taxonID	type	description	language	source
421169606031B338FF7B27455AE4BB17.taxon	discussion	Here, we compile photographs of several specimens of both sexes that were included in the phylogenetic tree published as fig. 2 in Zhang et al. (2025 a) to illustrate phenotypic variation and interspecies differences in the northernmost populations of the two species: Chlosyne palla (Boisduval, 1852) (type locality in USA: California, Plumas Co.) and Chlosyne flavula (W. Barnes & McDunnough, 1918) (type locality USA: Colorado, Garfield Co., Glenwood Springs) (Zhang et al. 2023 c, 2025 a), specifically, Chlosyne palla sterope (W. H. Edwards, 1870) (type locality in USA: Oregon, Wasco Co.) and Chlosyne flavula blackmorei Pelham, 2008 (type locality Canada: British Columbia, Lytton) (Fig. 1). The two species are best differentiated by females: C. palla sterope is typically darker with cream-colored to nearly white spots and bands and usually a more elongated forewing apex; and C. flavula blackmorei has orange and orange-yellow spots and bands and a rounder forewing apex. We were not able to find consistent wing pattern differences between the northern populations of C. palla sterope (Canada: British Columbia, Osoyoos and USA: Washington, Okanogan Co.) and its more southern populations (USA: Washington, Adams Co. and Oregon, Wasco Co.), with some specimens being rather similar (e. g., compare Fig. 1 d with Fig. 1 l or Fig. 1 m for males, and Fig. 1 g with Fig. 1 q for females). Females are particularly alike. However, males from the northern populations are more variable, with some being more extensively orange (e. g., Fig. 1 j) than typical C. palla sterope, or exhibiting cream-colored (rather than orange-yellow) discal bands (Fig. 1 o). Genetically, all these specimens cluster together, forming a nuclear genome clade that also includes the lectotype of C. palla sterope (sequenced as NVG- 21011 C 11 and used to assign this name to the clade), but partition into the northern and southern subclades; see fig. 2 in Zhang et al. (2025 a).	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606031B338FF2522865BDDB944.taxon	description	http: // zoobank. org / B 13 E 52 B 7 - 663 E- 4 C 16 - 9 ADD-D 57302 A 19744	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606031B338FF2522865BDDB944.taxon	discussion	A subgenus of Lasaia H. Bates, 1868, Locris Grishin, 2025 (type species Lasaia oileus Godman, 1903) is a junior homonym of Locris Stål, 1866 (type species Cercopis rubra Fabricius, 1794), currently a valid genus of froghoppers (Hemiptera: Cercopidae) (Stål 1866). Here, according to Article 60.3 of the International Code of Zoological Nomenclature (1999), Lochris Grishin is proposed as a new substitute name that replaces Locris Grishin, 2025. According to the ICZN Code Article 67.8, the type species of Lochris Grishin, nom. nov. is Lasaia oileus Godman, 1903.	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606031B33DFE2E203C5BB9BE10.taxon	description	http: // zoobank. org / 98452 F 55 - 3 EB 2 - 4 B 5 A- 84 B 1 - 4 D 1 C 8 FF 074 BE (Figs. 2 part, 3)	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606031B33DFE2E203C5BB9BE10.taxon	materials_examined	Type material. Holotype: ♂ deposited in the National Museum of Natural History, Washington, DC, USA (USNM), illustrated in Fig. 3 a, bears the following eight rectangular labels (1 st handprinted, others printed with handwritten text shown in italics; 4 th blue, 5 th yellow, the last red, others white): [Joinville | 18 · IX · 1982], [StaCatharina | Brazil], [Presented by | Robert E. Aronheim], [JHALL | - 00 05], [LEGS AWAY | FOR DNA], [DNA sample ID: | NVG- 18044 E 12 | c / o Nick V. Grishin], [USNMENT | {QR Code} | 01466379], and [HOLOTYPE ♂ | Emesis (Mandania) | mandarina Grishin]. Paratypes: 1 ♂ and 2 ♀♀ from Brazil, Santa Catarina (last one likely mislabeled): 1 ♂ NVG- 25013 H 04 Joinville, 4 - Mar- 1985, H. Miers leg. [MGCL] (Fig. 3 c); 1 ♀ NVG- 24032 A 05 Blumenau, old, coll. Staudinger [MFNB] (Fig. 3 b); and 1 ♀ NVG- 24032 A 12 “ Colombia | R. Magdalena s ”, old, ex coll. H. Stichel, number 3280 [MFNB]. Type locality. Brazil: Santa Catarina, Joinville.	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606031B33DFE2E203C5BB9BE10.taxon	etymology	Etymology. The name is a fusion given to this relative of mand [ana from Santa Cat] arina, and is treated as a noun in apposition.	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606031B33DFE2E203C5BB9BE10.taxon	distribution	Distribution. Currently known only from Santa Catarina in Brazil.	