes of B. shawii and P. chloridice differ by 8.5 % (51 bp). To restore monophyly of Pontia and, at the same time, keep the relative distinction of Baltia, we propose that Baltia Moore, 1878 is a subgenus of Pontia [Fabricius], 1807. If Baltia is a subgenus and not a synonym, clades comparable to it in prominence should be defined as subgenera. In addition to the nominal subgenus (Fig. 9 blue and pink), of which Synchloe Hübner, 1818 (type species Papilio callidice Hübner, [1800]) is kept as a junior subjective synonym, we propose that Pontieuchloia Verity, 1929 is a valid subgenus (Fig. 9 cyan). For these subgenera to be monophyletic, a new subgenus is proposed next for the clade with Pontia sisymbrii (Boisduval, 1852) (Fig. 9 magenta).	en	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): 1-55
03DC6105FFD36965FDC0A0E0FD15C888.taxon	type_taxon	Type species. Pieris sisymbrii Boisduval, 1852. Definition. This new subgenus differs from other subgenera of Pontia by the following combination of characters: forewing vein R 3 longer than in other subgenera, about half of vein R 4 + 5 length; androconia present in the forewing discal cell spot in males, this spot is narrower than in other subgenera, with a notch on the outer edge (smoothly curved of straight in Baltia) and without a line of white scales along the discal cross-vein that is curved less strongly than in most other subgenera towards the wing base; dorsal hindwing without prominent bar at the end of discal cell; aedeagus shorter and relatively broader than in other subgenera, prominently curved at phallobase; hindwing below with gray or brown (not green or yellow) scaling along yellowish veins; full-grown caterpillar with orange-yellow framed with black rings on grayish segments; univoltine in spring. See Chang (1963) for elaboration on and illustrations of some of these characters as they are given for P. sisymbrii.	en	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): 1-55
03DC6105FFD36965FDC0A0E0FD15C888.taxon	etymology	Etymology. The name is a feminine noun in the nominative singular, formed from the type species name. Species included. Only the type species. Parent taxon. Genus Pontia [Fabricius], 1807.	en	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): 1-55
03DC6105FFD36965FDC0A0E0FD15C888.taxon	discussion	Comments. The genomic tree reveals markedly uneven rates of evolution within Pontia (Fig. 9): the nominotypical subgenus evolves about 2 times faster than other subgenera. This observation combined with rather substantial genetic differentiation among Pontia, including the COI barcode, which in P. (Sisymbria) sisymbrii and P. (Pontia) daplidice differs by 8.7 % (57 bp), and close similarity in phenotypes of Pontia species creates a unique situation. On the one hand, Pontia (including Baltia) is a morphologically compact genus. On the other hand, strong genetic diversification behind this apparent phenotypic similarity may suggest elevating subgenera of Pontia to genera (which will return Baltia to the genus status), a step that we refrain from.	en	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): 1-55
03DC6105FFD36967FF28A4D5FA46CFD8.taxon	description	http: // zoobank. org / B 7717 ECE-C 015 - 48 AA-A 57 D-B 7 A 6 A 3 CCE 4 F 6	en	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): 1-55
03DC6105FFD36967FF28A4D5FA46CFD8.taxon	type_taxon	Type genus. Calopieris Aurivillius, 1898. Definition. In the genomic tree, this taxon is confidently placed as sister to the tribe Leptosiaini Braby, 2014 (Fig. 10, a monotypic tribe consisting of Leptosia Hübner, 1818) and shares veins M 1 and M 2 being connate with the hindwing discal cell, but genetically and phenotypically distant from it otherwise. Most notably, the wings are not rounded as in Leptosia and venation differs: forewing veins R 3 and R 4 + 5 stalked for less than half of their lengths and M 1 stalked with their stalk, resembling Coliadinae Swainson, 1821 (e. g., Phoebis Hübner, [1819]). In general appearance reminds more of Colotis, where it was formerly included as a subgenus (Klots 1933), but is distinguished from it by very short and slender palpi that do not protrude beyond the front of the head and are not visible from above. Antennae are short (about half of forewing discal cell length) with large and flattened clubs. The combination of the abovementioned characters of palpi, antennae and wing venation uniquely defines this new tribe. Genera included. Only the type genus.	en	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): 1-55
03DC6105FFD36967FF28A4D5FA46CFD8.taxon	description	Parent Taxon. Subfamily Pierinae Swainson, 1820.	en	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): 1-55
03DC6105FFD36967FF28A4D5FA46CFD8.taxon	discussion	Comments. Although grammatically correct formation of this tribe name calls for insertion of " id " before " ini ", these letters have been elided from the stem to agree with Pierini, under Art. 29.3.1.1 (ICZN 1999). Phrissura Butler, 1870 is a subgenus of Appias Hübner, [1819] Saletara Distant, 1885 is a subgenus of Catophaga Hübner, 1819 Frequently treated as a subgenus of Appias Hübner, [1819] (type species Papilio zelmira Stoll, 1780, currently a subspecies of Papilio libythea Fabricius, 1775), Catophaga Hübner, [1819] (type species Papilio paulina Cramer, [1777]) is not monophyletic with it and instead, together with Saletara Distant, 1885 (type species Pieris nathalia C. & R. Felder, 1862, currently a subspecies of Papilio liberia Cramer, [1779]), is sister to Aoa de Nicéville, 1898 (type and the only species Pieris affinis Vollenhoven, 1865) (Fig. 10). Therefore, in agreement with Wahlberg et al. (2014), we treat Catophaga as a valid genus distinct from Appias. Our genomic tree reveals close clustering of Saletara (Fig. 10 red) with Catophaga (Fig. 10 blue) at the level consistent with congeners, hence we propose that Saletara is a subgenus of Catophaga.	en	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): 1-55
03DC6105FFD16967FF3AA005FDD6CDB0.taxon	description	Papilio crataegi Linnaeus, 1758) rendering it paraphyletic (Fig. 11). Due to the close relationship of these taxa, it is not desirable to restore monophyly by breaking Aporia into at least 3 genera. Therefore, agreeing with previous studies (Ding and Zhang 2016; Todisco et al. 2020), genomic-scale phylogeny provides the ultimate evidence that Mesapia Gray, 1856 is a junior subjective synonym of Aporia Hübner, [1819].	en	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): 1-55
03DC6105FFD66960FDCDA002FA98C9BC.taxon	description	Appearance could be misleading, especially if unusual. Despite unique wing patterns, we find from the genomic analysis that Sandia Clench & P. Ehrlich, 1960 (type and the only species Callophrys (Sandia) mcfarlandi P. Ehrlich & Clench, 1960) originates within Deciduphagus K. Johnson, 1992 (type species Thecla augustinus Westwood, 1852), and is sister to Callophrys fotis (Strecker, [1878]) (Fig. 13). Because Deciduphagus is currently a junior subjective synonym of Incisalia Scudder, 1872 (type species Lycus niphon Hübner, [1819]) (Pelham 2008; Pelham 2020) due to evolutionary closeness, and because Sandia renders both Deciduphagus and Incisalia paraphyletic, we propose that Sandia is a junior subjective synonym of Incisalia. Furthermore, uniquely patterned Xamia Clench, 1961 (type species Thecla xami Reakirt, [1867]) also originates within Deciduphagus and is sister to the Deciduphagus core group that contains the type species. Although Xamia is more distant from other congeners than Sandia, it still falls within the prominent Incisalia clade. Thus leaving it as a valid subgenus would require a new name for at least Callophrys polios (Cook & F. Watson, 1907), which is sister to the clade Fig. 13. Callophrys subgenera: subgen. Greenie n. (red), Cisincisalia (pink), consisting of Xamia and the core Callophrys (blue), Mitoura (olive), and Incisalia (green) with its synonyms Deciduphagus species, but is not Deciduphagus (cyan), Xamia (orange), and Sandia (magenta). prominently distinct from its relatives. Therefore, we propose to treat Xamia as a junior subjective synonym of Incisalia. Finally, we note that Deciduphagus as originally defined (Fig. 13 cyan) (Johnson 1992) is also paraphyletic with respect to Incisalia (Fig. 13 green), and Callophrys henrici (Grote & Robinson, 1867) with Callophrys irus (Godart, [1824]) should be attributed to Incisalia sensu stricto instead of to Deciduphagus. As a result of this analysis (Fig. 13), we conclude that the genus Callophrys diversified into 5 prominent clades that correspond to the level of subgenus: Callophrys, Incisalia, Cisincisalia K. Johnson, 1992 (type species Cisincisalia moecki K. Johnson, 1992, a junior subjective synonym of Callophrys guatemalena Clench, 1981), Mitoura Scudder, 1872 (type species Thecla smilacis Boisduval & Le Conte, [1835]) and the fifth subgenus that includes Nearctic species superficially similar to Palearctic Callophrys, but instead is a confidently supported sister to Incisalia (Fig. 13 red) (ten Hagen and Miller 2010). This subgenus does not have a name available for it and therefore is new.	en	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): 1-55
