identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
03DC6105FFDF6969FE16A3D5FD72CB79.text	03DC6105FFDF6969FE16A3D5FD72CB79.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Boreographium Grishin 2021	<div><p>Boreographium Grishin, new subgenus</p><p>http://zoobank.org/ 32D82A5D-2DEB-4685-8179-D5FF9E5F2225</p><p>Type species. Papilio marcellus Cramer, 1777 .</p><p>Definition. As revealed by genome-scale phylogenetic trees (Zhang et al. 2019d; Zhang et al. 2019c), the type species of this new subgenus is in the same clade with the subgenus</p><p>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).</p><p>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.</p><p>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).</p><p>Species included. Only the type species.</p><p>Parent taxon. Genus Eurytides Hübner, [1821] .</p></div>	https://treatment.plazi.org/id/03DC6105FFDF6969FE16A3D5FD72CB79	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFDF696AFDF7A7A3FD61CF97.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Esperourus Grishin 2021	<div><p>Esperourus Grishin, new subgenus</p><p>http://zoobank.org/ DC085A9A-4391-49F3-B3AE-F2F57B0BB76C</p><p>Type species. Papilio esperanza Beutelspacher, 1975 .</p><p>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.</p><p>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.</p><p>Species included. Only the type species.</p><p>Parent taxon. Genus Pterourus Scopoli, 1777 .</p></div>	https://treatment.plazi.org/id/03DC6105FFDF696AFDF7A7A3FD61CF97	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFDC696AFDF0A3C6FC80CB91.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	(Hyppasonia) Zhang & Cong & Shen & Opler & Grishin 2021	<div><p>Hyppasonia Grishin, new subgenus</p><p>http://zoobank.org/ B37D0541-CD40-4824-AE21-BD6E37D37CB7</p><p>Type species. Papilio hyppason Cramer, 1775 .</p><p>Definition. Previously, we noted this unnamed subgenus in the COI barcode dendrogram</p><p>(Shiraiwa et al. 2014). Here, its definition is formalized. While the COI barcodes demonstrate its distinction from other subgenera of</p><p>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</p><p>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.</p><p>Etymology. The name is a feminine noun in the nominative singular, formed from the type species name.</p><p>Species included. Only the type species.</p><p>Parent taxon. Genus Heraclides Hübner, [1819] .</p></div>	https://treatment.plazi.org/id/03DC6105FFDC696AFDF0A3C6FC80CB91	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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
03DC6105FFDC696AFE9EA405FCE6C9AF.text	03DC6105FFDC696AFE9EA405FCE6C9AF.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Prestonia Schaus 1920	<div><p>Prestonia Schaus, 1920 belongs to Euremini Grote, 1898</p><p>The monotypic genus Prestonia Schaus, 1920 (type and the only species Prestonia clarki Schaus, 1920) has been tentatively placed near Phoebis Hübner, [1819] (type species Phoebis cypris Hübner, [1819], a junior subjective synonym of Papilio argante Fabricius, 1775) due to phenotypic similarity and no DNA sequences available for it. The genomic tree reveals that subfamily Coliadinae Swainson, 1821 splits into two clades that we treat as tribes: Coliadini Swainson, 1821 (includes Phoebis) and Euremini Grote, 1898 (Fig. 5). Genomic-scale phylogeny confidently places Prestonia as sister to Kricogonia Reakirt, 1863 (type species Colias lyside Godart, 1819) and therefore Prestonia belongs to Euremini Grote, 1898 (the clade that does not include Phoebis) and not to Coliadini.</p></div>	https://treatment.plazi.org/id/03DC6105FFDC696AFE9EA405FCE6C9AF	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFDD696BFE44A64BFA17C86E.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Pyrisitia westwoodii (Boisduval 1836)	<div><p>Pyrisitia westwoodii (Boisduval, 1836) is a species distinct from Pyrisitia dina (Poey, 1832)</p><p>Currently considered a subspecies of Pyrisitia dina (Poey, 1832) (type locality Cuba), Terias westwoodii Boisduval, 1836 (type locality Mexico) is prominently separated genetically from the insular taxa (Fig. 6).</p><p>Compared to Pyrisitia dina helios (M. Bates, 1934)</p><p>(type locality Bahamas) the Fst/Gmin statistics are</p><p>0.46/0.03 and their COI barcodes are 2.7% (18 bp)</p><p>different. Therefore, we propose that Pyrisitia Fig. 6. Pyrisitia westwoodii (red), and dina (blue)</p><p>and parvumbra (Jamaica).</p><p>westwoodii (Boisduval, 1836), reinstated status is a species-level taxon and employ Pyrisitia westwoodii gabriela Le Crom &amp; Llorente, 2004 comb. nov. We also confirm Pyrisitia parvumbra (Kaye, 1925) as a species (Fig. 6; 2.6%, 17 bp barcode difference).</p></div>	https://treatment.plazi.org/id/03DC6105FFDD696BFE44A64BFA17C86E	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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
03DC6105FFDD696BFDE9A165FA3DCA61.text	03DC6105FFDD696BFDE9A165FA3DCA61.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Terias Swainson 1821	<div><p>Terias Swainson, 1821 is a valid genus</p><p>Our genomic tree reveals that Eurema Hübner, [1819] (type species Papilio delia Cramer, 1780, a junior homonym: valid name is Pieris daira Godart, 1819) is paraphyletic with respect to Pyrisitia Butler, 1870 (type species Papilio proterpia Fabricius, 1775) (Fig. 5). To restore monophyly, we choose to keep Pyrisitia as a genus and therefore treat the Old World clade currently placed in Eurema as a distinct valid genus. Terias Swainson, 1821 (type species Papilio hecabe Linnaeus, 1758) is its oldest available name.</p></div>	https://treatment.plazi.org/id/03DC6105FFDD696BFDE9A165FA3DCA61	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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
03DC6105FFDD696BFDFAA4BFFA58C982.text	03DC6105FFDD696BFDFAA4BFFA58C982.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Zegris Boisduval 1836	<div><p>Zegris Boisduval, 1836 is a subgenus of Anthocharis Boisduval, Rambur, [Duménil] &amp; Graslin, [1833]</p><p>Zegris Boisduval, 1836 (type species Papilio eupheme Esper, [1804]) originates within Anthocharis Boisduval, Rambur, [Duménil] &amp; Graslin, [1833] (type species Papilio cardamines Linnaeus, 1758), rendering it paraphyletic, and is confidently placed as a sister to subgenus Paramidea Kuznetsov, 1929 (type species: Anthocharis scolymus Butler, 1866) (Fig. 7). Subgenera of Anthocharis are genetically close to each other and even to the sister genus Euchloe (type species Euchloe ausonia var. esperi W. F.</p></div>	https://treatment.plazi.org/id/03DC6105FFDD696BFDFAA4BFFA58C982	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFD26965FE81A14AFA2ECCBE.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Hesperocharis leucothea (Molina 1782) Zhang & Cong & Shen & Opler & Grishin 2021	<div><p>Hesperocharis leucothea (Molina, 1782) new combination</p><p>Papilio leucothea Molina, 1782 (type locality Chile), currently placed in the genus Mathania Oberthür, 1890, originates within Hesperocharis Felder, 1862 (type species Pieris erota Lucas, 1852) according to the genome-scale tree (Fig. 8), which implies Hesperocharis leucothea (Molina, 1782) comb. n.</p><p>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</p><p>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).</p></div>	https://treatment.plazi.org/id/03DC6105FFD26965FE81A14AFA2ECCBE	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFD36965FDC0A0E0FD15C888.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Sisymbria Grishin 2021	<div><p>Sisymbria Grishin, new subgenus</p><p>http://zoobank.org/ 28C486B5-3F65-4CDD-AC44-3FE386B58D0B</p><p>Type species. Pieris sisymbrii Boisduval, 1852 .</p><p>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 .</p><p>Etymology. The name is a feminine noun in the nominative singular, formed from the type species name.</p><p>Species included. Only the type species.</p><p>Parent taxon. Genus Pontia [Fabricius], 1807.</p><p>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.</p></div>	https://treatment.plazi.org/id/03DC6105FFD36965FDC0A0E0FD15C888	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFD36967FF28A4D5FA46CFD8.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Tatochila A. Butler 1870	<div><p>Tatochila A. Butler, 1870, Piercolias Staudinger, 1894, Hypsochila Ureta, 1955, Theochila W. D. Field, 1958, Pierphulia W. D. Field, 1958, and Infraphulia W. D. Field, 1958 are junior subjective synonyms of Phulia Herrich-Schäffer, 1867</p><p>Phulia Herrich-Schäffer, 1867 (type species Pieris nymphula Blanchard, 1852), Tatochila A. Butler, 1870 (type species Synchloe autodice Hübner, [1818]), Piercolias Staudinger, 1894 (type species Trifurcula huanaco Staudinger, 1894), Hypsochila Ureta, 1955 (type species Tatochila microdice f. wagenknechti Ureta, 1938), Theochila W. D. Field, 1958 (type species Pieris maenacte Boisduval, 1836), Pierphulia W. D. Field, 1958 (type species Phulia nysias Weymer, 1890), and Infraphulia W. D. Field, 1958 (type species Phulia nymphula var. illimani Weymer, 1890) cluster closely in the genomic tree without obvious separation into groups (Fig. 9 green). E.g., COI barcodes of P. nymphula and T. autodice differ by 4.0% (26 bp). However, being combined into one, all these genera together represent a prominently distinct genetic group that is sister to Ascia Scopoli, 1777 (type species Papilio monuste Linnaeus, 1764) and more distantly related to Ganyra Billberg, 1820 (type species Papilio amaryllis Fabricius, 1793, a junior homonym: valid name for this species is Pieris josephina Godart, 1819). With genetic similarity between these taxa being at the level of a species group, it may not be meaningful to consider these names as denoting valid subgenera, and therefore we propose that Tatochila A. Butler, 1870, Piercolias Staudinger, 1894, Hypsochila Ureta, 1955, Theochila W. D. Field, 1958, Pierphulia W. D. Field, 1958, and Infraphulia W. D. Field, 1958 are junior subjective synonyms of Phulia Herrich-Schäffer, 1867 .</p><p>Calopierini Grishin, new tribe</p><p>http://zoobank.org/ B7717ECE-C015-48AA-A57D-B7A6A3CCE4F6</p><p>Type genus. Calopieris Aurivillius, 1898 .</p><p>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.</p><p>Genera included. Only the type genus.</p><p>Parent Taxon. Subfamily Pierinae Swainson, 1820 .</p><p>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).</p><p>Phrissura Butler, 1870 is a subgenus of Appias Hübner, [1819] Saletara Distant, 1885 is a subgenus of Catophaga Hübner, 1819</p><p>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. &amp; 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 .</p></div>	https://treatment.plazi.org/id/03DC6105FFD36967FF28A4D5FA46CFD8	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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