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606035B33CFE1326015DF4B91A.taxon	description	http: // zoobank. org / E 971 DCB 7 - 6 DA 1 - 48 D 6 - BF 19 - C 1 E 866 DE 2226 (Figs. 2 part, 4 a)	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606035B33CFE1326015DF4B91A.taxon	materials_examined	Type material. Holotype: ♂ deposited in the Museum für Naturkunde, Berlin, Germany (MFNB), illustrated in Fig. 4 a, bears the following five printed rectangular labels (text in italics handwritten), four white: [Pto Cabello | Hahnel], [Coll. | Staudinger], [DNA sample ID: | NVG- 24033 B 06 | c / o Nick V. Grishin], [{QR Code} MfN URI | http: // coll. mfn- | berlin. de / u / | 09 f 2 f 9], and one red [HOLOTYPE ♂ | Emesis (Mandania) | mandela Grishin]. Type locality. Venezuela: Carabobo, Puerto Cabello.	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606035B33CFE1326015DF4B91A.taxon	etymology	Etymology. The name is a fusion given to this relative of mand [ana from Venezu] ela, and is treated as a noun in apposition.	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606035B33CFE1326015DF4B91A.taxon	distribution	Distribution. Currently known only from the holotype collected in coastal Venezuela.	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606035B33CFE1326015DF4B91A.taxon	discussion	Comment. This new species is the third Emesis (Mandania) species we recorded in Venezuela, in addition to Emesis (Mandania) mandana (Cramer, 1780) (type locality in Suriname) and Emesis (Mandania) mandora Grishin, 2024 (type locality in Ecuador: Santo Domingo) (Fig. 2 yellow highlight). Additional records of Emesis (Mandania) mantunga Grishin, 2025 Through expanded genomic sequencing (Fig. 2), we found three more specimens of Emesis (Mandania) mantunga Grishin, 2025 (type locality Ecuador: Tungurahua Province, Topo; originally described from five males), including the first confirmed female (NVG- 25013 E 02, Ecuador: Napo Province, Puerto Misahuallí, 6 - Nov- 1983, D. & J. Jenkins [MGCL], Fig. 4 c), an additional male from eastern Ecuador (NVG- 24033 A 11, Pastaza Province, Sarayacu, old, R. Haensch S., Stichel collection number 3282 [MFNB]), and extend the distribution of this species to the eastern slopes of the Andes in northern Peru (♂ NVG- 24033 A 10, San Martín Department, Tarapoto, old, Stichel collection number 4338 [MFNB]).	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606035B331FE9920855CACBB60.taxon	description	http: // zoobank. org / C 615 B 4 FA- 4 DD 4 - 491 D-B 7 EB- 0 C 54 C 481 D 1 DF (Figs. 5 part, 6 a)	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606035B331FE9920855CACBB60.taxon	materials_examined	Type material. Holotype: ♂ deposited in the McGuire Center for Lepidoptera and Biodiversity Collection, Gainesville, FL, USA (MGCL), illustrated in Fig. 6 a, bears the following four rectangular labels (1 st handwritten, others printed with handwritten text shown in italics), three white: [C. RICA: PUNTARENAS | Corcovado; 20. ix. 1976 | P. DeVries], [Allyn Museum | Acc. 19 78 - 20], [DNA sample ID: | NVG- 24073 H 04 | c / o Nick V. Grishin], and one red [HOLOTYPE ♂ | Telegonus (Rhabdoides) alector | obscuratus Grishin]. Type locality. Costa Rica: Puntarenas Province, Corcovado National Park.	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606035B331FE9920855CACBB60.taxon	etymology	Etymology. In Latin, obscuratus means darkened, made dark, or having become dark, and is given for the lack of the white smudge in the middle of the forewing in males characteristic of other T. alector subspecies. The name is a perfect passive participle.	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606035B331FE9920855CACBB60.taxon	distribution	Distribution. Currently known only from the holotype collected in southern Costa Rica.	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606038B330FF56222D5AD0BCA2.taxon	discussion	Genomic sequencing of additional specimens of Telegonus Hübner, [1819] (type species Papilio talus Cramer, 1777) reveals that Telegonus (Rhabdoides) missionus Grishin, 2025 (type locality USA: Texas, Hidalgo Co., Mission, holotype sequenced as NVG- 14111 E 04) is a species widely distributed in eastern Mexico, recorded from the states of Tamaulipas, Nuevo Leon, and Veracruz (Fig. 5 red), thus confirming it as a species-level taxon, and not an unusual single specimen. This recently described species (Zhang et al. 2025 a) is sympatric with Telegonus (Rhabdoides) gilberti H. Freeman, 1969 (type locality in Mexico, San Luis Potosí, holotype sequenced as NVG- 15104 B 08) over its range, with sequenced specimens of the latter species from the same three Mexican states shown in the trees (Fig. 5 blue). Both species (T. missionus and T. gilberti) have been recorded from Hidalgo County in