03DC6105FFD76961FEA9A738FA5AC905.taxon	description	Fig. 14. Nemeobiini genera: Hamearis (black), Styx by proposing that Corrachia Schaus, 1913 is a (red and green), Polycaena (blue and magenta), subgenus of Styx Staudinger, 1876. Considering these Dodona (olive), new genus (Espeland et al. 2015), two close relatives to be congeneric is more revealing Stiboges (cyan) and Zemeros (purple). about their evolutionary relationship than keeping them in two not prominently distinct monotypic genera.	en	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): 1-55
03DC6105FFD76961FDD3A2B2FD7ACDBD.taxon	type_taxon	Type species. Thecla sheridonii [sic] Edwards, 1877, presently Callophrys sheridanii (Edwards, 1877). Definition. Previously placed in the subgenus Callophrys Billberg, 1820 (type species Papilio rubi Linnaeus, 1758) but is not monophyletic with it, forming a prominent lineage of its own within the genus Callophrys (Fig. 13). In appearance, remarkably similar to some species (including the type) from the subgenus Callophrys in: wings rounded without tails, slightly lobed hindwings at anal angle, and green color of ventral surface with postdiscal variously complete to absent row of white markings. Distinguished from Callophrys by hindwing tornal area: less developed lobe and less crenulate outer margin, and generally smaller and rounder forewing androconial patch. Diagnosed among the genus Callophrys by a combination of the following characters in the nuclear genome: cce 2400.8.3: T 366 C, cce 8426.17.3: C 4290 T, cce 10587.6.6: G 181 C, cce 925.9.4: C 351 T, and cce 2041.25.2: T 1572 A; and the COI barcode: 479 C (not T) and 610 C (not T), differing from the subgenus Callophrys in 202 T (not A), 512 G (not T), and 556 T (not A). See <https: // osf. io / kj 4 es /> for the sequences with these characters.	en	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): 1-55
03DC6105FFD76961FDD3A2B2FD7ACDBD.taxon	etymology	Etymology. The name is a feminine noun in the nominative singular, Latinized verbatim from the affectionate English name of this butterfly group. Species included. The type species, Thecla viridis W. H. Edwards, 1862, Thecla dumetorum Boisduval, 1852, and Thecla affinis W. H. Edwards, 1862. Parent taxon. Genus Callophrys Billberg, 1820.	en	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): 1-55
03DC6105FFD56963FDA6A33FFC9DCB9B.taxon	type_taxon	Type species. Eurygona pelor Hewitson, [1853]. Definition. Currently within Euselasia Hübner, [1819] (type species Euselasia gelaena Hübner, [1819], which is Papilio gelon Stoll, 1787) but is not monophyletic with it (Fig. 15 magenta). Instead, an independent prominent lineage originating in the early radiation of Euselasiini Kirby, 1871 and a poorly supported sister to the clade that includes Methone Doubleday, 1847 (type species Papilio cecilia Cramer, 1777) (Fig. 15 brown, gray and red) and Erythia Hübner, [1819] (type species Papilio labdacus Stoll, 1780) (Fig. 15 olive). Therefore, it constitutes a new genus. Union of Stichel (1928) groups Peloriformes, Eusepiformes, Melaphaeoformes, Argenteoformes and Eubuliformes (in part). Characterized by wings rounder than in relatives, hindwing with undulate outer margin and checkered fringes, ventrally with postdiscal reddish line and a row of marginal black dots framed with white, reddish, or both; or nearly immaculate wings below, silvery to golden, may be with marginal black dots on hindwing and brown postdiscal line; or with 3 – 5 brown narrow bands over broader than bands pale background, lacking isolated large eyespots or dotted discal pattern, or with a row of small eyespots along hindwing margin. The following combination of nuclear genome characters is diagnostic: cne 2559.1.3: T 87 C, cne 3355. 8.1: T 362 C, cne 1314.4.1: A 351 G, cne 2022.5.2: C 448 A, and cne 599.10.1: T 5841 A.	en	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): 1-55
03DC6105FFD56963FDA6A33FFC9DCB9B.taxon	etymology	Etymology. The name is a feminine noun in the nominative singular formed as a fusion of the type species name and its former genus name: Pelo [r] + [Euse] lasia to keep the word at 9 letters, which is the number of letters in Euselasia and the number of genera Euselasia is being split into. Species included. The type species, Eurygona amphidecta Godman & Salvin, 1878, Eurygona argentea Hewitson, 1871, Eurygona artos Herrich-Schäffer, [1853], Eurygona aurantia Butler & Druce, 1872, Eurygona bettina Hewitson, 1869, Eurygona candaria Druce, 1904, Eurygona cataleuca R. Felder, 1869, Eurygona chrysippe Bates, 1866, Eurygona euboea Hewitson, [1853], Eurygona eumedia Hewitson, 1853, Eurygona eumenes Hewitson, 1853, Euselasia eupatra Seitz, 1916, Eurygona eusepus Hewitson, 1853, Eurygona fervida Butler, 1874, Euselasia hahneli Staudinger, [1887], Euselasia ignitus Stichel, 1924, Eurygona mazaca Hewitson, 1860, Erythia melaphaea Hübner, 1823, Eurygona mirania Bates, 1868, Euselasia misteriosa Salazar & J. Vargas, 2019, Euselasia nytua J. Hall & Willmott, 2009, Euselasia pellonia Stichel, 1919, Euselasia rubrocilia Lathy, 1926, and Euselasia seitzi Lathy, 1926. Parent taxon. Subfamily Euselasiinae Kirby, 1871.	en	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): 1-55
03DC6105FFD5697CFDACA7C2FC9DCD22.taxon	type_taxon	Type species. Eurygona mys Herrich-Schäffer, [1853]. Definition. Currently within Euselasia Hübner, [1819] (type species Euselasia gelaena Hübner, [1819], which is Papilio gelon Stoll, 1787) but is not monophyletic with it (Fig. 15 green). Instead, an independent prominent lineage in the same clade with and of the same rank as Methone Doubleday, 1847 (type species Papilio cecilia Cramer, 1777) (Fig. 15 brown, gray and red), thus is a new genus. Largely, the Hygeniiformes group of Stichel (1928), sharing its diagnostic characters. Hindwing rounded, below with a kinked-L-shaped central reddish band and an eyespot in the middle by the margin, but without well-developed marginal longitudinal dashes; forewing without eyespots and with 1 - 2 narrow reddish bands. The following combination of nuclear genome characters is diagnostic: cne 23605.2.5: C 468 T, cne 13338.5.3: A 334 C, cne 123.2.3: C 1009 T, cne 18035.2.1: C 172 A, and cne 81.14.5: A 756 G.	en	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): 1-55
03DC6105FFD5697CFDACA7C2FC9DCD22.taxon	etymology	Etymology. The name is a feminine noun in the nominative singular formed as a fusion of the type species name and its former genus name: Mys + [Eus] elasia to keep the word at 9 letters, which is the number of letters in Euselasia and the number of genera Euselasia is being split into. Species included. The type species, Eurygona alcmena Druce, 1878, Eurygona athena Hewitson, 1869, Eurygona cafusa Bates, 1868, Euselasia crinon Stichel, 1919, Euselasia cucuta (Schaus, 1902), Euselasia cyanofusa J. Hall & Willmott, 1998, Euselasia eberti Callaghan, 1999, Euselasia ella Seitz, 1916, Eurygona eulione Hewitson, 1856, Euselasia eustola Stichel, 1919, Euselasia gradata Stichel, 1927, Eurygona hieronymi Salvin & Godman, 1868, Papilio hygenius Stoll, 1787, Euselasia illarina J. Hall, Willmott & R. Busby, 1998, Eurygona inconspicua Godman & Salvin, 1878, Euselasia janigena Stichel, 1919, Euselasia jigginsi J. Hall & Willmott, 1998, Eurygona leucon Schaus, 1913, Euselasia mapatayna J. Hall & Willmott, 1998, Euselasia marica Stichel, 1919, Eurygona mystica Schaus, 1913, Euselasia nauca J. Hall & Willmott, 1998, Euselasia pance Callaghan, 1999, Eurygona procula Godman & Salvin, 1885, Euselasia pseudomys Callaghan, 1999, Euselasia pullata Stichel, 1927, Eurygona pusilla R. Felder, 1869, Euselasia rhodon Seitz, 1913, and Eurygona sergia Godman & Salvin, 1885. Parent taxon. Subfamily Euselasiinae Kirby, 1871.	en	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): 1-55
03DC6105FFCA697CFDA2A10CFC9DC816.taxon	type_taxon	Type species. Eurygona euryone Hewitson, 1856. Definition. Currently within Euselasia Hübner, [1819] (type species Euselasia gelaena Hübner, [1819], which is Papilio gelon Stoll, 1787) but is not monophyletic with it (Fig. 15 pink). Instead, an independent prominent lineage in the same clade with and of the same rank as Methone Doubleday, 1847 (type species Papilio cecilia Cramer, 1777) (Fig. 15 brown, gray and red), hence is a new genus. It is diagnosed by its distal ⅖ of hindwing dusted pale-yellow below, large round marginal eyespot in the middle of it, and either large eyespot in the middle by forewing margin below or diagonal orange patch on forewing above. The hindwing tornus is pale above in some species. The following combination of nuclear genomic characters is diagnostic: cne 9878.8.1: C 146 A, cne 178.3.20: C 595 T, cne 178.3.20: T 596 C, cne 7676.26.2: T 31 C, and cne 5931.2.1: A 478 G.	en	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): 1-55