03DC6105FFD16960FF46A454FB79CFD5.text	03DC6105FFD16960FF46A454FB79CFD5.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Catasticta Butler 1870	<div><p>Catasticta Butler, 1870 is a junior subjective synonym of Archonias Hübner, 1827</p><p>Catasticta Butler, 1870 (type species Euterpe nimbice Boisduval, 1836) clusters closely with Archonias Hübner, [1829] (type species Archonias marcias Hübner, 1825 which is currently treated as a subspecies of Papilio brassolis Fabricius, 1777 from Brazil: Bahia) in the genomic tree (Fig. 12). Genetic diversification between these genera is within the norm for most Pieridae genera, e.g., Mylothris shown in the same tree. Moreover, previously published phylogenetic trees reveal that Catasticta is not monophyletic (Padrón 2014; Wahlberg et al. 2014), although sometimes with low support. Furthermore, Padrón (2014) discussed this issue at length and suggested synonymy between Catasticta and Archonias . We agree with these conclusions, also supported by our genome-scale results. Although further studies are required to develop a meaningful classification within this very large genus, three independent studies provide phylogenetic data suggesting that Catasticta Butler, 1870 is a junior subjective synonym of Archonias Hübner, 1827 . This name change may not be welcomed by those used to the name Catasticta, but it will be necessary to accept at one point. While some may consider breaking this large genus into several genera, the genomic tree indicates that other relatives of Catasticta, such as Charonias Röber, 1908 (type species Euterpe eurytele Hewitson, 1853), Neophasia Behr, 1869 (type species Pieris menapia C. &amp; R. Felder, 1859), and Eucheira Westwood, 1834 (type and the only species Eucheira socialis Westwood, 1834) are all closely allied to each other and may be treated as subgenera of Archonias, along with some others currently placed in Catasticta . However, details of such classification will be revealed after genomic-scale sequencing of all major species groups in this complex is completed.</p></div>	https://treatment.plazi.org/id/03DC6105FFD16960FF46A454FB79CFD5	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFD16967FF3AA005FDD6CDB0.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Mesapia Gray 1856	<div><p>Mesapia Gray, 1856 is a junior subjective synonym of Aporia Hübner, [1819]</p><p>A monotypic genus Mesapia Gray, 1856 (type and the only species Pieris peloria Hewitson, 1853) originates within Aporia Hübner, [1819] (type species</p><p>Papilio crataegi Linnaeus, 1758) rendering it paraphyletic</p><p>(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] .</p></div>	https://treatment.plazi.org/id/03DC6105FFD16967FF3AA005FDD6CDB0	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFD66960FDCDA002FA98C9BC.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Lycaenidae	<div><p>Family Lycaenidae [Leach], [1815]</p><p>Sandia Clench &amp; P. Ehrlich, 1960 and Xamia Clench, 1961 are junior subjective synonyms of Incisalia Scudder, 1872</p><p>Appearance could be misleading, especially if unusual. Despite unique wing patterns, we find from the genomic analysis that Sandia Clench &amp; P. Ehrlich, 1960 (type and the only species Callophrys (Sandia) mcfarlandi P. Ehrlich &amp; 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.</p><p>Although Xamia is more distant from other congeners than Sandia,</p><p>it still falls within the prominent</p><p>Incisalia clade. Thus leaving it as a valid subgenus would require a new name for at least Callophrys polios (Cook &amp; F. Watson, 1907),</p><p>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 &amp; 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 &amp; 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.</p></div>	https://treatment.plazi.org/id/03DC6105FFD66960FDCDA002FA98C9BC	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFD76961FEA9A738FA5AC905.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Corrachia Schaus 1913	<div><p>Corrachia Schaus, 1913 is a subgenus of Styx Staudinger, 1876</p><p>Two monotypic genera Styx Staudinger, 1876 (type and the only species Styx infernalis Staudinger, 1875) and Corrachia Schaus, 1913 (type and the only species Corrachia leucoplaga Schaus, 1913) are close sisters (Fig. 14 red and green). Hindered by substantial difference in appearance, their close kinship has been revealed by DNA sequencing and discussed in detail (Espeland et al. 2015). Their COI barcodes differ by only 7.4% (49 bp). Here, we take the next step and eliminate the two monotypic genera</p><p>Fig. 14. Nemeobiini genera: Hamearis (black), Styx</p><p>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).</p><p>about their evolutionary relationship than keeping them in two not prominently distinct monotypic genera.</p></div>	https://treatment.plazi.org/id/03DC6105FFD76961FEA9A738FA5AC905	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFD76961FDD3A2B2FD7ACDBD.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Greenie Grishin 2021	<div><p>Greenie Grishin, new subgenus</p><p>http://zoobank.org/ 733681D0-34AF-4884-9396-BE415705E783</p><p>Type species. Thecla sheridonii [sic] Edwards, 1877, presently Callophrys sheridanii (Edwards, 1877) .</p><p>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:T366C, cce8426.17.3:C4290T, cce10587.6.6:G181C, cce925.9.4:C351T, and cce2041.25.2:T1572A; and the COI barcode: 479C (not T) and 610C (not T), differing from the subgenus Callophrys in 202T (not A), 512G (not T), and 556T (not A). See &lt;https://osf.io/kj4es/&gt; for the sequences with these characters.</p><p>Etymology. The name is a feminine noun in the nominative singular, Latinized verbatim from the affectionate English name of this butterfly group.</p><p>Species included. The type species, Thecla viridis W. H. Edwards, 1862, Thecla dumetorum Boisduval, 1852, and Thecla affinis W. H. Edwards, 1862 .</p><p>Parent taxon. Genus Callophrys Billberg, 1820 .</p></div>	https://treatment.plazi.org/id/03DC6105FFD76961FDD3A2B2FD7ACDBD	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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
03DC6105FFD76962FF76A56FFDDFCF38.text	03DC6105FFD76962FF76A56FFDDFCF38.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Hades Westwood 1851	<div><p>Hades Westwood, 1851 is a junior subjective synonym of Methone Doubleday, 1847</p><p>Despite their difference in appearance, monotypic genus Methone Doubleday, 1847 (type and the only species Papilio cecilia Cramer, 1777) is genetically close to Hades Westwood, 1851 (type species Hades noctula Westwood, 1851) (Fig. 15 brown and gray). Therefore, we propose that Hades is a junior subjective synonym of Methone and deduce that the phenotypic difference between Hades and Methone was caused by rapid evolution possibly driven by selection for mimetic appearance of their type species, each in a different mimetic complex.</p></div>	https://treatment.plazi.org/id/03DC6105FFD76962FF76A56FFDDFCF38	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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
03DC6105FFD76961FF74A627FA8DCAFB.text	03DC6105FFD76961FF74A627FA8DCAFB.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Takashia M. Okano & T. Okano 1985	<div><p>Takashia M. Okano &amp; T. Okano, 1985 is a subgenus of Polycaena Staudinger, 1886</p><p>Monotypic genus Takashia M. Okano &amp; T. Okano, 1985 (type and the only species Timelaea nana Leech, 1892) is a close sister to Polycaena Staudinger, 1886 (type species Polycaena tamerlana Staudinger, 1886) (Fig. 14 magenta and blue). To avoid yet another not truly distinct monotypic genus, we propose to place Takashia M. Okano &amp; T Okano, 1985 as a subgenus of Polycaena Staudinger, 1886 .</p></div>	https://treatment.plazi.org/id/03DC6105FFD76961FF74A627FA8DCAFB	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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
03DC6105FFD46962FF31A365FACACD3F.text	03DC6105FFD46962FF31A365FACACD3F.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Methone Doubleday 1847	<div><p>Methone Doubleday, 1847 receives 9 species from Euselasia Hübner, [1819]</p><p>We find that Euselasia Hübner, [1819] (type species Euselasia gelaena Hübner, [1819], which is Papilio gelon Stoll, 1787) as currently defined is paraphyletic with respect to Methone Doubleday, 1847 (type species Papilio cecilia Cramer, 1777), and a number of Euselasia species belong to the clade with Methone cecilia (Fig. 15 red and brown). The type species of Euselasia and Methone are genetically distant from each other, e.g., their COI barcodes differ by 11.5% (76 bp), which is typical for species in different genera. Therefore, to restore the monophyly, instead of placing Methone in Euselasia, we transfer Methone -clade species (Fig. 15 red) and their phenotypically close relatives from Euselasia to Methone to form the following new combinations: Methone euploea (Hewitson, [1855]), Methone eucerus (Hewitson, 1872), Methone hypophaea (Godman &amp; Salvin, 1878), Methone eubule (R. Felder, 1869), Methone onorata (Hewitson, 1869), Methone authe (Godman, 1903), Methone dolichos (Staudinger, [1887]), Methone baucis (Stichel, 1919), and Methone eucrates (Hewitson, 1872) .</p></div>	https://treatment.plazi.org/id/03DC6105FFD46962FF31A365FACACD3F	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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