Texas, USA (Fig. 5 highlighted yellow). In the nuclear genome trees, T. missionus is sister to Telegonus (Rhabdoides) hopfferi (Plötz, 1881) (type locality in Mexico, probably Oaxaca), a southern Mexico species, and specimens of both species were sequenced from Veracruz, (Fig. 5 a, b). As in our previous study (Zhang et al. 2025 a), we observe confidently supported incongruence between the three phylogenetic trees of the T. alector species group (Fig. 5). In the Z chromosome tree (Fig 5 b) and the mitochondrial genome tree (Fig 5 c), the T. alector group partitions into two prominently separated clades with T. gilberti being in the second clade, thus more strongly differentiated genetically from the three species in the first clade: Telegonus (Rhabdoides) alector (C. Felder & R. Felder, 1867) (type locality in Colombia), T. hopfferi, and T. missionus; while in the tree constructed from protein-coding regions of autosomes (Fig 5 a), all four species belong to the same clade, with T. gilberti being sister to the other three species. In addition to males, genomic sequencing also reveals females of T. missionus, and one is illustrated here for the first time (Fig. 7). Females are similar to males in their darker appearance compared to females of related species (but paler than a typical T. missionus male), with a reduced white band on the ventral forewing, which has a beige (i. e., white scales sprinkled over brown) costal area from the base to about a third of the forewing length; a white triangle partly overscaled with brown at the base of the ventral hindwing; weakly expressed but noticeable pale overscaling in the discal area of the dorsal forewing corresponding to the white ventral band; and darker (not prominently orange-yellow) ventral side of the body.	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606039B336FECA226A5B75BC4D.taxon	description	http: // zoobank. org / 16 CBDDCE- 329 A- 402 A- 9 C 25 - 25 D 6790 F 53 EC (Figs. 8 part, 9 b – c)	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606039B336FECA226A5B75BC4D.taxon	materials_examined	Type material. Holotype: ♀ deposited in the Museum für Naturkunde, Berlin, Germany (MFNB), illustrated in Fig. 9 b, bears the following seven rectangular labels (first four handwritten, others printed), six white: [Columbia | 86 Klbr.], [201.], [T. chiriquensis | ♀ var.], [chiriquensis | var.], [{QR Code} MfN URI | http: // coll. mfn- | berlin. de / u / | 09 ec 52], [DNA sample ID: | NVG- 24028 C 10 | c / o Nick V. Grishin], and one red [HOLOTYPE ♀ | Telegonus (Rhabdoides) | flavifimbro Grishin]. Paratype: 1 ♀: NVG- 24039 F 01 Colombia, Boyacá, Muzo, Mar- 1918, W. Gerstner leg. [SMNS] (Fig. 9 c). Type locality. Colombia, possibly in the eastern Andes.	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606039B336FECA226A5B75BC4D.taxon	etymology	Etymology. Formed similarly to flavimargo, the name is given for the orange-yellow fringes (fimbia in Latin), particularly noticeable in the holotype of this species. The name is also longer to indicate a more southern distribution of this species and is treated as a noun in apposition.	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606039B336FECA226A5B75BC4D.taxon	distribution	Distribution. Currently known only from Colombia.	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
42116960603FB336FF002505590CBAB4.taxon	discussion	Genomic analysis of over two dozen Telegonus (Rhabdoides) cretatus Hayward, 1939 (type locality in Ecuador: Napo) specimens from across the range reveals that they partition into two comb-like clades genetically differentiated at the species level (Fig. 10). The first clade includes specimens from French Guiana, Venezuela, Ecuador, Peru, Bolivia, and Amazonian Brazil and corresponds to the nominate subspecies. The second clade is composed of specimens from the Atlantic states of Brazil from Bahia to Santa Catarina and represents a taxon originally described as a subspecies Astraptes cretatus adoba Evans, 1952 (type locality in Brazil: Espírito Santo) and currently treated as such, but now placed in Telegonus (Rhabdoides). Although it differs by only 0.6 % (4 bp) from the nominate subspecies, this difference is consistent throughout the range, and the nuclear genome clades suggest a species-level distinction. Phenotypically, the Atlantic taxon is darker than the Amazonian (e. g., nearly lacking the ventral forewing green / white area by the costal margin at the base) and has a typically less robust serrated dorsal ridge of the harpe with a more rounded ventrocaudal angle. Therefore, we propose Telegonus (Rhabdoides) adoba (Evans, 1952), stat. nov. is a species distinct from Telegonus (Rhabdoides) cretatus Hayward, 1939.	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