03DC6105FFCA697CFDA2A10CFC9DC816.taxon	etymology	Etymology. The name is a feminine noun in the nominative singular formed as a fusion of the type species name and its former genus name: Eury [one] + [Euse] lasia to keep the word at 9 letters, which is the number of letters in Euselasia and the number of genera Euselasia is being split into. Species included. The type species, Eurygona effima Hewitson, 1869, Euselasia thusnelda Möschler, 1883, and Eurygona eunaeus Hewitson, 1855. Parent taxon. Subfamily Euselasiinae Kirby, 1871.	en	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): 1-55
03DC6105FFCA697DFDABA458FC9DCCB4.taxon	type_taxon	Type species. Euselasia albomaculiga Callaghan, 1999. Definition. The genus is sister to Euselasia Hübner, [1819] (Euselasia gelaena Hübner, [1819], which is Papilio gelon Stoll, 1787) and is prominently distinct from it genetically (Fig. 15 orange). Therefore, it constitutes a new genus. Distinguished from its relatives by the characters given on pages 1047 – 1048 and illustrated in Figs. 3 – 6, 46 – 47 for Euselasia albomaculiga by Callaghan (1999). In brief, its wings are rounded, without blue scaling, its forewings are with a large pale spot, and its ventral hindwing submarginal black spots are surrounded by grayish-white (not at the end of orange rays, except the central large one that is framed by yellow basad), and have yellow streaks along the veins not between them; valvae narrowing to a point, not bilobed, as long as tegumen with uncus, slightly wider than aedeagus, vinculum angled in lateral view, with spurs in the middle directed caudad. Additionally, the following combination of nuclear genomic characters is diagnostic: cne 2298.2.2: A 2104 A (not C), cne 5129.1.5: A 833 A (not G), cne 2685.14.3: A 934 A (not T), cne 1095.7.22: G 1463 G (not A), cne 4870.1.18: T 552 T (not A), cne 2885.9.10: A 4270 C, cne 1547.14.4: A 2756 G, cne 1696.1.1: A 2529 T, cne 7231.10.9: T 567 C, and cne 1036.6.11: G 3200 C.	en	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): 1-55
03DC6105FFCA697DFDABA458FC9DCCB4.taxon	etymology	Etymology. The name is a feminine noun in the nominative singular formed as a fusion of the type species name and its former genus name: [albo] Macu [liga] + [Euse] lasia to keep the word at 9 letters, which is the number of letters in Euselasia and the number of genera Euselasia is being split into. Species included. Only the type species. Parent taxon. Subfamily Euselasiinae Kirby, 1871.	en	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): 1-55
03DC6105FFCB697DFDACA0FAFC9DCB78.taxon	type_taxon	Type species. Eurygona eugeon Hewitson, 1856. Definition. This taxon is sister to the clade formed by Euselasia Hübner, [1819] (Euselasia gelaena Hübner, [1819], which is Papilio gelon Stoll, 1787) and Maculasia gen. n. and is prominently distinct from it genetically (Fig. 15 purple); hence it is a new genus. It is distinguished from its relatives by its inverted-drop shaped hindwing with extended anal lobe, as well as by its solid-brown color above and lack of eyespots below, fringes dark, legs yellow; and the following combination of nuclear genomic characters: cne 1999.2.1: C 203 A, cne 1015.3.2: T 555 C, cne 2803.19.1: G 88 A, cne 5471.1.1: T 261 C, and cne 703.2.8: A 1414 C.	en	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): 1-55
03DC6105FFCB697DFDACA0FAFC9DCB78.taxon	etymology	Etymology. The name is a feminine noun in the nominative singular formed as a fusion of the type species name and its former genus name: Euge [on] + [Euse] lasia to keep the word at 9 letters, which is the number of letters in Euselasia and the number of genera Euselasia is being split into. Species included. The type species and Euselasia brevicauda Lathy, 1926. Parent taxon. Subfamily Euselasiinae Kirby, 1871.	en	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): 1-55
03DC6105FFC9697FFDA8A2ADFC8FCDB3.taxon	type_taxon	Type species. Papilio eumene Cramer, 1776. Definition. Species in this clade are currently in Mesosemia Hübner, [1819] (type species Mesosemia phicoclessa Hübner, [1819], which is Papilio philocles Linnaeus, 1758), but are quite distant from it falling much outside of the Mesosemia radiation and originating during the earlier round of radiation (Fig. 16). Therefore this clade is a taxon of the same rank, a genus. This new genus differs from Mesosemia by the following combination of characters: hindwing not lobed in the middle, each wing with 2 broad and straight parallel bands: discal and postdiscal, bands do not encircle forewing eyespot as in most Mesosemia, but discal forewing band bends distad at costa, no striations, discal cell eyespots developed ventrally on both wings, elongated along the cell, with 3 white spots inside (sometimes 2 on hindwing). Furthermore, the following combination of nuclear genomic characters is diagnostic: cne 7048.1.3: T 2251 C, cne 3658.2.1: A 638 T, cne 2957.11.4: A 952 G, cne 3658.2.1: A 608 C, and cne 3461.2.10: C 1462 A.	en	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): 1-55
03DC6105FFC9697FFDA8A2ADFC8FCDB3.taxon	etymology	Etymology. The name is a feminine noun in the nominative singular formed by replacing prefix " Meso- " with " Ecto- " in the name of the former genus for these species. Species included. The type species, Mesosemia decolorata Lathy, 1932, Mesosemia erinnya Stichel, 1910, and Mesosemia steli Hewitson, 1858. Parent taxon. Subtribe Mesosemiina Bates, 1859.	en	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): 1-55
03DC6105FFC9697FFDD6A1FAFC8FC85D.taxon	type_taxon	Type species. Papilio ulrica Cramer, 1777. Definition. Species in this clade are currently in Mesosemia Hübner, [1819] (type species Mesosemia phicoclessa Hübner, [1819], which is Papilio philocles Linnaeus, 1758), but are quite distant from it falling much outside of the Mesosemia radiation and originating during the earlier round of radiation (Fig. 16). Therefore this clade is a taxon of the same rank, a genus. This new genus differs from both Mesosemia and Ectosemia gen. n. by the following combination of characters: hindwing not lobed in the middle, forewing eyespot not elongated along discal cell, discal band (if developed) narrow, bands do not encircle forewing eyespot, or bands diffuse and poorly formed. Furthermore, the following combination of nuclear genomic characters is diagnostic: cne 703.2.8: T 903 C, cne 1411.6.4: T 1025 A, cne 2651.14.5: A 4602 G, cne 12205.6.2: G 835 A, and cne 8028.2.1: T 1787 A.	en	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): 1-55
03DC6105FFC9697FFDD6A1FAFC8FC85D.taxon	etymology	Etymology. The name is a feminine noun in the nominative singular formed by replacing prefix " Meso- " with " Endo- " in the name of the former genus for these species. Species included. The type species and Mesosemia macella Hewitson, 1859. Parent taxon. Subtribe Mesosemiina Bates, 1859.	en	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): 1-55
03DC6105FFC96978FDEFA480FABCCE9B.taxon	description	Reassessment of Voltinia Stichel, 1910 and Napaea Hübner, [1819]	en	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): 1-55
03DC6105FFCE6978FEADA626FE93CB54.taxon	description	A genus comprised of two species, Voltinia Stichel, 1910 (type species Esthemopsis (?) radiata Godman & Salvin, 1886) (Fig. 17 gray) is a close sister to Cremna Doubleday, 1847 (type species Papilio actoris Cramer, 1776), which also consists of only two species: the type and Cremna heteroea Bates, 1867 (Fig. 17 purple); and Ionotus Hall, 2005 (type and the only species Hamanumida alector Geyer, 1837) (Fig. 17 olive) is sister to them combined. Hence, we reinstate Cremna as a valid genus (not a synonym of Napaea), and due to genetic similarities place Ionotus Hall, 2005 and Voltinia Stichel, 1910 as its subgenera.	en	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): 1-55
03DC6105FFCC697AFDBAA60BFD76C983.taxon	type_taxon	Type species. Symmachia virgatula Stichel, 1910. Definition. This group of species is currently placed in Symmachia Hübner, [1819] (type species Symmachia probetrix Hübner, [1819], which is Papilio probetor Stoll, 1782) but is not monophyletic with it (Fig. 18). It originates early in the radiation of the Symmachiini core group, a possible, but weakly supported sister to Phaenochitonia Stichel, 1910 (type species Papilio cingulus Stoll, 1790) sensu lato (see below), and therefore is a genus. Similar to Symmachia and Tigria gen. n. and is distinguished from them by the following combination of characters (at least in males): forewing costa concave in the middle, as in most Symmachia but different from Tigria gen. n., wings red-orange to yellow, bordered and partly striped or spotted at least along forewing costa with black or dark-brown, without white spots by the forewing apex. The following combination of nuclear genome characters is diagnostic: cne 4291.7.6: A 1077 G, cne 4291.7.6: A 1064 G, cne 4291.7.6: A 1051 G, cne 3461.2.5: A 1310 G, and cne 3461.1.15: A 3342 C.	en	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): 1-55