03DC6105FFD46963FF1FA16FFAA7CEEE.text	03DC6105FFD46963FF1FA16FFAA7CEEE.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Erythia Hubner 1819	<div><p>Erythia Hübner, [1819] and Marmessus Hübner, [1819] are valid genera</p><p>Hübner, 1823, which is a junior subjective synonym of Hesperia thucydides Fabricius, 1793, is in the Erythia clade, making Psalidopteris a junior subjective synonym of Erythia . Thus, four of these major clades have names and five do not, corresponding to five new genera that are proposed next.</p></div>	https://treatment.plazi.org/id/03DC6105FFD46963FF1FA16FFAA7CEEE	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFD56963FDA6A33FFC9DCB9B.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Pelolasia Grishin 2021	<div><p>Pelolasia Grishin, new genus</p><p>http://zoobank.org/ A9AC96C6-B5D0-416D-9C5E-0853B7BD8339</p><p>Type species. Eurygona pelor Hewitson, [1853] .</p><p>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:T87C, cne3355. 8.1:T362C, cne 1314.4.1:A351G, cne 2022.5.2:C448A, and cne599.10.1:T5841A.</p><p>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.</p><p>Species included. The type species, Eurygona amphidecta Godman &amp; Salvin, 1878, Eurygona argentea Hewitson, 1871, Eurygona artos Herrich-Schäffer, [1853], Eurygona aurantia Butler &amp; 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 &amp; J. Vargas, 2019, Euselasia nytua J. Hall &amp; Willmott, 2009, Euselasia pellonia Stichel, 1919, Euselasia rubrocilia Lathy, 1926, and Euselasia seitzi Lathy, 1926 .</p><p>Parent taxon. Subfamily Euselasiinae Kirby, 1871 .</p></div>	https://treatment.plazi.org/id/03DC6105FFD56963FDA6A33FFC9DCB9B	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFD5697CFDACA7C2FC9DCD22.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Myselasia Grishin 2021	<div><p>Myselasia Grishin, new genus</p><p>http://zoobank.org/ 61CDF024-ADCB-49D0-82FA-9956EA9E5553</p><p>Type species. Eurygona mys Herrich-Schäffer, [1853] .</p><p>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: cne23605.2.5:C468T, cne13338.5.3: A334C, cne123.2.3:C1009T, cne18035.2.1:C172A, and cne81.14.5:A756G.</p><p>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.</p><p>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 &amp; 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 &amp; Godman, 1868, Papilio hygenius Stoll, 1787, Euselasia illarina J. Hall, Willmott &amp; R. Busby, 1998, Eurygona inconspicua Godman &amp; Salvin, 1878, Euselasia janigena Stichel, 1919, Euselasia jigginsi J. Hall &amp; Willmott, 1998, Eurygona leucon Schaus, 1913, Euselasia mapatayna J. Hall &amp; Willmott, 1998, Euselasia marica Stichel, 1919, Eurygona mystica Schaus, 1913, Euselasia nauca J. Hall &amp; Willmott, 1998, Euselasia pance Callaghan, 1999, Eurygona procula Godman &amp; Salvin, 1885, Euselasia pseudomys Callaghan, 1999, Euselasia pullata Stichel, 1927, Eurygona pusilla R. Felder, 1869, Euselasia rhodon Seitz, 1913, and Eurygona sergia Godman &amp; Salvin, 1885 .</p><p>Parent taxon. Subfamily Euselasiinae Kirby, 1871 .</p></div>	https://treatment.plazi.org/id/03DC6105FFD5697CFDACA7C2FC9DCD22	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFCA697CFDA2A10CFC9DC816.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Eurylasia Grishin 2021	<div><p>Eurylasia Grishin, new genus</p><p>http://zoobank.org/ 44399705-A1CF-43AF-B35E-F510E964BDE0</p><p>Type species. Eurygona euryone Hewitson, 1856 .</p><p>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: cne9878.8.1:C146A, cne178.3.20:C595T, cne178.3.20:T596C, cne7676.26.2: T31C, and cne5931.2.1:A478G.</p><p>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.</p><p>Species included. The type species, Eurygona effima Hewitson, 1869, Euselasia thusnelda Möschler, 1883, and Eurygona eunaeus Hewitson, 1855 .</p><p>Parent taxon. Subfamily Euselasiinae Kirby, 1871 .</p></div>	https://treatment.plazi.org/id/03DC6105FFCA697CFDA2A10CFC9DC816	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFCA697DFDABA458FC9DCCB4.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Maculasia Zhang & Cong & Shen & Opler & Grishin 2021	<div><p>Maculasia Grishin, new genus</p><p>http://zoobank.org/ 20121C52-93F8-4540-9070-FF2971B2D960</p><p>Type species. Euselasia albomaculiga Callaghan, 1999 .</p><p>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:A2104A (not C), cne5129.1.5: A833A (not G), cne2685.14.3:A934A (not T), cne 1095.7.22:G1463G (not A), cne4870.1.18:T552T (not A), cne 2885.9.10:A4270C, cne1547.14.4:A2756G, cne 1696.1.1:A2529T, cne7231.10.9:T567C, and cne 1036.6.11:G3200C.</p><p>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.</p><p>Species included. Only the type species.</p><p>Parent taxon. Subfamily Euselasiinae Kirby, 1871 .</p></div>	https://treatment.plazi.org/id/03DC6105FFCA697DFDABA458FC9DCCB4	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFCB697DFDACA0FAFC9DCB78.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Eugelasia Grishin 2021	<div><p>Eugelasia Grishin, new genus</p><p>http://zoobank.org/ 36F889D1-2E27-40BB-95B9-7619031A223E</p><p>Type species. Eurygona eugeon Hewitson, 1856 .</p><p>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:C203A, cne 1015.3.2:T555C, cne2803.19.1:G88A, cne5471.1.1:T261C, and cne703.2.8:A1414C.</p><p>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.</p><p>Species included. The type species and Euselasia brevicauda Lathy, 1926 .</p><p>Parent taxon. Subfamily Euselasiinae Kirby, 1871 .</p></div>	https://treatment.plazi.org/id/03DC6105FFCB697DFDACA0FAFC9DCB78	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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
03DC6105FFC8697EFEA1A345FA94CFC9.text	03DC6105FFC8697EFEA1A345FA94CFC9.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Semomesia Westwood 1851	<div><p>Semomesia Westwood, 1851, Mesophthalma Westwood, 1851, Perophthalma Westwood, 1851 and Leucochimona Stichel, 1909</p><p>are junior subjective synonyms of Mesosemia Hübner, [1819]</p></div>	https://treatment.plazi.org/id/03DC6105FFC8697EFEA1A345FA94CFC9	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFC9697FFDA8A2ADFC8FCDB3.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Ectosemia Zhang & Cong & Shen & Opler & Grishin 2021	<div><p>Ectosemia Grishin, new genus</p><p>http://zoobank.org/ 8AB8B269-CAE8-417D-BF0D-E46043B20CBE</p><p>Type species. Papilio eumene Cramer, 1776 .</p><p>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: cne7048.1.3: T2251C, cne3658.2.1:A638T, cne 2957.11.4:A952G, cne3658.2.1:A608C, and cne3461.2.10:C1462A.</p><p>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.</p><p>Species included. The type species, Mesosemia decolorata Lathy, 1932, Mesosemia erinnya Stichel, 1910, and Mesosemia steli Hewitson, 1858 .</p><p>Parent taxon. Subtribe Mesosemiina Bates, 1859 .</p></div>	https://treatment.plazi.org/id/03DC6105FFC9697FFDA8A2ADFC8FCDB3	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFC9697FFDD6A1FAFC8FC85D.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Endosemia Grishin 2021	<div><p>Endosemia Grishin, new genus</p><p>http://zoobank.org/ AA1AC39C-2696-46A8-9927-2A8CDBB14D28</p><p>Type species. Papilio ulrica Cramer, 1777 .</p><p>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: cne703.2.8:T903C, cne 1411.6.4:T1025A, cne2651.14.5: A4602G, cne12205.6.2: G835A, and cne8028.2.1:T1787A.</p><p>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.</p><p>Species included. The type species and Mesosemia macella Hewitson, 1859 .</p><p>Parent taxon. Subtribe Mesosemiina Bates, 1859 .</p></div>	https://treatment.plazi.org/id/03DC6105FFC9697FFDD6A1FAFC8FC85D	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFC96978FDEFA480FABCCE9B.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Eucorna Strand 1932	<div><p>Eucorna Strand, 1932 is a valid genus</p><p>Presently, Eucorna Strand, 1932 (type species Voltinia sanarita (Schaus, 1902)) is a junior subjective synonym of Voltinia Stichel, 1910 (type species Esthemopsis (?) radiata Godman &amp; Salvin, 1886), but is not monophyletic with it, and is sister to all other Napaeina Hall, 2003 except Hyphilaria Hübner, [1819] (type species Hyphilaria nicia Hübner, [1819]) (Fig. 17). Hence, Eucorna is a valid genus.</p><p>Reassessment of Voltinia Stichel, 1910 and Napaea Hübner, [1819]</p></div>	https://treatment.plazi.org/id/03DC6105FFC96978FDEFA480FABCCE9B	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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
03DC6105FFCE6978FF3FA799FB43C8E3.text	03DC6105FFCE6978FF3FA799FB43C8E3.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Hermathena Hewitson 1874	<div><p>Hermathena Hewitson, 1874 is a subgenus of Ithomiola C. &amp; R. Felder, 1865</p><p>Despite its mostly white coloration, much different from its relatives, Hermathena Hewitson, 1874 (type species Hermathena candidata Hewitson, 1874) (Fig. 17 cyan) clusters closely with Ithomiola C. &amp; R. Felder, 1865 (type species Ithomiola floralis C. &amp; R. Felder, 1865) (Fig. 17 green), which even now includes species considerably different in appearance (Hall 2005). The male genitalia of these taxa are rather similar as illustrated by Hall (2005). To achieve a more internally consistent classification, we place Hermathena Hewitson, 1874 as a subgenus of Ithomiola C. &amp; R. Felder, 1865.</p></div>	https://treatment.plazi.org/id/03DC6105FFCE6978FF3FA799FB43C8E3	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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
03DC6105FFCE6979FF06A4CBFF4CCED3.text	03DC6105FFCE6979FF06A4CBFF4CCED3.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Lucillella Strand 1932	<div><p>Lucillella Strand, 1932 is a subgenus of Esthemopsis C. &amp; R. Felder, 1865</p><p>Lucillella Strand, 1932 (type species Lucilla camissa Hewitson, 1870) (Fig. 18 pale blue) and Esthemopsis C. &amp; R. Felder, 1865 (type species Esthemopsis clonia C. &amp; R. Felder, 1865) (Fig. 18 gray) are closely related sisters that form a clade prominently separated from others (Fig. 18) and sister to Mesene Doubleday, 1847 (type species Papilio phareus Cramer, 1777) (Fig. 18 olive). Because Mesene is already more diverse than Lucillella and Esthemopsis combined, to achieve better consistency of classification, we propose treating Lucillella Strand, 1932 as a subgenus of Esthemopsis C. &amp; R. Felder, 1865.</p></div>	https://treatment.plazi.org/id/03DC6105FFCE6979FF06A4CBFF4CCED3	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFCE6978FEADA626FE93CB54.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Voltinia Stichel 1910	<div><p>Ionotus Hall, 2005 and Voltinia Stichel, 1910 are subgenera of Cremna Doubleday, 1847, which is a valid genus</p><p>A genus comprised of two species, Voltinia Stichel, 1910 (type species Esthemopsis (?) radiata Godman &amp; 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.</p></div>	https://treatment.plazi.org/id/03DC6105FFCE6978FEADA626FE93CB54	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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