42116960603FB335FF25207A5CD4B9E6.taxon	discussion	Genomic analysis enables us to confidently propose new species from a single specimen of either sex that is genetically differentiated from others at the species level (Zhang et al. 2025 a). However, even fortified with the whole genome shotgun dataset, this single-specimen approach carries a risk of describing a hybrid or a contaminated dataset as a “ species, ” despite all the precautions and careful analysis we undertake. Therefore, we strive to find and sequence additional specimens of the newly proposed species and investigate them further. Here, we confirm the species-level status of Telegonus (Rhabdoides) flavimargo Grishin, 2025 (type locality in Costa Rica: Limón, 2 additional specimens) (Fig. 8 green) and Telegonus (Rhabdoides) tatus Grishin, 2005 (type locality in Panama: Panamá, 6 additional specimens) (Fig. 10 maroon) proposed from a single specimen and Telegonus (Rhabdoides) panamus Grishin, 2025 (type locality in Panama: Barro Colorado Island, 4 additional specimens) originally described from the holotype and the paratype (Fig. 10 purple). For all these species, additional specimens group closely with the holotypes in the genomic trees, resulting in comb-like clades characteristic of conspecific specimens. Although for T. flavimargo all three known specimens are from the same locality (Costa Rica: Limón Province, Guapiles), we are able to extend the known distribution for the other two species: T. panamus has been recorded from both central (around the Panama Canal) and eastern Panama (Darien) (Fig. 10 purple); and two specimens from northern Ecuador fall in the same clade among T. tatus specimens from central Panama and therefore, we identify them as this species (Fig. 10 maroon).	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
42116960603CB334FF0C20D459DABDB1.taxon	discussion	Additional genomic sequencing and analyses suggest several revisions to our preliminary taxonomic list of species from the subgenus Rhabdoides Scudder, 1889 (only from the clade we have analyzed). The list from Zhang et al. (2025 a) is updated below, using the same rationale and format. We introduced a new species and a new subspecies and revised the status of one subspecies to species. Furthermore, tweaks to the order were made. For instance, Telegonus chuchuvianus Grishin, 2025 was moved next to its most likely sister group of two species: Telegonus meretrix (Hewitson, 1876) and Telegonus fulvimargo Grishin, 2025, as evidenced by strong statistical support in both the Z chromosome and the mitochondrial genome trees (Fig. 10). The former species was placed before the latter two, because it lacks the yellower ventral hindwing margin characteristic of the latter, and the next species group in the list (the latimargo species group) starts with pale-margined species. In the following arrangement from Zhang et al. (2025 a), refined below, species of Rhabdoides excluding the clades with Telegonus anaphus (Cramer, 1777) and Telegonus cellus (Boisduval & Le Conte, [1837] are given. The list also includes species discovered by Steinhauser (1987) (“ four new species will be added to the group ”) that fall within these species groups but remain unpublished, shown in gray font. Type localities (general area only: state, region, department, or county) are in gray font. New taxa described in this study and the category of taxonomic change are in red font. Taxonomic treatment before this work (for valid names) and comments are given in smaller font following a vertical bar | after the type locality; an equal sign = precedes synonyms given in their original genus combination; and a double dagger ‡ marks unavailable names. The list covers 48 valid taxa and 4 yet undescribed species, comprising 46 species (1 newly proposed here and 1 formerly treated as a subspecies) and 6 additional subspecies (1 new).	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
42116960603DB334FF2F255159E0BCEE.taxon	type_taxon	type species Eudamus cellus Boisduval & Le Conte, [1837] alector species group	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
42116960603DB334FF5F25B45CE8BC9D.taxon	distribution	Colombia: Bogotá	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
42116960603DB334FF5F226F5C10BB46.taxon	distribution	Ecuador: Esmeraldas	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
42116960603DB334FF5F25D15C3FBCB0.taxon	distribution	Costa Rica: Puntarenas	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