03DC6105FFCC697AFDBAA60BFD76C983.taxon	etymology	Etymology. The name is a feminine noun in the nominative singular, formed from the beginning of the former genus name of these species, prefixing it with " a " for " not ", because these species cannot possibly belong to Symmachia: A + symma [chia]. Species included. The type species, Synapta arion C. & R. Felder, 1865, Symmachia giffordi P. Jauffret & J. Jauffret, 2010, Symmachia hippodice Godman, 1903, Symmachia pardalis Hewitson, 1867, Symmachia satana J. Hall & Harvey, 2007, Symmachia stigmosissima Stichel, 1910, and Symmachia virgaurea Stichel, 1910. Parent taxon. Tribe Symmachiini Reuter, 1896.	en	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): 1-55
03DC6105FFCC697AFD89A33EFD76CA21.taxon	type_taxon	Type species. Mesene xypete Hewitson, 1870. Definition. A sister clade to other Symmachia Hübner, [1819] (type species Symmachia probetrix Hübner, [1819], which is Papilio probetor Stoll, 1782), but prominently distinct from it, more so than Symmachia sensu stricto species are from each other (Fig. 18 blue), and genetically distant from them at a level where other Symmachiini genera are defined (Fig. 18), is therefore a genus. It is similar to Symmachia and distinguished from it by the following combination of characters (at least in males): forewing costa rather straight, not concave, apex produced, hindwing typically with angular tornus; antennae long, about ¾ of forewing length; eyes bare; wings red-orange, bordered and partly striped at least along forewing costa with black or dark-brown, dark areas could take half of wings, no pale spot mid-costa. The following combination of nuclear genome characters is diagnostic: cne 1935.6.1: A 2889 G, cne 3461.1.14: G 922 C, cne 3437.1.9: A 1260 G, cne 3461.1.14: A 846 G, and cne 2170.2.1: T 2751 C.	en	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): 1-55
03DC6105FFCC697AFD89A33EFD76CA21.taxon	etymology	Etymology. The name is a feminine noun in the nominative singular given for the red-orange tiger-striped appearance of these species. Species included. The type species, Polystichtis rubrica Stichel, 1929, Cricosoma phaedra Bates, 1868 and Metacharis elinas Rebillard, 1958. Parent taxon. Tribe Symmachiini Reuter, 1896.	en	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): 1-55
03DC6105FFCD697BFE2EA389FBA6CCD6.taxon	description	(type locality Brazil: Para) is not monophyletic with it, and instead is sister to Mesene argentea Bates, 1866, the type species of Chimastrum Godman & Salvin, 1886 that originates within Mesene Doubleday, 1847 (type species Papilio phareus Cramer, 1777) (Fig. 18, sequenced specimen in Fig. 19), implying Mesene aeolia (Bates, 1868) comb. nov.	en	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): 1-55
03DC6105FFCD697BFEB5A473FE4DC982.taxon	description	1910 (type species Pterographium aphaniodes Stichel, 1910, which is Panara sicora Hewitson, 1875) sensu lato as it is defined above (Fig. 18, sequenced specimen shown in Fig. 20), hence Pterographium hypochloris (Bates, 1868) comb. nov.	en	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): 1-55
03DC6105FFC26974FEDAA7CCFD71C979.taxon	description	Echenais Hübner, [1819] (type species Lemonias alphaea Hübner, [1808], which is Papilio thelephus Cramer, 1775) is genetically close to Imelda Hewitson, 1870 (type species Imelda glaucosmia Hewitson, 1870, which is a subspecies of Nymphidium mycea Hewitson, 1865) (Fig. 23). E. g., COI Fig. 23. Echenais (blue) subgenus Imelda (red). barcodes of their type species differ by 8.4 % (55 bp). Therefore we propose treating Imelda Hewitson, 1870 as a subgenus of Echenais Hübner, [1819].	en	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): 1-55
03DC6105FFC26974FE8DA3D9FE3ACBE1.taxon	description	1777), Hesperia florus Fabricius, 1793 (type locality " Indiis ", likely SE Brazil) is not monophyletic with it and originates within Phaenochitonia Stichel, 1910 (type species Papilio cingulus Stoll, 1790) (Fig. 18 yellow, highlighted in violet, sequenced specimen shown in Fig. 21), implying Phaenochitonia florus (Fabricius, 1793) comb. nov. Distinctness of this species from Mesene came as a surprise considering how well it was fitting in that genus on the basis of wing colors, patterns and shape, even reminding of Mesene type species. Reassessment of Ourocnemis Bethune-Baker, 1887 and Anteros Hübner, [1819] The tree of Helicopini Stichel, 1928 revealed a distribution of species between the two genera Ourocnemis Bethune-Baker, 1887 (type species Anteros axiochus Hewitson, 1867) and Anteros Hübner, [1819] (type species Papilio formosus Cramer, 1777) not as presently attributed (Fig. 22). A number of species placed in Anteros were in the same clade with Ourocnemis, confirming the suspicion of Hall (1998) that Anteros may be paraphyletic. Using this phylogenetic result and phenotypic similarities (cream vs. dark background of wings below) for species with missing DNA data, we transfer four species from	en	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): 1-55
03DC6105FFC36975FF52A25EFDDECC31.taxon	description	(type species Monethe johnstoni Dannatt, 1904) originates deep within Baeotis Hübner, [1819] (type species Baeotis hisbaena Hübner, [1819] which is	en	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): 1-55
03DC6105FFC36975FF70A07DFD34C9B9.taxon	type_taxon	Type species. Charis argyrea Bates, 1868. Definition. Currently within Detritivora Hall & Harvey, 2002 (type species Charis matic Harvey & Hall, 2002), but is not monophyletic with it, and instead is sister to the clade that includes Detritivora with a number of other genera such as Charis Hübner, [1819] (type species Charis ania Hübner, Fig. 26. Putridivora gen. n. (green), Detritivora [1819], which is Papilio anius Cramer, 1776) and (brown), Chadia (pink), Charis (olive), Inkana Calephelis Grote & Robinson, 1869 (type species Erycina gen. n. (red), and Lasaia (blue). virginiensis Guérin-Méneville, [1832]) (Fig. 26), hence a new genus. Similar to Detritivora as described by Hall and Harvey (2002), and distinguished from it by 3 (not 4) dark marks in discal cell and signa markedly elongate along corpus bursae wall, as described by Hall and Harvey (2001), see their Fig. 6 a.	en	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): 1-55
03DC6105FFC36975FF70A07DFD34C9B9.taxon	etymology	Etymology. The name is a feminine noun in the nominative singular formed by replacing prefix " Detriti- " with " Putridi- " in the name of the former genus for these species, but keeping the meaning. Species included. The type species and Charis smalli Hall & Harvey, 2001. Parent taxon. Tribe Riodinini Grote, 1895.	en	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): 1-55
03DC6105FFC06976FDB0A25EFD34CCEF.taxon	type_taxon	Type species. Charis cadytis Hewitson, 1866. Definition. Currently one of the two species of Charis Hübner, [1819] (type species Charis ania Hübner, [1819], which is Papilio anius Cramer, 1776), but is not monophyletic with the second (and the type) species, and instead sister to the clade that includes a number of other genera such as Charis and Calephelis Grote & Robinson, 1869 (type species Erycina virginiensis Guérin-Méneville, [1832]) (Fig. 26). Therefore, it constitutes a new genus. Distinguished from its relatives by atypical for the group more elongated wings (without pointed apex), orange band at forewing apex, prominent silver wing margins in males below; exceptionally curved, earlobe-shaped aedeagus and very broad valvae, as broad as long.	en	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): 1-55
03DC6105FFC06976FDB0A25EFD34CCEF.taxon	etymology	Etymology. The name is a feminine noun in the nominative singular formed from the beginning of the type species name with insertion of h for Charis. Species included. Only the type species. Parent taxon. Tribe Riodinini Grote, 1895.	en	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): 1-55
03DC6105FFC06977FDB0A13EFC45CDB5.taxon	type_taxon	Type species. Charis incoides Schaus, 1902. Definition. Currently in the genus Lasaia Bates, 1868 (type species Papilio meris Stoll, 1781) but is not monophyletic with it and is far removed from it in the tree, being sister to the clade with several genera such as Pheles Herrich-Schäffer, [1858] (type species Pheles heliconides Herrich-Schäffer, [1853]) and Parcella Stichel, 1910 (type and the only species Amblygonia amarynthina C. & R. Felder, 1865) (Fig. 26) and therefore is a genus of its own. Similar to Lasaia in wing shape and color and male genitalia. Description and diagnostic characters for this new genus are as given for Lasaia cutisca Hall & Willmott, 1998 and Lasaia incoides on pages 23 – 24 and illustrated in Figs. 3 – 4, 9 – 10 by Hall & Willmott (1998). In brief, brown above and lacks blue or green scaling of Lasaia males, wings below uniform in background without paler patches and bands of most Lasaia species, hindwing outer margin lacks prominent concavity of Lasaia at vein M 2 (manifested also as a " tooth " at M 1); eyes setose; genitalic valvae narrow, about 3 times longer than broad, processus superior with rounded projection from ventral margin, processus inferior long and narrow, with a pointed tip.	en	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): 1-55