03DC6105FFCF697AFEFEA319FD82CEEF.text	03DC6105FFCF697AFEFEA319FD82CEEF.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Xynias Hewitson 1874	<div><p>Xynias Hewitson, 1874 is a junior subjective synonym of Mesenopsis Godman &amp; Salvin, 1886, which along with Xenandra C. &amp; R. Felder, 1865 are subgenera of Symmachia Hübner, [1819], that contains Stichelia J. Zikán, 1949 as its junior subjective synonym</p><p>Symmachia, Xenandra, and Mesenopsis . Finally, some species currently in Symmachia fall outside this genus (e.g., Fig. 18 violet and magenta clades). Two of such major clades do not have names and are proposed as new genera here.</p></div>	https://treatment.plazi.org/id/03DC6105FFCF697AFEFEA319FD82CEEF	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFCC697AFDBAA60BFD76C983.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Asymma Grishin 2021	<div><p>Asymma Grishin, new genus</p><p>http://zoobank.org/ 901035D7-E6D1-4F16-A1FD-BA85C48AAC50</p><p>Type species. Symmachia virgatula Stichel, 1910 .</p><p>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: cne4291.7.6: A1077G, cne4291.7.6:A1064G, cne4291.7.6:A1051G, cne3461.2.5:A1310G, and cne3461.1.15:A3342C.</p><p>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].</p><p>Species included. The type species, Synapta arion C. &amp; R. Felder, 1865, Symmachia giffordi P. Jauffret &amp; J. Jauffret, 2010, Symmachia hippodice Godman, 1903, Symmachia pardalis Hewitson, 1867, Symmachia satana J. Hall &amp; Harvey, 2007, Symmachia stigmosissima Stichel, 1910, and Symmachia virgaurea Stichel, 1910 .</p><p>Parent taxon. Tribe Symmachiini Reuter, 1896 .</p></div>	https://treatment.plazi.org/id/03DC6105FFCC697AFDBAA60BFD76C983	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFCC697AFD89A33EFD76CA21.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Tigria Zhang & Cong & Shen & Opler & Grishin 2021	<div><p>Tigria Grishin, new genus</p><p>http://zoobank.org/ 1665C4A1-7AF7-4426-855A-1799CC9F59E4</p><p>Type species. Mesene xypete Hewitson, 1870 .</p><p>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:A2889G, cne3461.1.14: G922C, cne3437.1.9:A1260G, cne3461.1.14:A846G, and cne 2170.2.1:T2751C.</p><p>Etymology. The name is a feminine noun in the nominative singular given for the red-orange tiger-striped appearance of these species.</p><p>Species included. The type species, Polystichtis rubrica Stichel, 1929, Cricosoma phaedra Bates, 1868 and Metacharis elinas Rebillard, 1958 .</p><p>Parent taxon. Tribe Symmachiini Reuter, 1896 .</p></div>	https://treatment.plazi.org/id/03DC6105FFCC697AFD89A33EFD76CA21	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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
03DC6105FFCD697BFF6DA107FAB5C82B.text	03DC6105FFCD697BFF6DA107FAB5C82B.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Alethea Nielsen & Salazar 2018	<div><p>Alethea Nielsen &amp; Salazar, [2018] is a junior subjective synonym of Pirascca J. Hall &amp; Willmott, 1996, which is a subgenus of Pterographium Stichel, 1910, that contains Panaropsis J. Hall, 2002 as its junior subjective synonym</p><p>The monotypic Alethea Nielsen &amp; Salazar, [2018] (type and the only species Siseme pedias Godman, 1903) (Fig. 18 dark blue), Panaropsis J. Hall, 2002 (type species Panara elegans Schaus, 1920) (Fig. 18 cyan-green), Pirascca J. Hall &amp; Willmott, 1996 (type species Papilio sagaris Cramer, 1775) (Fig. 18 green) and monotypic Pterographium Stichel, 1910 (type and the only species Pterographium aphaniodes Stichel, 1910, which is Panara sicora Hewitson, 1875) (Fig. 18 brown) taken together form a prominent clade in the tree that originated early in the diversification of Symmachiini Reuter, 1896 into genera and therefore corresponds to genus rank. This genus gets the name Pterographium as the oldest available for its species. This species-rich Pterographium sensu lato splits into 2 well-defined and strongly supported but not very prominent clades (100% support for each clade, Fig. 18) that we treat as subgenera. The nominal subgenus (upper clade in Fig. 18) contains Panaropsis as sister to its type species, and therefore Panaropsis becomes a junior subjective synonym of Pterographium, along with some other species such as Phaenochitona [sic] interrupta Lathy, 1932 (currently in Pirascca). The second subgenus of Pterographium (lower clade in Fig. 18) is Pirascca, a close sister to monotypic and unusually patterned Alethea, which we place as junior subjective synonym of Pirascca due to genetic similarities. We reason that considerable phenotypic differences of Pterographium (Pirascca) pedias (Godman, 1903) comb. nov. from other Pirascca are caused by rapid evolution towards a different mimicry complex.</p></div>	https://treatment.plazi.org/id/03DC6105FFCD697BFF6DA107FAB5C82B	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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
03DC6105FFCD697BFE07A25EFE12CFA3.text	03DC6105FFCD697BFE07A25EFE12CFA3.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Chimastrum Godman & Salvin 1886	<div><p>Chimastrum Godman &amp; Salvin, 1886 is a junior subjective synonym of Mesene Doubleday, 1847</p><p>Chimastrum Godman &amp; Salvin, 1886 (type species Mesene argentea Bates, 1866) (Fig. 18 red) originates within Mesene Doubleday, 1847 (type species Papilio phareus Cramer, 1777) (Fig. 18 olive), thus rendering it paraphyletic. To restore the monophyly, instead of breaking Mesene (that is currently already assembled from close relatives) into several genus-group taxa, we treat Chimastrum as a junior subjective synonym of Mesene .</p></div>	https://treatment.plazi.org/id/03DC6105FFCD697BFE07A25EFE12CFA3	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFCD697BFE2EA389FBA6CCD6.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Mesene aeolia (Bates 1868) Zhang & Cong & Shen & Opler & Grishin 2021	<div><p>Mesene aeolia (Bates, 1868), new combination</p><p>Kept in Esthemopsis C. &amp; R. Felder, 1865 (type species Esthemopsis clonia C. &amp; R. Felder, 1865) since its description, E. aeolia Bates, 1868</p><p>(type locality Brazil: Para) is not monophyletic with it, and instead is sister to Mesene argentea Bates, 1866, the type species of Chimastrum</p><p>Godman &amp; Salvin, 1886 that originates within Mesene Doubleday, 1847</p><p>(type species Papilio phareus Cramer, 1777) (Fig. 18, sequenced specimen in Fig. 19), implying Mesene aeolia (Bates, 1868) comb. nov.</p></div>	https://treatment.plazi.org/id/03DC6105FFCD697BFE2EA389FBA6CCD6	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFCD697BFEB5A473FE4DC982.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Pterographium hypochloris (Bates 1868) Zhang & Cong & Shen & Opler & Grishin 2021	<div><p>Pterographium hypochloris (Bates, 1868), new combination</p><p>Currently in Symmachia Hübner, [1819] (type species Symmachia probetrix Hübner, [1819], which is Papilio probetor Stoll, 1782), Emesis hypochloris Bates, 1868 (type locality Brazil: Amazonas) is not monophyletic with it and instead originates within Pterographium Stichel,</p><p>1910 (type species Pterographium aphaniodes Stichel, 1910, which is</p><p>Panara sicora Hewitson, 1875) sensu lato as it is defined above (Fig. 18,</p><p>sequenced specimen shown in Fig. 20), hence Pterographium hypochloris</p><p>(Bates, 1868) comb. nov.</p></div>	https://treatment.plazi.org/id/03DC6105FFCD697BFEB5A473FE4DC982	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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
03DC6105FFC26974FE97A2ADFCE2CF94.text	03DC6105FFC26974FE97A2ADFCE2CF94.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Comphotis Stichel 1910	<div><p>Comphotis Stichel, 1910 is a junior subjective synonym of Phaenochitonia Stichel, 1910</p><p>Comphotis Stichel, 1910 (type species Cricosoma irroratum Godman, 1903) (Fig. 18 pale orange) is paraphyletic with respect to Phaenochitonia Stichel, 1910 (type species Papilio cingulus Stoll, 1790) (Fig. 18 bright cyan), but taken together, they form a prominent lineage consistent with other Symmachiini Reuter, 1896 genera. Therefore we place Comphotis Stichel, 1910 -XII-31 as a junior subjective synonym of Phaenochitonia Stichel, 1910 -IX.</p></div>	https://treatment.plazi.org/id/03DC6105FFC26974FE97A2ADFCE2CF94	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFC26974FEDAA7CCFD71C979.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Imelda Hewitson 1870	<div><p>Imelda Hewitson, 1870 is a subgenus of Echenais Hübner, [1819]</p><p>Despite the differences in appearance, monotypic genus</p><p>Echenais Hübner, [1819] (type species Lemonias alphaea</p><p>Hübner, [1808], which is Papilio thelephus Cramer, 1775)</p><p>is genetically close to Imelda Hewitson, 1870 (type species</p><p>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).</p><p>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] .</p></div>	https://treatment.plazi.org/id/03DC6105FFC26974FEDAA7CCFD71C979	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFC26974FE8DA3D9FE3ACBE1.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Phaenochitonia florus (Fabricius 1793) Zhang & Cong & Shen & Opler & Grishin 2021	<div><p>Phaenochitonia florus (Fabricius, 1793), new combination</p><p>Currently in Mesene Doubleday, 1847 (type species Papilio phareus Cramer,</p><p>1777), Hesperia florus Fabricius, 1793 (type locality " Indiis ", likely SE Brazil)</p><p>is not monophyletic with it and originates within Phaenochitonia Stichel, 1910</p><p>(type species Papilio cingulus Stoll, 1790) (Fig. 18 yellow, highlighted in violet,</p><p>sequenced specimen shown in Fig. 21), implying Phaenochitonia florus</p><p>(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.</p><p>Reassessment of Ourocnemis Bethune-Baker, 1887 and Anteros Hübner, [1819]</p><p>The tree of Helicopini Stichel, 1928 revealed a distribution of species between the two genera Ourocnemis Bethune-Baker, 1887 (type species</p><p>Anteros axiochus Hewitson, 1867) and Anteros</p><p>Hübner, [1819] (type species Papilio formosus Cramer,</p><p>1777) not as presently attributed (Fig. 22). A number of species placed in Anteros were in the same clade with</p><p>Ourocnemis, confirming the suspicion of Hall (1998)</p><p>that Anteros may be paraphyletic. Using this phylogenetic result and phenotypic similarities (cream</p><p>vs. dark background of wings below) for species with missing DNA data, we transfer four species from</p><p>Anteros to Ourocnemis to form the following new combinations: Ourocnemis carausius (Westwood, 1851), Ourocnemis principalis (Hopffer, 1874), Ourocnemis renaldus (Stoll, 1790), and Ourocnemis aerosus (Stichel, 1924) .</p></div>	https://treatment.plazi.org/id/03DC6105FFC26974FE8DA3D9FE3ACBE1	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFC36975FF52A25EFDDECC31.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Colaciticus Stichel 1910	<div><p>Colaciticus Stichel, 1910 is junior subjective synonym of Baeotis Hübner, [1819]</p><p>Despite different coloration and wing shape, which are probably due to mimicry, Colaciticus Stichel, 1910</p><p>(type species Monethe johnstoni Dannatt, 1904)</p><p>originates deep within Baeotis Hübner, [1819] (type species Baeotis hisbaena Hübner, [1819] which is</p><p>Papilio hisbon Cramer, 1775) (Fig. 24). Baeotis is a well-defined prominent genus that we do not wish to split. Therefore, to restore the monophyly, we propose that Colaciticus Stichel, 1910 is a junior subjective synonym of Baeotis Hübner, [1819] .</p></div>	https://treatment.plazi.org/id/03DC6105FFC36975FF52A25EFDDECC31	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFC36975FF70A07DFD34C9B9.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Nahida coenoides (Hewitson 1870)	<div><p>Nahida coenoides (Hewitson, 1870) is conspecific with Ithomeis aurantiaca H. Bates, 1862 and Nahida Kirby, 1871 is a junior subjective synonym of Ithomeis Bates, 1862</p><p>Putridivora Grishin, new genus http://zoobank.org/ AD6A1874-503C-4A07-B079-79A326060635</p><p>Type species. Charis argyrea Bates, 1868 .