42116960603DB334FF0F23695BD0BA78.taxon	distribution	Brazil: Rondônia	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
42116960603DB334FF0F23A65C83B96B.taxon	distribution	Guatemala: San Gerónimo	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
42116960603DB334FF0F21E15C44B8C0.taxon	distribution	unknown, likely SE or S Brazil	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
42116960603DB334FF0F209F5CA0B818.taxon	distribution	no data [likely SE or S Brazil]	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
42116960603DB334FF0F22FF5B97BBD6.taxon	distribution	Mexico: San Luis Potosí	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
42116960603DB334FF0F22425B3FBB01.taxon	distribution	Mexico [likely C or S Mexico]	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
42116960603DB334FF0F22045B85BBED.taxon	distribution	USA: Texas, Hidalgo Co.	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
42116960603DB334FF0F22B55B4FBB9C.taxon	distribution	Peru: Piura	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
42116960603DB334FF0F207C5B1EB955.taxon	distribution	Panama: Darién	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
42116960603DB334FF0F22D25B22BBB3.taxon	distribution	Panama: Panamá	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
42116960603DB334FF0F200A5BF6B91B.taxon	distribution	Brazil: Santa Catarina	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606022B32BFF0F22435D3DBB22.taxon	distribution	Brazil: Espírito Santo | was a subspecies of T. cretatus	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606022B32BFF0F20775BFEB95E.taxon	distribution	Brazil: Rio de Janeiro	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606022B32BFF0F21175B08B8FE.taxon	distribution	Costa Rica: Irazú	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606022B32BFF0F27165B09BEFF.taxon	distribution	Mexico: Chiapas	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606022B32BFF0F27CA5CE2BE84.taxon	distribution	Panama: Chiriquí	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606022B32BFF0F22265B81BB0F.taxon	distribution	Ecuador: Esmeraldas	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606022B32BFF0F27805B0DBD61.taxon	distribution	Colombia: Cauca	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606022B32BFF0F25B75C82BB7B.taxon	distribution	Ecuador: Napo	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606022B32BFF0F24865AA7BC6F.taxon	distribution	Suriname	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606022B32BFF0F245F5B22BD36.taxon	distribution	Ecuador: Balzapamba	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606022B32BFF0F24C85C14BDD9.taxon	distribution	Colombia [likely eastern Andes]	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606022B32BFF0F24315BCABD10.taxon	distribution	Costa Rica: Limón	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606022B32BFF0F22FD5B18BBD4.taxon	distribution	Peru: Cuzco	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606022B32BFF0F21335B20B812.taxon	distribution	Peru: Chanchamayo	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606022B32BFF0F26AF5B30BF86.taxon	distribution	Panama: Chiriquí	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606022B32BFF0F202A5AE1B93B.taxon	distribution	Cuba	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606022B32BFF0F200D5DDBB9AD.taxon	distribution	" Brazil " [Dominican Republic]	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606022B32BFF0F22195B43BBE8.taxon	distribution	Ecuador	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606022B32BFF0F25F95C45BCC8.taxon	distribution	Panama: Barro Colorado Island	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606022B32BFF0F25065B3FBCEF.taxon	distribution	likely Suriname	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606022B32BFF0F27655BF5BE4C.taxon	distribution	Peru: Madre de Dios	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606022B32BFF0F257A5BC6BC4B.taxon	distribution	Brazil: Santa Catarina	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606022B32BFF0F27585DE3BE15.taxon	distribution	Mexico: Veracruz	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606022B32BFF0F21605DC2B82A.taxon	distribution	Ecuador: Chimborazo	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606022B32BFF0F25DC5B73BCB5.taxon	distribution	Panama: Panamá	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606022B32BFF0F24A35AB7BD82.taxon	distribution	Brazil: Pará	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606023B32AFF4726165A70BEBF.taxon	discussion	In the original description of Urbanus (Urbanoides) elma Grishin, 2025 (type locality in Venezuela: Mérida) we provided a phylogenetic tree showing specimens of Urbanus (Urbanoides) elmina Evans, 1952 (type locality in Ecuador: Rio Pastaza) from Ecuador, Peru, and Argentina (Zhang et al. 2025 a). The only specimen from Colombia (without further locality details) was the paratype of U. elma. To date, we have sequenced specimens of U. elmina from additional localities in Ecuador and also from western Colombia (Valle del Cauca and Cauca Departments), thus confirming this species from Colombia (Fig. 11). While we have not found more specimens of U. elma, we stumbled upon a specimen of a new species, which is described next.	