03DC6105FFC06977FDB0A13EFC45CDB5.taxon	etymology	Etymology. The name is a feminine noun in the nominative singular formed from the type species name with k for c to avoid a homonym. Species included. The type species and Lasaia cutisca Hall & Willmott, 1998. Parent taxon. Tribe Riodinini Grote, 1895.	en	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): 1-55
03DC6105FFC06977FDB0A13EFC45CDB5.taxon	description	Callistiumini Grishin, new tribe http: // zoobank. org / 3 BFAAAD 2 - 83 E 4 - 48 EF- 9952 - CA 85 DA 8 DEB 9 E	en	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): 1-55
03DC6105FFC06977FDB0A13EFC45CDB5.taxon	type_taxon	Type genus. Callistium Stichel, 1911. Definition. Presently, Callistium (type species Charis cleadas Hewitson, 1866) is kept in incertae sedis (Seraphim et al. 2018). Genomic analysis reveals that the type species of Callistium may be sister to Calydnini Seraphim, Freitas & Kaminski, 2018, albeit with very weak support (Fig. 27), while other species currently placed in Callistium belong to other genera (see below). Therefore, the Callistium lineage is of ancient origin, not confidently associated with any Riodinidae tribes and thus is a distinct Comments. Callistus Bonelli, 1810 (Coleoptera: Carabidae) is the type genus of Callistini. 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 " - ini ", avoids homonymy. However, it does not avoid confusion, including internet search engines that correct possible spelling errors. Therefore, the choice was to form the name as Callistiumini using the entire word as a stem (Art. 29.1.), instead of Callistiini.	en	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): 1-55
03DC6105FFC16977FF45A1F8FAE6CBBB.taxon	description	Placed in Callistium (type species Charis cleadas Hewitson, 1866) by Hall (2018), Calydna castanea Prittwitz, 1865 is not monophyletic with its type species, but instead is sister to the type species of Zabuella Stichel, 1911, which is Lemonias tenellus Burmeister, 1878 (Fig. 28), and is genetically close to it. Therefore, we agree Fig. 28. Zabuellina genera: Zabuella (cyan), Hallonympha with Seraphim et al. (2018) who reached the genera (magenta: Behemothia), and Teenie (magenta gen. n with. (orange subgenus); and Calicosama Theopina same conclusion, and confirm the combination in cyan), Archaeonympha (black), Theope (blue), Zabuella castanea (Prittwitz, 1865). Pseudotinea (black), and Petrocerus (red).	en	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): 1-55
03DC6105FFC66970FE74A725FAE1C86F.taxon	description	is not monophyletic with its type species, but is sister to	en	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): 1-55
03DC6105FFC66970FDB5A25EFBAECC8A.taxon	type_taxon	Type species. Calydna tinea Bates, 1868. Definition. Currently in the genus Zabuella Stichel, 1911 (type species Lemonias tenellus Burmeister, 1878) as suggested by Hall (2018), this group is not monophyletic with it and is sister to Hallonympha Penz & DeVries, 2006 (type species Apodemia paucipuncta Spitz, 1930), but prominently differs from it genetically and thus constitutes a genus (Fig. 28). Diagnostic characters for the new genus are as those detailed for Z. tinea and Z. argiella in Hall (2018). Briefly, similar to Zabuella and Hallonympha in wing patterns and genitalia, but aedeagus narrower, and signa highly asymmetrical both in position and size.	en	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): 1-55
03DC6105FFC66970FDB5A25EFBAECC8A.taxon	etymology	Etymology. The name is a feminine noun in the nominative singular inspired by the type species name. Species included. The type species (including Lemonias eudocia Godman & Salvin, 1897 as it synonym) and Calydna argiella Bates, 1868. Parent taxon. Subtribe Zabuellina Seraphim, Freitas & Kaminski, 2018.	en	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): 1-55
03DC6105FFC66971FD99A4B2FD34CF4D.taxon	type_taxon	Type species. Symmachia ocellata Hewitson, 1867. Definition. Currently in the genus Seco Hall & Harvey, 2002 (type species Charis calagutis Hewitson, 1871), but is not monophyletic with its type species, and instead forms an independent lineage in early radiation of the core Riodinini group (Fig. 29). Diagnostic characters for this new genus are as those given for Seco (except the characters specific to either Seco calagutis (Hewitson, 1871) or Exoplisia aphanis (Stichel, 1910) comb. nov. [then placed in Seco]) on pages 415 - 417 in Hall and Harvey (2002). Readily distinguished from its relatives by a black yellow-bordered eyespot at forewing apex.	en	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): 1-55
03DC6105FFC66971FD99A4B2FD34CF4D.taxon	etymology	Etymology. The name is a feminine noun in the nominative singular formed as a fusion of the type species name with the name of the genus it was formerly placed in: O [cellata] + [Se] co for its eyespots. Species included. Only the type species. Parent taxon. Tribe Riodinini Grote, 1895.	en	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): 1-55
03DC6105FFC76971FDCBA390FCD5CD2B.taxon	description	Willmott, 1995 (type species Machaya obstinata Hall & Willmott, 1995) originates deep within Pachythone Bates, 1868 (type species Pachythone erebia Bates, 1868) at the time of rapid diversification of its crown group (Fig. 30). Therefore, to restore the monophyly, we propose that Machaya is a junior subjective synonym of Pachythone.	en	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): 1-55
03DC6105FFC7694CFE5AA4D4FD7DC891.taxon	description	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 (2019 a), 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 / 46 A 68 C 99 - BBC 2 - 4 F 6 C-BB 06 - 4 CA 2 AD 2343 B 8	en	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): 1-55
03DC6105FFC7694CFE5AA4D4FD7DC891.taxon	type_taxon	Type genus. Callerebia Butler, 1867. Definition. Several genera of uncertain tribal placement (Wahlberg 2019 a) 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: hm 2009277 - RA. 13: T 2076 C, hm 2009277 - RA. 13: A 130 C, hm 2015715 - RA. 2: C 2137 T, hm 2009379 - RA. 4: T 1915 C, ahm 2002906 - RA. 2: A 128 G, where the part before the first dot (e. g. hm 2009277 - 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 A 130 C means base pair C at position 130, changed from A in the ancestor. See <https: // osf. io / kj 4 es /> 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.	en	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): 1-55
03DC6105FFC7694CFE5AA4D4FD7DC891.taxon	description	Parent Taxon. Tribe Satyrini Boisduval, 1833. Gyrocheilina Grishin, new subtribe http: // zoobank. org / EF 7 C 543 A- 88 B 9 - 4 DCC- 9847 - 139 CC 7763 B 84	en	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): 1-55
03DC6105FFC7694CFE5AA4D4FD7DC891.taxon	type_taxon	Type genus. Gyrocheilus Butler, 1867. Definition. Previously placed in Pronophilina Reuter, 1896 (Wahlberg 2019 a), 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.	en	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): 1-55
03DC6105FFC7694CFE5AA4D4FD7DC891.taxon	description	Parent Taxon. Tribe Satyrini Boisduval, 1833. Calistina Grishin, new subtribe http: // zoobank. org / F 1 D 9 A 7 DC-F 549 - 4 EC 9 - 8 C 68 - D 8032 DC 4 FB 95	en	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): 1-55
03DC6105FFC7694CFE5AA4D4FD7DC891.taxon	type_taxon	Type genus. Calisto Hübner, [1823]. Genera included. Only the type genus. Parent Taxon. Tribe Satyrini Boisduval, 1833.	en	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): 1-55
03DC6105FFC7694CFE5AA4D4FD7DC891.taxon	discussion	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.	en	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): 1-55
03DC6105FFFA694DFF72A4DCFA3FCE8A.taxon	description	COI barcodes between their type species (GenBank accession GQ 357203 for P. ornata) differ by 9.4 % (62 bp). Therefore Harsiesis Fruhstorfer, 1911 is a subgenus of Platypthima Rothschild & Jordan, 1905.	en	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): 1-55
03DC6105FFFB694DFF3BA2D3FC67CCB1.taxon	description	homonym, valid name Erebia merula Hewitson, 1875) cluster closely with Argyrophenga Doubleday, 1845 (type species Argyrophenga antipodum Doubleday, 1845) (Fig. 34), all being from New Fig. 34. Platypthima (magenta) and Argyrophenga (blue). Zealand and phenotypically similar. Therefore, Percnodaimon Butler, 1876 and Erebiola Fereday, 1879 are junior subjective synonyms of Argyrophenga Doubleday, 1845.	en	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): 1-55