</p><p>Definition. Currently within Detritivora Hall &amp; Harvey, 2002 (type species Charis matic Harvey &amp; 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 &amp; 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. 6a.</p><p>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.</p><p>Species included. The type species and Charis smalli Hall &amp; Harvey, 2001 .</p><p>Parent taxon. Tribe Riodinini Grote, 1895 .</p></div>	https://treatment.plazi.org/id/03DC6105FFC36975FF70A07DFD34C9B9	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFC06976FDB0A25EFD34CCEF.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Chadia Grishin 2021	<div><p>Chadia Grishin, new genus</p><p>http://zoobank.org/ 10295164-45F5-4C4E-8323-FBC0D37E802D</p><p>Type species. Charis cadytis Hewitson, 1866 .</p><p>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 &amp; 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.</p><p>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 .</p><p>Species included. Only the type species.</p><p>Parent taxon. Tribe Riodinini Grote, 1895 .</p></div>	https://treatment.plazi.org/id/03DC6105FFC06976FDB0A25EFD34CCEF	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFC06977FDB0A13EFC45CDB5.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Inkana Grishin 2021	<div><p>Inkana Grishin, new genus</p><p>http://zoobank.org/ 4FEC8FF0-3B35-4B6A-BEC4-5CF06240CA5C</p><p>Type species. Charis incoides Schaus, 1902 .</p><p>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. &amp; 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 &amp; Willmott, 1998 and Lasaia incoides on pages 23–24 and illustrated in Figs. 3–4, 9–10 by Hall &amp; 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.</p><p>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.</p><p>Species included. The type species and Lasaia cutisca Hall &amp; Willmott, 1998 .</p><p>Parent taxon. Tribe Riodinini Grote, 1895 .</p><p>Callistiumini Grishin, new tribe</p><p>http://zoobank.org/ 3BFAAAD2-83E4-48EF-9952-CA85DA8DEB9E</p><p>Type genus. Callistium Stichel, 1911 .</p><p>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 &amp; 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.</p></div>	https://treatment.plazi.org/id/03DC6105FFC06977FDB0A13EFC45CDB5	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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
03DC6105FFC16977FE91A431FEECC9A8.text	03DC6105FFC16977FE91A431FEECC9A8.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Hallonympha maculosa (Bates 1868) Zhang & Cong & Shen & Opler & Grishin 2021	<div><p>Hallonympha maculosa (Bates, 1868), new combination</p><p>Although synonymized with Zabuella Stichel, 1911 (type species Lemonias tenellus Burmeister, 1878) by Hall (2018), Hallonympha Penz &amp; DeVries, 2006 (type species Apodemia paucipuncta Spitz, 1930) is genetically distant from it (Fig. 28), in agreement with Seraphim et al. (2018). E.g., COI barcodes of the type species of these genera differ by 9.1% (60 bp) and their divergence dates to about 20 Mya (Seraphim et al. 2018). Therefore, we reinstate Hallonympha as a valid genus. Currently in Callistium (type species Charis cleadas Hewitson, 1866), Calydna maculosa Bates, 1868 (type locality Brazil: Amazonas) is not monophyletic with its type species and instead is sister to the type species of Hallonympha (Fig. 28). Their COI barcodes are only 7.4% (49 bp) different. Hence, we propose Hallonympha maculosa (Bates, 1868) comb. nov.</p></div>	https://treatment.plazi.org/id/03DC6105FFC16977FE91A431FEECC9A8	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFC16977FF45A1F8FAE6CBBB.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Zabuella castanea (Prittwitz 1865)	<div><p>Zabuella castanea (Prittwitz, 1865),</p><p>confirmed combination</p><p>Placed in Callistium (type species Charis cleadas</p><p>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</p><p>Lemonias tenellus Burmeister, 1878 (Fig. 28),</p><p>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</p><p>(magenta</p><p>: Behemothia</p><p>), and Teenie</p><p>(magenta gen. n</p><p>with</p><p>. (orange subgenus</p><p>); and</p><p>Calicosama</p><p>Theopina same conclusion, and confirm the combination in cyan), Archaeonympha (black), Theope (blue), Zabuella castanea (Prittwitz, 1865) . Pseudotinea (black), and Petrocerus (red).</p></div>	https://treatment.plazi.org/id/03DC6105FFC16977FF45A1F8FAE6CBBB	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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
03DC6105FFC66970FF0AA1DFFD56CAF8.text	03DC6105FFC66970FF0AA1DFFD56CAF8.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Calicosama J. Hall & Harvey 2001	<div><p>Calicosama J. Hall &amp; Harvey, 2001 is a subgenus of Behemothia Hall, 2000</p><p>Monotypic genus Behemothia Hall, 2000 (type and the only species Pandemos godmanii Dewitz, 1877) is closely related to nearly monotypic Calicosama J. Hall &amp; Harvey, 2001 (type species Nymphidium lilina Butler, 1870) (Fig. 28). Both genera are mostly Central American. To highlight their close kinship and reduce the number of unnecessarily monotypic genera we propose that Calicosama J. Hall &amp; Harvey, 2001 is a subgenus of Behemothia Hall, 2000 .</p></div>	https://treatment.plazi.org/id/03DC6105FFC66970FF0AA1DFFD56CAF8	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFC66970FE74A725FAE1C86F.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Exoplisia aphanis (Stichel 1910) Zhang & Cong & Shen & Opler & Grishin 2021	<div><p>Exoplisia aphanis (Stichel, 1910), new combination</p><p>Placed in Seco Hall &amp; Harvey, 2002 (type species Charis calagutis Hewitson, 1871), Charmona aphanis Stichel, 1910</p><p>is not monophyletic with its type species, but is sister to</p><p>Exoplisia Godman &amp; Salvin, 1886 (type species Amarynthis hypochalybe C. &amp; R. Felder, 1861) and is not prominently distinct from it (Fig. 29). Therefore, we place it in this genus Fig. 29. Seco (purple), Oco gen. n. (magenta), to form Exoplisia aphanis (Stichel, 1910) comb. nov. Exoplisia (cyan and orange).</p></div>	https://treatment.plazi.org/id/03DC6105FFC66970FE74A725FAE1C86F	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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
03DC6105FFC66970FEADA0D2FA20CD8F.text	03DC6105FFC66970FEADA0D2FA20CD8F.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Petrocerus Callaghan 1979	<div><p>Petrocerus Callaghan, 1979 belongs to Theopina Clench, 1955</p><p>Currently in incertae sedis, Petrocerus Callaghan, 1979 (type species Calydna catiena Hewitson, 1875) is a confident sister to Pseudotinea Hall &amp; Callaghan, 2003 (type species Calydna volcanicus Callaghan &amp; Salazar, 1997) (Fig. 28). Together they are sister to the clade with Theope Doubleday, 1847 (type species Polyommatus terambus Godart, [1824]), and therefore, Petrocerus belongs to Theopina Clench, 1955.</p></div>	https://treatment.plazi.org/id/03DC6105FFC66970FEADA0D2FA20CD8F	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFC66970FDB5A25EFBAECC8A.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Teenie Grishin 2021	<div><p>Teenie Grishin, new genus</p><p>http://zoobank.org/ BDF61309-B594-4355-B2ED-306025A2E8D9</p><p>Type species. Calydna tinea Bates, 1868 .</p><p>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 &amp; 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.</p><p>Etymology. The name is a feminine noun in the nominative singular inspired by the type species name.</p><p>Species included. The type species (including Lemonias eudocia Godman &amp; Salvin, 1897 as it synonym) and Calydna argiella Bates, 1868 .</p><p>Parent taxon. Subtribe Zabuellina Seraphim, Freitas &amp; Kaminski, 2018 .</p></div>	https://treatment.plazi.org/id/03DC6105FFC66970FDB5A25EFBAECC8A	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFC66971FD99A4B2FD34CF4D.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Oco Zhang & Cong & Shen & Opler & Grishin 2021	<div><p>Oco Grishin, new genus</p><p>http://zoobank.org/ 1D2F2454-EFC9-473A-A91D-17097BB3DB03</p><p>Type species. Symmachia ocellata Hewitson, 1867 .</p><p>Definition. Currently in the genus Seco Hall &amp; 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</p><p>(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.</p><p>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.</p><p>Species included. Only the type species.</p><p>Parent taxon. Tribe Riodinini Grote, 1895 .</p></div>	https://treatment.plazi.org/id/03DC6105FFC66971FD99A4B2FD34CF4D	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFC76971FDCBA390FCD5CD2B.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Machaya Hall & Willmott 1995	<div><p>Machaya Hall &amp; Willmott, 1995 is a junior subjective synonym of Pachythone Bates, 1868</p><p>Despite the difference in appearance, Machaya Hall &amp;</p><p>Willmott, 1995 (type species Machaya obstinata Hall &amp;</p><p>Willmott, 1995) originates deep within Pachythone Bates,</p><p>1868 (type species Pachythone erebia Bates, 1868) at the time of rapid diversification of its crown group (Fig. 30).</p><p>Therefore, to restore the monophyly, we propose that</p><p>Machaya is a junior subjective synonym of Pachythone .</p></div>	https://treatment.plazi.org/id/03DC6105FFC76971FDCBA390FCD5CD2B	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFC7694CFE5AA4D4FD7DC891.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Siderone Hubner 1823	<div><p>Siderone Hübner, [1823] and Phantos Dias, 2018 are subgenera of Zaretis Hübner, [1819]</p><p>Genetic differentiation within the clade of three genera: Siderone Hübner, [1823] (type species Siderone ide Hübner, [1823], a junior subjective synonym of Papilio nemesis Illiger, 1801), Phantos Dias, 2018 (type species Nymphalis callidryas R. Felder, 1869) and Zaretis Hübner, [1819] (type species Papilio isidora Cramer, 1779) is less than that of Memphis and Anaea (sensu lato, as above) (Fig. 31), suggesting that Siderone and Phantos are subgenera of Zaretis to restore internal consistency of the classification. According to Fig. 3 in Toussaint et al. (2019), all non-monotypic genera of Anaeini as we define them ( Anaea, Memphis, Consul and Zaretis) diversified around the same time 17-20 Mya, indicating internal consistency of our definition; and monotypic genera Coenophlebia C. &amp; R. Felder, 1862 (type and the only species Siderone archidona Hewitson, 1860, we have not sequenced this species yet) and Hypna Hübner, [1819] (type species Papilio clytemnestra Cramer, 1777) are more distant from others to maintain their distinction.</p><p>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</p><p>Listed among the "Subtribe uncertain" genera by Wahlberg (2019a), Paralasa Moore, 1893 (type species Erebia kalinda Moore, 1865) is a confident sister to Ypthimina Reuter, 1896 (Fig. 32). Therefore, instead of proposing a monotypic subtribe for this genus, we include Paralasa in the subtribe Ypthimina .