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606023B32FFE14208059D1BED8.taxon	description	http: // zoobank. org / 89 F 465 B 0 - F 9 CE- 4 BB 0 - A 902 - 257402570 AD 3 (Figs. 11 part, 12 b, 13)	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606023B32FFE14208059D1BED8.taxon	materials_examined	Type material. Holotype: ♀ deposited in the McGuire Center for Lepidoptera and Biodiversity Collection, Gainesville, FL, USA (MGCL), illustrated in Fig. 12 b (genitalia Fig. 13), bears the following five rectangular labels (2 nd handwritten, others printed with handwritten text shown in italics), four white: [COLOMBIA: TOLIMA | La Aurora, R. Cambrin | 1300 m.; 17. XI. 1974 | S. & L. Steinhauser], [GENIT. PREP. | SRS- 406], [A. C. Allyn | Acc. 1975 - 17], [DNA sample ID: | NVG- 24111 A 06 | c / o Nick V. Grishin], and one red [HOLOTYPE ♀ | Urbanus (Urbanoides) dolus Grishin]. Genitalia of the holotype, misidentified as Urbanus elmina, were illustrated in fig. 58 by Steinhauser (1981). Type locality. Colombia: Tolima, La Aurora, Río Cambrín, elevation 1300 m.	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606023B32FFE14208059D1BED8.taxon	etymology	Etymology. In Greek, δόλος (dólos) means deceit, treachery, guile, or craftiness; the same meaning carries to Latin dolus. The name is given for the deceitful appearance of this species, which is sister to U. viridis, but in the ventral hindwing pattern looks more similar to U. elmina or U. elma and was identified as U. elmina by Steinhauser (1981). The name is an adjective.	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606023B32FFE14208059D1BED8.taxon	distribution	Distribution. Currently known only from the holotype collected on the western slopes of the eastern Andes of Colombia.	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606026B32EFE01279B5D93BB41.taxon	discussion	Currently, Burnsius communis (Grote, 1872) (type locality in USA: central Alabama) is partitioned into two subspecies: the nominate, distributed throughout most of the range from Canada to Mexico; and Burnsius communis albescens (Plötz, 1884) (type locality in Mexico), known from the southern parts of the range (e. g., Oaxaca and Veracruz in Mexico). In line with this treatment, B. communis albescens is a confidently supported sister to all other populations (98 % bootstrap value) in the nuclear genome tree constructed from more than 10 million positions in autosome protein-coding genes (Fig. 14 a). However, due to their overall lack of prominent genetic differentiation and continuing gene flow within a species, subspecies do not always stand out as major clades in phylogenetic trees. Accordingly, the clade of B. communis albescens specimens, although supported with 100 % ultra-fast bootstrap value (Minh et al. 2013), is found within the nominate clade in the Z chromosome tree (Fig. 14 b), likely due to poor phylogenetic resolution (low support for most bifurcations) caused by DNA similarity and gene flow. We note that mitochondrial DNA does not consistently segregate several species of Burnsius Grishin, 2019 (type species Syricthus [sic] communis Grote, 1872), such as B. communis, B. albezens Grishin, 2022 (type locality in USA: AZ, Cochise Co.), and B. burnsi Grishin, 2022 (type locality in Mexico: Veracruz), and its utility is equally limited for subspecies delimitation. Additional sequencing of B. communis specimens across the range revealed that specimens from Stanislaus National Forest in Tuolumne County, California, were partitioned between different clades (Fig. 14 yellow highlight). While several specimens were in a confidently supported (99 % – 100 % bootstrap value) nuclear genome clade of specimens from the northwestern part of the range, one specimen was in a clade that included more western and southern specimens. This latter specimen was collected at the same locality and on the same day with one of the specimens from the northwestern clade (near Mill Creek Campground, elevation 6525 ’, GPS 38.312 6, − 119.939 8, 21 - Jul- 2022, W. R. Dempwolf leg.). These specimens were different in size and wing pattern (Fig. 15 h vs. Fig. 15 