03DC6105FFFB694FFDAAA0FCFA10CF1D.taxon	type_taxon	Type species. Erebia magdalena Strecker, 1880. broad with many small teeth in distal half (E. discoidalis), or long, more than 4 times longer than wide and with a broad-W-shaped dorsal margin (E. rossii (J. Curtis, 1835 )). The following combination of characters in the COI barcode region is diagnostic: 271 C (not T), 274 C (not T), 421 T or C (not A), 424 T (not A), 451 T or C (not A), 484 not T, and 622 T (not A).	en	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): 1-55
03DC6105FFFB694FFDAAA0FCFA10CF1D.taxon	etymology	Etymology. The name is a feminine noun in the nominative singular, formed from the type species name. Species included. The type species, Hipparchia cyclopius Eversmann, 1844, Papilio disa Thunberg, 1791, Hipparchia discoidalis Kirby, 1837, Erebia edda Ménétriés, 1851, Papilio embla Thunberg, 1791, Erebia erinnyn Warren, 1932, Erebia fasciata Butler, 1868, Erebia mackinleyensis Gunder, 1932, Erebia mancinus Doubleday, 1849, Hipparchia rossii Curtis, 1835, Erebia magdalena sachaensis Dubatolov, 1992, Erebia tristis tristior Goltz, 1937, and Erebia wanga Bremer, 1864. Parent taxon. Genus Erebia Dalman, 1816.	en	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): 1-55
03DC6105FFFB694FFDAAA0FCFA10CF1D.taxon	description	Additional genomic support for the broadly defined genera Cercyonis Scudder, 1875, Maniola Schrank, 1801 and Erebia Dalman, 1816 related phylogenetic lineages diversified around the same time, and if we chose the names of genera to reflect the pivotal common point in their evolution, the three-genus classification fits it best. As an alternative, we would be left with many small lineages as genera, and for internal consistency (i. e. taxonomic category represents a level in the classification), would need to split Erebia into at least 3 distinct genera, and Maniola into at least 5 (Fig. 36). We prefer to treat these smaller lineages as subgenera, as they do not represent the most prominent level of diversification, but the next one to it.	en	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): 1-55
03DC6105FFF9694FFF58A340FD3FCCF5.taxon	description	Moore, 1866) rendering it paraphyletic (Fig. 37). To restore monophyly and considering genetic closeness of these species, we propose that Hestinalis Bryk, 1938 is a junior subjective synonym of Mimathyma Moore, 1896. Curiously, Hestina Westwood, 1850 (type species Papilio assimilis Linnaeus, 1758) is in a different clade and thus is not synonymous with Hestinalis.	en	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): 1-55
03DC6105FFF96948FF6CA6C7FB67CDF4.taxon	description	The following taxa are junior subjective synonyms: Catacore Dillon, 1948 of Diaethria Billberg, 1820; Mesotaenia Kirby, 1871 with Orophila Staudinger, 1886 of Perisama Doubleday, 1849; and Paulogramma Dillon, 1948 of Catagramma Boisduval, 1836 Genomic sequencing and analysis of the type species of available genus-group names of the subtribe Callicorina Orfila, 1952 resulted in a fully resolved confident phylogeny (Fig. 39) that revealed close relationships between some of them. For instance, even the most distinctive of the type species for available genus-group names in the Diaethria Billberg, 1820 (type species Papilio clymena Cramer, 1775) clade sometimes separated in the monotypic genus Catacore Dillon, 1948 (type and the only species Catagramma kolyma Hewitson, 1851) clusters closely with Diaethria. The COI barcodes of Diaethria and Catacore type species are only 6.7 % (44 bp) different. In agreements with Wahlberg (2019 a), we treat Catacore Dillon, 1948 as a junior subjective synonym of Diaethria Billberg, 1820. Next, the three genera Perisama Doubleday, 1849 (type species Catagramma bomplandii Guérin-Méneville, [1844]), Mesotaenia Kirby, 1871 (type species Callitaenia doris C. & R. Felder, 1861, a subspecies of Catagramma vaninka Hewitson, 1855) and Orophila Staudinger, 1886 (type species Cybelis campaspe Hewitson, 1869, a subspecies of Cybdelis cardases Hewitson, 1869) cluster closely together (Fig. 39). Their COI barcodes also indicate a close relationship, e. g., the type species of Mesotaenia and Perisama are only 6.2 % (41 bp) different. Therefore, Fig. 39. Callicorina genera: Lucinia (black), Haematera (red: subgenus Antigonis, and cyan), we agree with Wahlberg (2019 a) and confirm that Catagramma (olive & pink), and Callicore (purple & Mesotaenia Kirby, 1871 and Orophila Staudinger, 1886 yellow: subgenus Diaethria, magenta & green: are junior subjective synonyms of Perisama Doubleday, subgenus Perisama, and blue). 1849. Furthermore, the genomic tree confirms the close relationship between Paulogramma Dillon, 1948 (type species Nymphalis pyracmon Godart, [1824]) and Catagramma Boisduval, 1836 (type species Catagramma hydaspes Boisduval, 1836, which is a junior subjective synonym of Nymphalis pygas Godart, [1824]). The reasons why Freitas et al. (2014) resurrected Paulogramma in favor of the older name Catagramma that they did not use are unclear. Their tree (Freitas et al. 2014: Fig. 3) shows " Callicore pygas ", the current name for the type species of Catagramma, inside Paulogramma as they define it. Furthermore, Wahlberg (2019 a) lists Paulogramma in synonymy with Catagramma. Therefore, we confirm that Paulogramma Dillon, 1948 is a junior subjective synonym of Catagramma Boisduval, 1836. Finally, we confirm that Lucinia Hübner, [1823] belongs to Callicorina (Fig. 39).	en	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): 1-55
03DC6105FFFE6948FE08A60EFABAC89C.taxon	description	Inspection of genetic diversification and prominence of tree branches, we find that, contrary to their wing pattern similarity, Catagramma Boisduval, 1836 (type species Catagramma hydaspes Boisduval, 1836, which is a junior subjective synonym of Nymphalis pygas Godart, [1824]) is a prominent genus strongly separated from others in the Callicore group, in agreement with Freitas et al. (2014) who called this genus by its junior synonym Paulogramma Dillon, 1948 (type species Nymphalis pyracmon Godart, [1824]) (Fig. 39). Therefore, we retain Catagramma as a genus. However, the other three taxa: Callicore Hübner, [1819] (type species Papilio astarte Cramer, 1779), Diaethria Billberg, 1820 (type species Papilio clymena Cramer, 1775) and Perisama Doubleday, 1849 (type species Catagramma bomplandii Guérin-Méneville, [1844]) cluster more closely and are less prominently separated from each other in the tree (Fig. 39), e. g., COI barcodes of the type species of differ by 8.8 % (58 bp). Therefore, we propose that Diaethria Billberg, 1820 and Perisama Doubleday, 1849 are subgenera of Callicore Hübner, [1819]. This adjustment of status (from genus to subgenus) makes their classification more internally consistent, and this newly broader Callicore becomes similar in genetic divergence to its relatives Eunica Hübner, [1819], Hamadryas Hübner, [1806], and Dynamine Hübner, [1819]. In wing patterns and shapes, the currently proposed subgenera of Callicore are rather similar to each other, and a number of erroneous attributions to former genera have been made in classifying these species (Freitas et al. 2014).	en	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): 1-55
03DC6105FFFE6949FEE8A4C0FA2FCE8A.taxon	description	39), e. g., their COI barcodes differ by 9.3 % (61 bp), suggesting that Antigonis C. Felder, 1861 is a subgenus of Haematera Doubleday, 1849, thus eliminating two not truly distinctive monotypic genera.	en	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): 1-55
03DC6105FFFF6949FEC2A2D3FE0ECDFC.taxon	description	Temenis Hübner, [1819] (type species Papilio merione Fabricius, 1781, which is Papilio laothoe Cramer, 1777) may not be monophyletic, with its type species being sister to Asterope Hübner, [1819] (type species Fig. 40. Temenis (colored) with its subgenera Asterope (purple), Nica (green), Callicorina (red), Peria (cyan). Oreas sapphira Hübner, [1816]) with medium support, and Temenis pulchra Hewitson, 1861, the type species of Callicorina Smart, 1976 possibly originating among other related genera, two of which are monotypic: Nica Hübner, [1826] (type and the only species Nymphalis flavilla Godart, [1824]) and Peria Kirby, 1871 (type and the only species Papilio lamis Cramer, 1779). Species of this group are close to each other, e. g., COI barcodes of the type species of Peria and Temenis differ by only 6.8 % (45 bp) and taken together prominently separate from the rest of the subtribe. Even if Temenis is monophyletic (i. e., if the topology in Fig. 40 is incorrect), it is not prominently distinct from its relatives, and therefore we propose that they all are congeneric. Thus, Asterope Hübner, [1819], Nica Hübner, [1826], Peria Kirby, 1871, and Callicorina Smart, 1976 are subgenera of Temenis Hübner, [1819]. Curiously, the subgenus Callicorina does not belong to the subtribe Callicorina.	en	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): 1-55