</p><p>Callerebiina Grishin, new subtribe</p><p>http://zoobank.org/ 46A68C99-BBC2-4F6C-BB06-4CA2AD2343B8</p><p>Type genus. Callerebia Butler, 1867 .</p><p>Definition. Several genera of uncertain tribal placement (Wahlberg 2019a) confidently grouped with others sometimes placed in Ypthimina Reuter, 1896 (Fig. 32 red). Close relationship of Callerebia, Loxerebia and Argestina that are in the red clade has been reported before (Yang and Zhang 2015). This prominent red clade is indeed a weakly supported sister to Ypthimina (Fig. 32 blue and cyan). However, due to the weak statistical support (therefore possibly erroneous sister relationship with Ypthimina) and the origin of both clades near the rapid radiation of many Satyrini subtribes, this clade is proposed as a new subtribe. A phenotypically diverse assembly of species, generally characterized by Erebia -like appearance, typically with a row of orange black-centered eyespots on each wing, but these could be reduced to a couple of eyespots, similar to Ypthimina, or wings could be unspotted brown or even mostly white above (with brown borders) in some species; gnathos developed (absent in many Ypthimina), but arms shorter than uncus, saccus reduced, valva rather stout, flattened and rounded or excavated near the apex, aedeagus rather short, shorter than valva, typically bent and twisted, boomerang-shaped differentiating the subtribe from Erebia Dalman, 1816 . Due to phenotypic diversity, best diagnosed by the DNA characters in the nuclear genome: hm2009277-RA.13:T2076C, hm2009277-RA.13:A130C, hm2015715-RA.2:C2137T, hm2009379-RA.4:T1915C, ahm2002906-RA.2:A128G, where the part before the first dot (e.g. hm2009277-RA) is the protein ID, next number (e.g. 13) is exon of the Heliconius melpomene genome assembly (Davey et al. 2016), and combination like A130C means base pair C at position 130, changed from A in the ancestor. See &lt;https://osf.io/kj4es/&gt; for the sequences of these exons.</p><p>Genera included. The type genus, Proterebia Roos &amp; Arnscheid, 1980, Argestina Riley, 1923, Loxerebia Watkins, 1925, and Physcaeneura Wallengren, 1857 .</p><p>Parent Taxon. Tribe Satyrini Boisduval, 1833 .</p><p>Gyrocheilina Grishin, new subtribe</p><p>http://zoobank.org/ EF7C543A-88B9-4DCC-9847-139CC7763B84</p><p>Type genus. Gyrocheilus Butler, 1867 .</p><p>Definition. Previously placed in Pronophilina Reuter, 1896 (Wahlberg 2019a), but is not monophyletic with it. Instead, it originates in early radiation of the clade that is sister to Satyrina Boisduval, 1833 (Figs. 32, 33 orange), likely prior to divergence of subtribes Ypthimina Reuter, 1896 and Erebiina Tutt, 1896, and therefore is a subtribe. Diagnosed by the combination of the following characters: most prominently, female foreleg is much reduced, with 2 tarsal subsegment, not spined; then forewings apically rounded with somewhat undulate outer margin, hindwing with strongly undulate, almost toothed outer margins; forewing discal cell short and broad: slightly longer than half of wing, width ⅓–½ of its length; forewing vein R 1 starts at distal ¼ of discal cell, R 2 very near discal cell apex (only slightly stalked), R 3 and R 4 stalked for ⅓ of their length, R 4 and R 5 stalked for about half of their length, recurrent vein in discal cell from the middle of discocellular vein for ¼ of discal cell length, discocellular vein straight between R 2 and M 1 and between M 2 and M 3 origins, but V-shaped between M 1 and M 2 origins; forewing with 4 (rarely 3) white-centered round eyespots, hindwing without such eyespots but frequently with several cream-colored ovals or crescents; palpi long and porrect, scales on the second segment long, up to 5 times the width of the segment, eyes bare, antennae shorter than half of forewing; pupa suspended, spindleshaped, smooth, head capsule apically extended similar to cremaster, forked at the tip.</p><p>Genera included. Only the type genus.</p><p>Parent Taxon. Tribe Satyrini Boisduval, 1833 .</p><p>Calistina Grishin, new subtribe</p><p>http://zoobank.org/ F1D9A7DC-F549-4EC9-8C68-D8032DC4FB95</p><p>Type genus. Calisto Hübner, [1823] .</p><p>Genera included. Only the type genus.</p><p>Parent Taxon. Tribe Satyrini Boisduval, 1833 .</p><p>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.</p></div>	https://treatment.plazi.org/id/03DC6105FFC7694CFE5AA4D4FD7DC891	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFFA694DFF72A4DCFA3FCE8A.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Harsiesis Fruhstorfer 1911	<div><p>Harsiesis Fruhstorfer, 1911 is a subgenus of Platypthima Rothschild &amp; Jordan, 1905</p><p>Harsiesis Fruhstorfer, 1911 (type species Hypocista [sic] hygea Hewitson, 1863) and Platypthima Rothschild &amp; Jordan, 1905 (type species Platypthima ornata Rothschild &amp; Jordan, 1905) are characterized by an elevated evolutionary rate as revealed by their longer branches (Fig. 34). Despite their faster evolution, the two are more genetically similar to each other than other pairs of sister genera, e.g.,</p><p>COI barcodes between their type species (GenBank accession GQ357203 for P. ornata) differ by 9.4% (62 bp). Therefore Harsiesis Fruhstorfer, 1911 is a subgenus of Platypthima Rothschild &amp; Jordan, 1905.</p></div>	https://treatment.plazi.org/id/03DC6105FFFA694DFF72A4DCFA3FCE8A	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFFB694DFF3BA2D3FC67CCB1.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Percnodaimon Butler 1876	<div><p>Percnodaimon Butler, 1876 and Erebiola Fereday, 1879 are junior subjective synonyms of Argyrophenga Doubleday, 1845</p><p>Two monotypic genera Erebiola Fereday, 1879 (type and the only species Erebiola butleri Fereday, 1879) and Percnodaimon Butler, 1876 (type and the only species Percnodaimon pluto Butler, 1876,</p><p>homonym, valid name Erebia merula Hewitson,</p><p>1875) cluster closely with Argyrophenga Doubleday,</p><p>1845 (type species Argyrophenga antipodum</p><p>Doubleday, 1845) (Fig. 34), all being from New Fig. 34. Platypthima (magenta) and Argyrophenga (blue).</p><p>Zealand and phenotypically similar. Therefore, Percnodaimon Butler, 1876 and Erebiola Fereday, 1879 are junior subjective synonyms of Argyrophenga Doubleday, 1845 .</p></div>	https://treatment.plazi.org/id/03DC6105FFFB694DFF3BA2D3FC67CCB1	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFFB694FFDAAA0FCFA10CF1D.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Magda Grishin 2021	<div><p>Magda Grishin, new subgenus</p><p>http://zoobank.org/ 4AF74BDC-1FCD-42B6-8BFB-A316F0F09D93</p><p>Type species. Erebia magdalena Strecker, 1880 .</p><p>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: 271C (not T), 274C (not T), 421T or C (not A), 424T (not A), 451T or C (not A), 484not T, and 622T (not A).</p><p>Etymology. The name is a feminine noun in the nominative singular, formed from the type species name.</p><p>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 .</p><p>Parent taxon. Genus Erebia Dalman, 1816 .</p><p>Additional genomic support for the broadly defined genera Cercyonis Scudder, 1875, Maniola Schrank, 1801 and Erebia Dalman, 1816</p><p>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.</p></div>	https://treatment.plazi.org/id/03DC6105FFFB694FFDAAA0FCFA10CF1D	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFF9694FFF58A340FD3FCCF5.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Hestinalis Bryk 1938	<div><p>Hestinalis Bryk, 1938 is a junior subjective synonym of Mimathyma Moore, 1896</p><p>In the genomic tree Hestinalis Bryk, 1938 (type species Hestina mimetica Butler, 1874) originates within Mimathyma Moore, 1896 (type species Athyma chevana</p><p>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 .</p></div>	https://treatment.plazi.org/id/03DC6105FFF9694FFF58A340FD3FCCF5	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFF96948FF6CA6C7FB67CDF4.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Biblis aganisa Boisduval 1836	<div><p>Biblis aganisa Boisduval, 1836 is a species distinct from Biblis hyperia (Cramer, 1779)</p><p>Genetic diversification between the nominotypical Biblis (Biblis) hyperia (Cramer, 1779) (type locality St. Thomas) and Biblis (Biblis) hyperia aganisa Boisduval, 1836 (type locality inferred as Mexico) is at the level of that between two species from the subgenus Vila: Biblis (Vila) azeca (Doubleday, [1848]) and Biblis (Vila) eueidiformis (Joicey &amp; Talbot, 1918) (Fig. 38). The COI barcodes of the two taxa differ by 4.6% (30 bp). The nominotypical B. hyperia is characterized by more prominent dorsal forewing marginal bands than B. b. aganisa . Taken together, these arguments suggest that Biblis aganisa Boisduval, 1836 reinstated status is a species distinct from Biblis hyperia (Cramer, 1779) .</p><p>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</p><p>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 (2019a), 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. &amp; R. Felder, 1861, a subspecies of Catagramma vaninka Hewitson, 1855)</p><p>and Orophila Staudinger, 1886 (type species Cybelis campaspe Hewitson, 1869, a subspecies of Cybdelis cardases Hewitson, 1869) cluster closely together (Fig.</p><p>39). Their COI barcodes also indicate a close relationship, e.g., the type species of Mesotaenia and</p><p>Perisama are only 6.2% (41 bp) different. Therefore, Fig. 39. Callicorina genera: Lucinia (black), Haematera (red: subgenus Antigonis, and cyan),</p><p>we agree with Wahlberg (2019a) and confirm that Catagramma (olive &amp; pink), and Callicore (purple &amp; Mesotaenia Kirby, 1871 and Orophila Staudinger, 1886 yellow: subgenus Diaethria, magenta &amp; green:</p><p>are junior subjective synonyms of Perisama Doubleday, subgenus Perisama, and blue).</p><p>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 (2019a) 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).</p></div>	https://treatment.plazi.org/id/03DC6105FFF96948FF6CA6C7FB67CDF4	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFFE6948FE08A60EFABAC89C.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Callicore Hubner 1819	<div><p>are subgenera of Callicore Hübner, [1819]</p><p>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).</p></div>	https://treatment.plazi.org/id/03DC6105FFFE6948FE08A60EFABAC89C	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFFE6949FEE8A4C0FA2FCE8A.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Antigonis C. Felder 1861	<div><p>Antigonis C. Felder, 1861 is a subgenus of Haematera Doubleday, 1849</p><p>A monotypic genus Antigonis C. Felder, 1861 (type and the only species Cybdelis pharsalia Hewitson, 1852) is rather closely related to another monotypic genus Haematera Doubleday, 1849 (type species Haematera thysbe Doubleday, 1849, which is a subspecies of Callidula pyrame Hübner, [1819]) (Fig.</p><p>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.