l). Moreover, a similar scenario was later found for four specimens of the same size from south of Lake Tahoe, El Dorado County, California, with two being in the northwestern clade and two (collected several miles to the east) falling in the southeastern clade (Fig. 14 orange highlight; Fig. 15 j, k, m, n). Therefore, the northwestern and southeastern clades represent different taxa. Although it is possible that they are species-level taxa, it is also possible that they may be subspecies that intergrade at certain localities and elevations. Currently, several taxa with similar relationships are treated as subspecies (type localities in parentheses): Hesperia colorado sublima A. Warren & Calhoun, 2015 (USA: Colorado, Clear Creek Co.) vs. Hesperia colorado colorado (Scudder, 1874) (USA: Colorado, Lake Co.) and Apodemia virgulti dialeucoides J. Emmel, T. Emmel & Pratt, 1998 vs. Apodemia virgulti nigrescens J. Emmel & T. Emmel, 1998 (both USA: California, San Bernardino Co.) (Pelham 2023). Hence, and due to the lack of prominent genetic differentiation characteristic of species, the new taxa of Burnsius are conservatively described here as subspecies of B. communis, pending more detailed analyses and further studies. Warren (2005) reviewed Oregon populations and also noted phenotypic differences among them.	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606027B323FEC222095DD8BE68.taxon	description	http: // zoobank. org / E 738 CD 38 - E 871 - 4 B 6 A-A 77 E- 4 C 6 C 5 ED 5969 E (Figs. 14 part, 15 a – d, 16 a – b)	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606027B323FEC222095DD8BE68.taxon	materials_examined	Type material. Holotype: ♂ deposited in the McGuire Center for Lepidoptera and Biodiversity Collection, Gainesville, FL, USA (MGCL), illustrated in Fig. 15 a (genitalia Fig. 16 a, b), bears the following four printed rectangular labels (handwritten text shown in italics), three white: [W. of Little Deschutes Riv. | near Crescent El. 4400 ' | Klamath Co, Oregon | August 21, 2004 | COLLECTORS JUNE | & FLOYD PRESTON], [MGCL Accession | 2010 - 33 | J. & F. Preston], [DNA sample ID: | NVG- 24064 E 07 | c / o Nick V. Grishin], [genitalia: | NVG 241111 - 10 | c / o Nick V. Grishin] and one red [HOLOTYPE ♂ | Burnsius communis | tenebrunis Grishin]. Paratypes: 3 ♂♂ and 3 ♀♀: USA, J. & F. Preston leg. [MGCL]: Oregon: 1 ♂ NVG- 23058 A 02 Jackson Co., 3.2 mi S of OR- 66 on Soda Mt. Rd., 5000 ’, 31 - May- 1996; Klamath Co., Deschutes National Forest, Little Deschutes River nr. Mowich, 4700 ’: 1 ♀ NVG- 23058 A 06 29 - Jun- 2002 (Fig. 15 b), 1 ♀ NVG- 24064 G 09 27 - Jun- 2007, and 1 ♂ NVG- 24064 E 08 29 - Jun- 2008; and 1 ♂ NVG- 23058 A 05 Lake Co., Warner Mts., Fremont National Forest, FR 3915 4.4 rd. mi S of Camas Creek, 6000 ’, 7 - Jul- 2008; and California: 1 ♀ NVG- 23058 A 03 Del Norte Co., 2 rd. mi E of Rowdy Creek Rd. on Low Divide Rd., 1600 ’, 1 - Sep- 2001. Other specimens: Due to genetic similarity (Fig. 14), we currently attribute the following three sequenced specimens to this subspecies but exclude them from the type series, as they exhibit stronger phenotypic differences — being paler and larger — compared to the population at the type locality: British Columbia [CNC]: 1 ♂ NVG- 24012 E 09, CNCLEP _ 00163304 Kaslo, 19 - Jun- 1900, J. W. Cockle (Fig. 15 d) and 1 ♂ NVG- 24012 E 08, CNCLEP _ 00163301 Fernie, 12 - Jun- 1934, H. B. Leech (Fig. 15 c) and 1 ♀ NVG- 23058 A 07 USA, Oregon, Wallowa Co., 12 road mi NE of Joseph, on road to Imnaha, 3700 ’, 26 - Jul- 2007, J. & F. Preston leg. [MGCL]. Type locality. USA: Oregon, Klamath Co., Deschutes National Forest, west of Little Deschutes River, nr. Crescent, 4400 ’.	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606027B323FEC222095DD8BE68.taxon	etymology	Etymology. In Latin, tenebrosus means dark, gloomy, or shadowy, and brunneus means brown. The name is formed as a fusion: tene [brosus] + brun [neus] + [commun] is, given for the darker and browner (not greener) aspect of this subspecies, and is treated as an adjective.	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606027B323FEC222095DD8BE68.taxon	distribution	Distribution. From British Columbia (Canada), through Oregon to northwestern California (USA).	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
42116960602AB323FE11272F5A50B8CB.taxon	description	http: // zoobank. org / 5 DF 6 A 207 - 4270 - 414 E- 91 DB- 8104 C 19 B 2452 (Figs. 14 part, 15 g – k, 16 c – d)	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