03DC6105FFFF6944FEDEA620FA58C995.taxon	description	Five genera from the subfamily Nymphalinae Rafinesque, 1815 were placed as incertae sedis by Wahlberg (2019 a): 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 (2019 b), 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 / D 5 C 0 AE 41 - 2615 - 4852 - 9 B 59 - 8533 C 27 E 4798	en	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): 1-55
03DC6105FFFF6944FEDEA620FA58C995.taxon	type_taxon	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.	en	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): 1-55
03DC6105FFFF6944FEDEA620FA58C995.taxon	description	Parent Taxon. Subfamily Nymphalinae Rafinesque, 1815. Rhinopalpini Grishin, new tribe http: // zoobank. org / 4 A 3 ACC 99 - A 761 - 49 B 9 - BA 00 - 00 F 83 FF 443 DE	en	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): 1-55
03DC6105FFFF6944FEDEA620FA58C995.taxon	type_taxon	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.	en	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): 1-55
03DC6105FFFF6944FEDEA620FA58C995.taxon	description	Parent Taxon. Subfamily Nymphalinae Rafinesque, 1815.	en	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): 1-55
03DC6105FFFF6944FEDEA620FA58C995.taxon	discussion	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 / 22 C 74 A 4 F- 6 B 24 - 4502 - 99 BB-B 8 E 3 C 66 AFABE	en	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): 1-55
03DC6105FFFF6944FEDEA620FA58C995.taxon	type_taxon	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.	en	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): 1-55
03DC6105FFFF6944FEDEA620FA58C995.taxon	description	Parent Taxon. Subfamily Nymphalinae Rafinesque, 1815. Vanessulini Grishin, new tribe http: // zoobank. org / D 584 B 74 C- 8 DC 8 - 479 A-AD 25 - 9197 A 03 A 843 B	en	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): 1-55
03DC6105FFFF6944FEDEA620FA58C995.taxon	type_taxon	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.	en	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): 1-55
03DC6105FFFF6944FEDEA620FA58C995.taxon	description	Parent Taxon. Subfamily Nymphalinae Rafinesque, 1815. Doleschalliaini Grishin, new tribe http: // zoobank. org / 88 D 32044 - DB 8 E- 486 F- 8 C 09 - 0 D 188 E 5 CDC 93	en	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): 1-55
03DC6105FFFF6944FEDEA620FA58C995.taxon	type_taxon	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.	en	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): 1-55
03DC6105FFFF6944FEDEA620FA58C995.taxon	description	Parent Taxon. Subfamily Nymphalinae Rafinesque, 1815.	en	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): 1-55
03DC6105FFFF6944FEDEA620FA58C995.taxon	discussion	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 29 A 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	en	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): 1-55
03DC6105FFF36945FF5FA427FA9CC998.taxon	description	Napeocles Bates, 1864 (type species Hamadryas jucunda Hübner, [1808]) clusters very closely with Siproeta Hübner, [1823] (type species Siproeta trayja Hübner, [1823], currently a subspecies of Siproeta epaphus (Latreille, [1813] )) (Fig. 44). The 4 species in this clade clump together, suggesting that Napeocles Bates, 1864 is a junior subjective synonym of Siproeta Hübner, [1823] despite profound differences in wing shape and patterns, a result of apparently convergent similarity with dead leaf mimics.	en	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): 1-55
03DC6105FFF16940FF32A663FBA2CB45.taxon	description	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.	en	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): 1-55
03DC6105FFF66941FDABA7A8FD54CFB2.taxon	type_taxon	Type species. Eresia orthia Hewitson, 1864. Definition. Previously placed in Ortilia Higgins, 1981 (type species Papilio liriope Cramer, 1775), this group is not monophyletic with it. Instead it is sister to all other Eresia sensu lato, and is close to them (Fig. 45). Previously discovered and defined as the Brazilian " Ortilia " clade by Wahlberg and Freitas (2007), this phylogenetic group was also confirmed and discussed in a more recent study suggesting " that this clade requires a new name " (Long et al. 2014). Due to its genetic closeness to Eresia sensu lato, this distinctive lineage is named as a subgenus of Eresia rather than a separate genus pending further analysis. It keys out to Ortilia in Higgins (1981) sharing the following diagnostic combination of characters with it: antennal club pyriform, aedeagus end without a pair of twisted processes, tegumen reduced, scaphial extensions small, without hooks and spines at angles, saccus single, narrow, finger-like, not expanding terminally, without a cleft. Differs from Ortilia in shorter and straighter harpe projecting directly caudodorsad (not arched at its origin changing direction from cephalodorsad) and less extensive or absent fulvous markings on wings above.	en	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): 1-55
03DC6105FFF66941FDABA7A8FD54CFB2.taxon	etymology	Etymology. The name is a feminine noun in the nominative singular, formed as a fusion Not + [Ort] ilia to indicate distinction from Ortilia. Species included. The type species, Phyciodes orticas Schaus, 1902, Phyciodes sejona Schaus, 1902, Eresia velica Hewitson, 1864, Eresia dicoma Hewitson, 1864, and Phyciodes polinella Hall, 1928. Parent taxon. Genus Eresia Boisduval, 1836.	en	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): 1-55
03DC6105FFF76941FDD9A3FBFD54CA12.taxon	type_taxon	Type species. Eresia levina Hewitson, 1872. Definition. Previously placed in Eresia Boisduval, 1836 (type species Nereis eunice Hübner, [1807]), this group is not monophyletic with it. Instead it is sister to all other Eresia sensu lato but Notilia subgen. n. (Fig. 45). It is a phenotypically distinctive subgenus diagnosed by wings rounder than Eresia with broad discal band across forewing, blue in the type species. Its male genitalia are distinctive, with diagnostic characters for this new subgenus as described for Eresia levina on page 150 and illustrated in Figs. 430 – 431 in Higgins (1981): i. e., genital capsule larger than in other Eresia, tegumen sclerotized, shoulders well-developed, scaphial extension expanded, sclerotized and terminally with many small teeth, valvae with fine teeth on inner sides near apex, aedeagus terminally rounded, without ostium keel.	en	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): 1-55
03DC6105FFF76941FDD9A3FBFD54CA12.taxon	etymology	Etymology. The name is a feminine noun in the nominative singular, formed from the type species name. Species included. Only the type species. Parent taxon. Genus Eresia Boisduval, 1836.	en	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): 1-55
03DC6105FFF76941FDA5A65CFD54C9A9.taxon	type_taxon	Type species. Phyciodes ithra Kirby, 1900. Definition. Previously placed in Ortilia Higgins, 1981 (type species Papilio liriope Cramer, 1775), this group is not monophyletic with it. Instead it is sister to Dagon Higgins, 1981 (type species Eresia catula Hopffer, 1874), but not with decisive statistical support (Fig. 45), and therefore is a distinctive lineage of the same rank, i. e., subgenus. Diagnosed by male genitalia and the characters for this subgenus are as those given for Ortilia ithra on page 120 and illustrated in Figs. 318 – 320 in Higgins (1981). Keys out to Ortilia in Higgins (1981) due to the following combination of characters this new subgenus shares with Ortilia and Notilia subgen. n.: pear-shaped (not extended) antennal club, no elongated processes at aedeagus end, reduced tegumen with small scaphial extensions not armed with hooks or spines, saccus single, terminally narrower, without a cleft. Differs from Ortilia and Notilia subgen. n. in having larger genital capsule, longer and bulkier saccus, prominent but short in dorsal view tegumen with very short scaphial extensions, short apical sections of valvae with thicker and rather straight harpes directed caudodorsad, aedeagus narrowing from phallobase caudad in basal half.	en	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): 1-55
03DC6105FFF76941FDA5A65CFD54C9A9.taxon	etymology	Etymology. The name is a feminine noun in the nominative singular, tautonymous with the type species name. Species included. Only the type species. Parent taxon. Genus Eresia Boisduval, 1836.	en	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): 1-55