</p></div>	https://treatment.plazi.org/id/03DC6105FFFE6949FEE8A4C0FA2FCE8A	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFFF6949FEC2A2D3FE0ECDFC.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Asterope Hubner 1819	<div><p>Asterope Hübner, [1819], Nica Hübner, [1826], Peria Kirby, 1871, and Callicorina Smart, 1976 are subgenera of Temenis Hübner, [1819]</p><p>We obtained and analyzed genomic data for the type species of all 10 available genus-group names of the subtribe Epiphilina Jenkins, 1987 that resulted in a well-resolved phylogeny (Fig. 40). We find that</p><p>Temenis Hübner, [1819] (type species Papilio merione</p><p>Fabricius, 1781, which is Papilio laothoe Cramer,</p><p>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,</p><p>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 .</p></div>	https://treatment.plazi.org/id/03DC6105FFFF6949FEC2A2D3FE0ECDFC	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFFF6944FEDEA620FA58C995.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Panacea Godman & Salvin 1883	<div><p>Panacea Godman &amp; Salvin, 1883 is a junior subjective synonym of Batesia C. Felder &amp; R. Felder, 1862</p><p>Introductory comments on Nymphalinae</p><p>Five genera from the subfamily Nymphalinae Rafinesque, 1815 were placed as incertae sedis by Wahlberg (2019a): Pycina Doubleday 1849 (type and the only species Pycina zamba Doubleday, [1849]), Rhinopalpa C. &amp; R. Felder 1860 (type and the only species Rhinopalpa fulva C. &amp; R. Felder, 1860, a junior subjective synonym of Vanessa eudoxia Guérin-Méneville, 1840, with is a subspecies of Papilio polynice Cramer, 1779)), Kallimoides Shirôzu &amp; 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. &amp; R. Felder 1860 (type species Papilio bisaltide Cramer, 1777). The reasons behind the incertae sedis placement, such as ancient origin of these taxa forming long branches in the trees, were given by Wahlberg (2019b), who concluded: "Clearly more data are needed for these ancient taxa." To shed light on their classification, we obtained whole genome shotgun for the type species of these five genera and placed them in the phylogenetic context of other Nymphalinae . The tree constructed from protein-coding genes of the Z chromosome is illustrated (Fig. 42). In this mostly well-supported phylogeny, all currently defined tribes receive 100% statistical support. The five taxa in question are found in deeper radiation near the origins of Nymphalinae and are not closely associated with any of the currently defined tribes. Pycina zamba and Rhinopalpa polynice are confidently placed prior to divergence of other tribes such as Victorinini Scudder, 1893 and Junoniini Reuter, 1896 (Fig. 42). Doleschallia bisaltide is a confident sister to Melitaeini Herrich-Schäffer, 1843, but is prominently separated from them, originating well prior to radiation of Melitaeini into subtribes. Notably, Melitaeini are characterized by an elevated evolutionary rate reflected in much longer branches within the tribe compared to those of most other Nymphalinae, including Doleschallia . Finally, although Kallimoides rumia and Vanessula milca are confidently placed in the clade consisting of Victorinini, Junoniini, Melitaeini and Doleschallia, their exact phylogenetic position remains weakly supported because they originated during a rapid radiation event near the last common ancestor of this clade. It is likely that K. rumia is a distant sister of Victorinini. However, the association of V. milca with Junoniini is questionable. It is not likely that the lack of confidence in this phylogenic placement is caused by the shortage of sequence data. It is plausible that these taxa are of hybrid origin, or their evolution involved incomplete lineage sorting resulting in phylogenetic incongruence among different genes. Regardless of their exact phylogenetic origins, and exactly due to this phylogenetic uncertainty, these taxa do not belong to any existing tribes. For all the reasons stated above, each of these five lineages represents a tribe of Nymphalinae, as named below.</p><p>Pycinini Grishin, new tribe</p><p>http://zoobank.org/ D5C0AE41-2615-4852-9B59-8533C27E4798</p><p>Type genus. Pycina Doubleday 1849 .</p><p>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 &amp; 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.</p><p>Genera included. Only the type genus.</p><p>Parent Taxon. Subfamily Nymphalinae Rafinesque, 1815 .</p><p>Rhinopalpini Grishin, new tribe</p><p>http://zoobank.org/ 4A3ACC99-A761-49B9-BA00-00F83FF443DE</p><p>Type genus. Rhinopalpa C. &amp; R. Felder 1860 .</p><p>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.</p><p>Genera included. Only the type genus.</p><p>Parent Taxon. Subfamily Nymphalinae Rafinesque, 1815 .</p><p>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).</p><p>Kallimoidini Grishin, new tribe</p><p>http://zoobank.org/ 22C74A4F-6B24-4502-99BB-B8E3C66AFABE</p><p>Type genus. Kallimoides Shirôzu &amp; Nakanishi 1984 .</p><p>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 &amp; 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.</p><p>Genera included. Only the type genus.</p><p>Parent Taxon. Subfamily Nymphalinae Rafinesque, 1815 .</p><p>Vanessulini Grishin, new tribe</p><p>http://zoobank.org/ D584B74C-8DC8-479A-AD25-9197A03A843B</p><p>Type genus. Vanessula Dewitz 1887 .</p><p>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 .</p><p>Genera included. Only the type genus.</p><p>Parent Taxon. Subfamily Nymphalinae Rafinesque, 1815 .</p><p>Doleschalliaini Grishin, new tribe</p><p>http://zoobank.org/ 88D32044-DB8E-486F-8C09-0D188E5CDC93</p><p>Type genus. Doleschallia C. &amp; R. Felder 1860 .</p><p>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 .</p><p>Genera included. Only the type genus.</p><p>Parent Taxon. Subfamily Nymphalinae Rafinesque, 1815 .</p><p>Comments. Doleschalla Walker 1861 ( Diptera: Tachinidae) is the type genus of Doleschallini . According to Art. 55.4. of the ICZN Code (ICZN 1999), one letter difference, which in this case would be an extra "i" from the suffix "-ina", avoids homonymy. However, it does not avoid confusion, including internet search engines that correct possible spelling errors, especially in this case, because it is a duplication of a letter in the suffix formation from the correctly determined stem. The Code gives a choice to alleviate the problem: Art. 29.4. (original stem formation to be maintained) and Recommendation 29A are followed here, and the entire name of the type genus is taken as a stem with hopes to diminish confusions. The choice of being ridiculed for grammatically incorrect stem formation is a "lesser evil" compared to increased confusion of those who use the name.</p><p>Additional genomic support to partition Nymphalis genus group into 4 genera: Hypanartia Hübner, [1821], Vanessa [Fabricius], 1807, Antanartia Rothschild &amp; Jordan, 1903, and Nymphalis Kluk, 1780</p><p>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</p></div>	https://treatment.plazi.org/id/03DC6105FFFF6944FEDEA620FA58C995	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFF36945FF5FA427FA9CC998.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Napeocles Bates 1864	<div><p>Napeocles Bates, 1864 is a junior subjective synonym of Siproeta Hübner, [1823]</p><p>We obtained genomic data for all 10 known species of Victorinini Scudder, 1893, and the resulting phylogenetic tree revealed that the monotypic genus</p><p>Napeocles Bates, 1864 (type species Hamadryas jucunda Hübner, [1808]) clusters very closely with</p><p>Siproeta Hübner, [1823] (type species Siproeta trayja</p><p>Hübner, [1823], currently a subspecies of Siproeta epaphus (Latreille, [1813])) (Fig. 44). The 4 species in this clade clump together, suggesting that Napeocles</p><p>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.</p></div>	https://treatment.plazi.org/id/03DC6105FFF36945FF5FA427FA9CC998	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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
03DC6105FFF16947FE8CA2A2FC92CA3A.text	03DC6105FFF16947FE8CA2A2FC92CA3A.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Microtia H. Bates 1864	<div><p>are junior subjective synonyms of Microtia H. Bates, 1864</p><p>Microtia H. Bates, 1864 (type and the only species Microtia elva H. Bates, 1864), Dymasia Higgins, 1960 (type and the only species Melitaea dymas W. H. Edwards, 1877) and Texola Higgins, 1959 (type species Eresia elada Hewitson, 1868) cluster closely in the genomic tree (Fig. 45). Our result agrees with the previous assessment based on gene markers (Wahlberg et al. 2005; Wahlberg and Freitas 2007; Long et al. 2014) and morphological study by Kons (2000), who already synonymized Texola and Dymasia with Microtia, a suggestion cited by Wahlberg et al. (2005) five years later but not followed either in that work (Wahlberg et al. 2005) or in subsequent publications (Pelham 2008; Long et al. 2014) and many on-line resources (Warren et al. 2016; North American Butterfly Association 2018; Wahlberg 2019a; Pelham 2020). Finally, more than two decades after these studies, we reach the same conclusion on a much larger DNA dataset: Texola Higgins, 1959 and Dymasia Higgins, 1960 are junior subjective synonyms of Microtia H. Bates, 1864 . A curious observation is that Microtia [= Texola] coracara (Dyar, 1912) appears to be more distant from other Texola than Dymasia from Microtia (Fig. 45), further supporting unification of these species. The unification is a more meaningful solution, because this clade actually consists of four semi-equal lineages, which are (1) M. elada (Hewitson, 1868) species group; (2) M. anomalus (Godman &amp; Salvin, 1897) species group that includes M. coracara; (3) M. dymas; and (4) M. elva; rather than the traditional three ( Texola, Dymasia and Microtia). An alternative could be to split the group into these 4, rather than 3, evolutionary lineages, but each of these 4 groups is nearly monotypic (or monotypic under some species concepts) and is a species group rather than a genus or even subgenus. Finally, Microtia sensu lato is characterized by an elevated evolutionary rate compared to its sister Chlosyne Butler, 1870 (type species Papilio janais Drury, 1782): in Fig. 45, branches of Microtia (green) are longer (i.e., stick out to the right more) than branches of Chlosyne (black). This elevated rate may be behind more pronounced phenotypic differences between Microtia species compared to Chlosyne that resulted in the oversplit classification of Microtia into genera.</p></div>	https://treatment.plazi.org/id/03DC6105FFF16947FE8CA2A2FC92CA3A	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFF16940FF32A663FBA2CB45.