03DC6105FFF46942FE53A25EFC73C893.taxon	description	Confident paraphyly of the species Phystis simois (Hewitson, 1864) (type locality Brazil, Fig. 47 left and right) with respect to Phystis fontus (A. Hall, 1928), comb. n. was one of the more unsettling results in this project (Fig. 48). However, COI barcodes of the two subspecies Phystis simois pratti (A. Hall, 1935) (type locality North Peru, Fig. 47 left) and Phystis simois variegata (Röber, 1913) (type locality Argentina, Fig. 47 right) differ by 6.4 % (42 bp), which is more than the difference between some of the Higgins genera (Higgins 1981), see above. Inspection of specimens (Fig. 47 left and right) reveals marked difference in wing shapes of the two subspecies, not commonly found within species. Therefore, these two subspecies are species that are also distinct from the nominotypical Phystis simois (Hewitson, 1864) (type locality Brazil) due to phenotypic differences and COI barcode differences (7.1 % & 8.1 %) between our specimens and P. simois available from GenBank (accession EF 493956) (Wahlberg and Freitas 2007). Hence, the names for these species are Phystis variegata (Röber, 1913) stat. nov. and Phystis pratti (A. Hall, 1935) stat. nov. We have not studied Phyciodes chinchipensis Hayward, 1964 (type locality Peru: Rio Chinchipe) currently treated as a subspecies of P. simois. Nevertheless, examination of the holotype photographs (Warren et al. 2016) suggests that it is not P. simois. Due to wing pattern resemblance and locality (both are from North Peru), we tentatively place it as a subspecies of P. pratti instead, keeping in mind that it may be a distinct species and not a mere color variation.	en	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): 1-55
03DC6105FFF46943FE4BA4DAFA85CCD7.taxon	description	(42 bp), the same divergence as between the two former subspecies of Phystis simois (see above), and therefore are congeneric. The Actinote clade is separated from them by a prominent gap (Carvalho et al. 2020), and the barcodes of A. thalia and A. neleus differ by 8.5 % (56 bp), which would suggest their attribution to different subgenera. However, species phylogeny is at odds with intuitive phenotypic assessment (Lamas 2004): there is an additional third clade unexpected from phenotypes (Silva-Brandao et al. 2008; Carvalho et al. 2020). Therefore, until these inconsistencies between phenotypic and genetic classifications of this group are sorted out, we propose placing Abananote Potts, 1943 and Altinote Potts, 1943 as junior subjective synonyms of Actinote Hübner, [1819]. This treatment is consistent with the unified, and more genetically diverse, Old World genus Telchinia Hübner, [1819] (type species Papilio serena Fabricius, 1775) that is sister to Actinote sensu lato (Fig. 49).	en	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): 1-55
03DC6105FFF56943FE96A107FA20CB23.taxon	description	and Ceratinia Hübner, 1816 (type species Nerëis neso Hübner, [1806]) (Willmott and Freitas 2006): the two genera are closely related and Episcada renders Ceratinia paraphyletic (Fig. 50), in agreement with other studies (Chazot et al. 2020). COI barcodes of the Episcada and Ceratinia type species differ by only 4.4 % (29 bp), less than between the two former subspecies of Phystis simois (6.4 %, see above). Therefore, we propose treating Episcada Godman & Salvin, 1879 as a junior subjective synonym of Ceratinia Hübner, 1816.	en	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): 1-55
03DC6105FFF5695CFF73A7B1FA23CC1F.taxon	description	(type locality Mexico) and Phocides lincea (HerrichSchäffer, 1869) (type locality not stated, probably the Guianas), and suggesting that P. pigmalion is a Fig. 51. Phocides batabano (red), bicolora (magenta), complex of several species. While the inclusion of pigmalion (blue), lincea (green) and perkinsi (cyan). P. belus in the pigmalion group is expected because Evans (1952) treated it as a subspecies of P. pigmalion, P. lincea was a surprise. Linked by Evans (1952) with Phocides perkinsi (Kaye, 1931) (type locality Jamaica) as its subspecies that was elevated to species only recently (Turner and Turland 2017), and placed at the end of Evans' key, P. lincea has not been associated with the pigmalion group before. Our genomic results definitively confirm P. perkinsi (Fig. 51 cyan) as a species-level taxon, because it is far removed from P. lincea (Fig. 51 green). Furthermore, due to genetic and genitalic differences, we reinstate Phocides batabano (Lucas, 1857) (type locality Cuba) and Phocides bicolora (Boddaert, 1783) (type locality not stated, likely Haiti) as species. The COI barcodes of P. batabano and P. bicolora differ by 2 % (13 bp), and P. pigmalion (from Ecuador) and P. batabano by 3.3 % (22 bp). Wing patterns agree with this partitioning of P. pigmalion as it was defined by Mielke (2005) previously, into at least three species: forewing hyaline spots absent (P. batabano), present but narrow and wings green-striped (P. bicolora), and present and broader, wings blue-striped (P. pigmalion). To accommodate this treatment, we revise species-subspecies combinations as: Phocides batabano okeechobee (Worthington, 1881) and Phocides batabano batabanoides (W. Holland, 1902).	en	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): 1-55
03DC6105FFEA695CFE0BA04FFA26CB23.taxon	description	(2004) concluded that " valeriana ... is clearly conspecific with T. mysie. " We sequenced the lectotypes of L. valeriana and L. mysie and found that their COI barcodes differ by 2.1 % (14 bp). Furthermore, the genomic tree of L. valeriana specimens from across the range revealed their partitioning into two clades, each clade with its type specimen: mysie from the US (Fig. 52 purple), and valeriana from various localities in Mexico (Fig. 52 blue). Fst / Gmin statistics for these clades were 0.41 / 0.04, suggesting that they represent distinct species (Cong et al. 2019 a; Zhang et al. 2020). Therefore, we reinstate Lobotractus mysie (Dyar, 1904) as a species different from Lobotractus valeriana (Plötz, 1881), and consequently exclude L. valeriana from the US fauna. Curiously, Burns (1996) and (2001) reached similar conclusion as far as the US fauna is concerned, but likely for incorrect reasons.	en	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): 1-55
03DC6105FFEA695CFF5EA709FAA6C9BC.taxon	description	Lake around Dalton Hwy mi. 274 in Alaska is not monophyletic with North American populations, but instead belongs to the Old World lineage (Fig. 53). Therefore, it is not Pyrgus centaureae freija (B. Warren, 1924) (type locality Labrador), but Pyrgus Fig. 53. Pyrgus centaureae: Palearctic lineage (blue) centaureae dzekh Gorbunov, 2007 (type locality and Nearctic lineage (magenta). Russia: Chukotka). Hence, we add this subspecies to North American fauna. The tree reveals partitioning of P. centaureae into two clades: mostly Palearctic lineage (Fig. 53 blue) and Nearctic lineage (Fig. 53 magenta). However, COI barcodes of specimens from different lineages (e. g. Sweden vs. USA: WV) differ by only 0.15 % (1 bp) and Fst / Gmin statistics of the two lineages are 0.21 / 0.08, suggesting rather limited genetic differentiation and continuing gene exchange between them. On the basis of statistics from this small sample of specimens we sequenced, we cannot yet support the two lineages as distinct species, and the Nearctic assemblage of subspecies may be conceptualized as a semi-species instead.	en	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): 1-55
03DC6105FFEB695DFED4A02EFE56C85C.taxon	description	Fabricius, 1793). Confirming our previous assessment (Zhang et al. 2019 c), we see that Polites Scudder, 1872 (type species Hesperia peckius W. Kirby, 1837) is genetically close to Wallengrenia Berg, 1897 (type species Hesperia premnas Wallengren, 1860) and is not separated from it by a long internal branch (Fig. 54). Both branches that are labeled " Polites " in Fig. 54 are longer than the branch between them and therefore assigning a taxonomic rank to them seems more appropriate than to the shorter branch between them. Moreover, COI barcodes of W. otho and P. peckius differ by 5.2 % (34 bp) further confirming their close relationship. Therefore, we propose treating Wallengrenia Berg, 1897 as a subgenus of Polites Scudder, 1872. The resulting classification of the Hylephila Billberg, 1820 (type species Papilio phyleus Drury, 1773) group is marked on the tree rooted with Hesperia comma (Linnaeus, 1758) (Fig. 54). Hylephila is sister to all others in the group. Then, there are two major levels in the tree. First, the genus Polites that includes Wallengrenia originates at about the same level as other three genera in the group: Pompeius Evans, 1955 (type species Hesperia pompeius Latreille, [1824]), Hedone Scudder, 1872 (type species Hesperia brettus Boisduval & Le Conte, [1837], a junior subjective synonym of Thymelicus vibex Geyer, 1832), and Limochores Scudder, 1872 (type species Hesperia manataaqua Scudder, 1863, which is a junior subjective synonym of Hesperia origenes Fabricius, 1793). This diversification event dates to about 15 Mya (Zhang et al. 2019 d) and therefore corresponds to genera. Second, Polites diversifies into 4 prominent lineages: Polites, Yvretta Hemming, 1935 Coa Grishin, 2019 and Wallengrenia Berg, 1897. This more recent diversification (~ 10 Mya) corresponds to subgenera.	en	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): 1-55