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Tisona Higgins 1981	<div><p>Tisona Higgins, 1981 is a junior subjective synonym of Ortilia Higgins, 1981</p><p>The monotypic genus Tisona Higgins, 1981 (type and the only species Phyciodes saladillensis Giacomelli, 1911) was proposed on the basis of genitalic differences, most significantly "penis apex with crossing ostium-folds" (Higgins 1981), i.e., with two twisted finger-like processes at the tip. Higgins likened Tisona to Tegosa Higgins,</p><p>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 ||</p><p>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</p><p>Papilio liriope Cramer, 1775), away from Tegosa (Fig. 45).</p><p>COI barcodes of O. liriope and T. saladillensis differ by 5.5%</p><p>(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</p><p>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. &amp; 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. &amp; 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.</p></div>	https://treatment.plazi.org/id/03DC6105FFF16940FF32A663FBA2CB45	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFF66941FDABA7A8FD54CFB2.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	(Notilia) Zhang & Cong & Shen & Opler & Grishin 2021	<div><p>Notilia Grishin, new subgenus</p><p>http://zoobank.org/ 3A8FA139-BE8C-444A-A212-6108DA6D3EAB</p><p>Type species. Eresia orthia Hewitson, 1864 .</p><p>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.</p><p>Etymology. The name is a feminine noun in the nominative singular, formed as a fusion Not + [Ort] ilia to indicate distinction from Ortilia .</p><p>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 .</p><p>Parent taxon. Genus Eresia Boisduval, 1836 .</p></div>	https://treatment.plazi.org/id/03DC6105FFF66941FDABA7A8FD54CFB2	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFF76941FDD9A3FBFD54CA12.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Levinata Grishin 2021	<div><p>Levinata Grishin, new subgenus</p><p>http://zoobank.org/ 5C69C6A2-7ADD-4E42-8818-23CCB9A042CE</p><p>Type species. Eresia levina Hewitson, 1872 .</p><p>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.</p><p>Etymology. The name is a feminine noun in the nominative singular, formed from the type species name.</p><p>Species included. Only the type species.</p><p>Parent taxon. Genus Eresia Boisduval, 1836 .</p></div>	https://treatment.plazi.org/id/03DC6105FFF76941FDD9A3FBFD54CA12	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFF76941FDA5A65CFD54C9A9.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Phyciodes ithra Kirby 1900	<div><p>Ithra Grishin, new subgenus</p><p>http://zoobank.org/ BADF7F37-1CFB-4846-AC56-6B125E8B6708</p><p>Type species. Phyciodes ithra Kirby, 1900 .</p><p>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.</p><p>Etymology. The name is a feminine noun in the nominative singular, tautonymous with the type species name.</p><p>Species included. Only the type species.</p><p>Parent taxon. Genus Eresia Boisduval, 1836 .</p></div>	https://treatment.plazi.org/id/03DC6105FFF76941FDA5A65CFD54C9A9	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFF46942FE53A25EFC73C893.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Phystis fontus (A. Hall 1928) Zhang & Cong & Shen & Opler & Grishin 2021	<div><p>Phystis fontus (A. Hall, 1928), new combination</p><p>Phystis variegata (Röber, 1913) and Phystis pratti (A. Hall, 1935) are species distinct from Phystis simois (Hewitson, 1864)</p><p>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),</p><p>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% &amp; 8.1%) between our specimens and P. simois available from GenBank (accession EF493956) (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.</p></div>	https://treatment.plazi.org/id/03DC6105FFF46942FE53A25EFC73C893	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFF46943FE4BA4DAFA85CCD7.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Abananote Potts 1943	<div><p>Abananote Potts, 1943 and Altinote Potts, 1943 are junior subjective synonyms of Actinote Hübner, [1819]</p><p>In agreement with previous studies (Silva-Brandao et al. 2008; Carvalho et al. 2020), we find that genera Actinote Hübner, [1819] (type species Papilio thalia Linnaeus, 1758), Abananote Potts, 1943 (type species Acraea abana Hewitson, 1868) and Altinote Potts, 1943 (type species Heliconius neleus Latreille, [1813]) are closely related and some of them are not monophyletic as currently circumscribed (Fig. 49). The type species of Abananote and Altinote are quite closely related: e.g. their COI barcodes differ by 6.4%</p><p>(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</p><p>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).</p></div>	https://treatment.plazi.org/id/03DC6105FFF46943FE4BA4DAFA85CCD7	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFF56943FE96A107FA20CB23.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Episcada Godman & Salvin 1879	<div><p>Episcada Godman &amp; Salvin, 1879 is a junior subjective synonym of Ceratinia Hübner, 1816</p><p>Genome-level phylogeny confirms problems with the current classification of Episcada Godman &amp; Salvin, 1879 (type species Ithomia salvinia Bates, 1864)</p><p>and Ceratinia Hübner, 1816 (type species Nerëis neso</p><p>Hübner, [1806]) (Willmott and Freitas 2006): the two genera are closely related and Episcada renders Ceratinia paraphyletic (Fig. 50), in agreement with other studies</p><p>(Chazot et al. 2020). COI barcodes of the Episcada and</p><p>Ceratinia type</p><p>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 &amp; Salvin, 1879 as a junior subjective synonym of Ceratinia Hübner, 1816 .</p></div>	https://treatment.plazi.org/id/03DC6105FFF56943FE96A107FA20CB23	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFF5695CFF73A7B1FA23CC1F.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Phocides batabano (Lucas 1857)	<div><p>Phocides batabano (Lucas, 1857) and Phocides bicolora (Boddaert, 1783) are species distinct from Phocides pigmalion (Cramer, 1779)</p><p>In his key, Evans (1952) stated that uncus flanges in genitalia of continental subspecies of Phocides pigmalion (Cramer, 1779) (type locality Suriname) are as long as the uncus, but are shorter in the island subspecies, being similar to other Phocides Hübner, [1819] (type species Phocides cruentus Hübner, [1819], which is Hesperia polybius Fabricius, 1793). Consistently with this notable genitalic difference, the genomic tree partitions P. pigmalion into several groups (Fig. 51) rendering it paraphyletic with respect to Phocides belus Godman &amp; Salvin, 1893</p><p>(type locality Mexico) and Phocides lincea (HerrichSchäffer, 1869) (type locality not stated, probably the Guianas), and suggesting that P. pigmalion is a</p><p>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</p><p>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).</p></div>	https://treatment.plazi.org/id/03DC6105FFF5695CFF73A7B1FA23CC1F	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFEA695CFE0BA04FFA26CB23.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Lobotractus mysie (Dyar 1904)	<div><p>Lobotractus mysie (Dyar, 1904) is a species distinct from Lobotractus valeriana (Plötz, 1881)</p><p>Correctly associating then " Codatractus " mysie (Dyar, 1904) with " Codatractus " valeriana (Plötz, 1881) as close relatives by visual inspection of photographs of their primary type specimens, Mielke and Warren</p><p>(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).</p><p>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. 2019a; 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.</p></div>	https://treatment.plazi.org/id/03DC6105FFEA695CFE0BA04FFA26CB23	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFEA695CFF5EA709FAA6C9BC.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Pyrgus centaureae subsp. dzekh Gorbunov 2007	<div><p>Pyrgus centaureae dzekh Gorbunov, 2007 is a new subspecies for North America</p><p>The genomic tree of Pyrgus centaureae (Rambur, 1839) (type locality Sweden) taxa reveals that a specimen collected by J. L. Harry west of Galbraith</p><p>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) .</p><p>Therefore, it is not Pyrgus centaureae freija (B.</p><p>Warren, 1924) (type locality Labrador), but Pyrgus Fig. 53. Pyrgus centaureae: Palearctic lineage (blue) centaureae dzekh Gorbunov, 2007 (type locality and Nearctic lineage (magenta).</p><p>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.</p></div>	https://treatment.plazi.org/id/03DC6105FFEA695CFF5EA709FAA6C9BC	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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
03DC6105FFEB695DFF35A2B5FCE2CFFF.text	03DC6105FFEB695DFF35A2B5FCE2CFFF.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Appia Evans 1955	<div><p>Appia Evans, 1955 is a junior subjective synonym of Pompeius Evans, 1955</p><p>The monotypic genus Appia Evans, 1955 (type and the only species Appia appia Evans, 1955) originates within Pompeius Evans, 1955 (type species Hesperia pompeius Latreille, [1824]) being sister to Pompeius amblyspila (Mabille, 1897) and rendering Pompeius paraphyletic (Fig. 54). All of these species are genetically close, e.g., COI barcodes of the type species of Appia and Pompeius differ by only 5% (33 bp). Therefore, to restore the monophyly, we treat Appia Evans, 1955 as a junior subjective synonym of Pompeius Evans, 1955 . Both taxa were proposed in the same work issued on the same date (Evans 1955), and using ICZN Code Art. 24, we give priority to Pompeius, because this name was used more frequently in the literature and the genus is not monotypic as Appia .</p></div>	https://treatment.plazi.org/id/03DC6105FFEB695DFF35A2B5FCE2CFFF	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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.text	03DC6105FFEB695DFED4A02EFE56C85C.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Wallengrenia Berg 1897	<div><p>Wallengrenia Berg, 1897 is a subgenus of Polites Scudder, 1872</p><p>We sequenced all major phenotypically distinct taxa from the Hylephila Billberg, 1820 (type species Papilio phyleus Drury, 1773) group of genera (Fig. 54, rooted with Hesperia</p><p>Fabricius, 1793). Confirming our previous assessment (Zhang et al.</p><p>2019c), we see that Polites Scudder,</p><p>1872 (type species Hesperia peckius</p><p>W. Kirby, 1837) is genetically close to</p><p>Wallengrenia Berg, 1897 (type species Hesperia premnas</p><p>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% (34bp) 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 &amp; 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. 2019d) 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.</p></div>	https://treatment.plazi.org/id/03DC6105FFEB695DFED4A02EFE56C85C	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Zhang, Jing;Cong, Qian;Shen, Jinhui;Opler, Paul A.;Grishin, Nick V.	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