42116960602AB323FE11272F5A50B8CB.taxon	materials_examined	Type material. Holotype: ♂ deposited in the McGuire Center for Lepidoptera and Biodiversity Collection, Gainesville, FL, USA (MGCL), illustrated in Fig. 15 g, (genitalia Fig. 16 c, d), bears the following six printed rectangular labels, five white: [CALIF.: Tuolumne Co. | 4 mi. S of Mill Creek | at CA Hwy. 108, 6500 | ft.; 30. vi. 1987 | L. D. & J. Y. Miller | sta. no. 4], [Allyn Museum | Acc. 1987 – 8], [DNA sample ID: | NVG- 23057 F 09 | c / o Nick V. Grishin], [DNA sample ID: | NVG- 24067 E 03 | c / o Nick V. Grishin], [genitalia: | NVG 241111 - 29 | c / o Nick V. Grishin] and one red [HOLOTYPE ♂ | Burnsius communis | altus Grishin]. The first DNA sample refers to the extraction from a leg (sequenced) and the second is from the abdomen (stored) prior to genitalia dissection. Paratypes: 3 ♂♂ and 3 ♀♀: USA, California: Placer Co., B. O’Hara leg. [MGCL]: 1 ♂ NVG- 23058 D 09 Soda Springs Rd. 2.9 - 4.1 mi S. of Donner Pass Rd., 27 - Jul- 1992 and 1 ♂ NVG- 23058 D 08 Squaw Valley Ski Area, 8200 - 8500 ’, 6 - Jun- 1994; El Dorado Co. [CNC]: 1 ♂ NVG- 24012 E 04, CNCLEP _ 00163011 Fallen Leaf, 13 - Jul- 1961, J. G. Chillcott leg. (Fig. 15 j) and 1 ♀ NVG- 24012 E 05, CNCLEP _ 00163017 Echo Lake, 13 - Jul- 1961, B. H. Poole leg (Fig. 15 k); and Tuolumne Co., Stanislaus National Forest, 21 - Jul- 2022, W. R. Dempwolf leg. [WRD]: 1 ♀ NVG- 23032 C 11, WRD 22431 near Niagara Campground, 6870 ’, 38.320 6, − 119.911 4 (Fig. 15 i) and 1 ♀ NVG- 23032 C 12, WRD 22432 near Mill Creek Campground, 6525 ’, 38.312 6, − 119.939 8 (Fig. 15 h). Type locality. USA: California, Tuolumne Co., 4 mi south of Mill Creek at SR 108, elevation 6500 ft.	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
42116960602AB323FE11272F5A50B8CB.taxon	etymology	Etymology. In Latin, altus means high, deep, or tall, and is given for the typical habitat of this subspecies at higher elevations. The name is an adjective.	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
42116960602AB323FE11272F5A50B8CB.taxon	distribution	Distribution. Currently known from the central Sierra Nevada in California, USA (Placer, El Dorado, and Tuolumne Counties), where it comes close to and may overlap at times with the nominate subspecies, especially at lower elevations.	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
42116960602BB322FEA326015C7DBDCF.taxon	description	confirm it as a species-level taxon	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
42116960602BB322FEA326015C7DBDCF.taxon	discussion	Genomic analysis of Heliopetes Billberg, 1820 (type species Papilio niveus Cramer, 1775, which is a junior subjective synonym of Papilio arsalte Linnaeus, 1758) reveals four additional specimens of Heliopetes (Heliopetes) acuta Grishin, 2024 (type locality Mexico, Oaxaca, Candelaria Loxicha), all from the type locality and collected by E. C. Welling (Fig. 17 red), thus confirming that it is a species-level taxon and not an unusual single specimen. In all three trees, H. acuta is sister to the clade consisting of three species: Heliopetes (Heliopetes) lana Grishin, 2023 (type locality in Guatemala), Heliopetes (Heliopetes) alana (Reakirt, 1868) (type locality in Colombia), and Heliopetes (Heliopetes) chimbo Evans, 1953 (type locality in Ecuador: Chimbo), and, therefore, is the most distinct species in this group. Here, we illustrate male genitalia of H. acuta (Fig. 18) and they differ from those of H. lana, a species closest in distribution or possibly even sympatric with it (see figs. 302 – 303 in Zhang et al. (2023 a) for its genitalia photographs), by a terminally rounder and flatter (shell-shaped in the dorsal half) harpe, which is not prominently turning inward and bears coarser serrations mostly along its dorsoposterior margin; a more robust ampulla; and slightly narrower, more separated uncus arms.	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606028B326FE46261D592ABE9F.taxon	discussion	Previously treated as a single species Ochlodes sylvanoides (Boisduval, 1852) (type locality in USA: California, Plumas Co.), this species group is currently placed in the subgenus Ochluma Grishin, 2025 (type species Hesperia yuma W. H. Edwards, 1873) and, as here defined, consists of three genetically differentiated species: Ochlodes (Ochluma) santacruza J. Scott, 1981 (type locality USA: California, Santa Barbara Co., Santa Cruz Island), O. sylvanoides, and Ochlodes (Ochluma) napa (W. H. Edwards, 1865) (type locality in USA: Colorado, Clear Creek Co.) (Zhang et al. 2023 b). We expanded genomic sequencing of this group by sampling additional specimens across the range from southwestern Canada through all 13 western states of the USA to Baja California, Mexico (Fig. 19). The results confirm the phylogenetic separation of the group into three species and reveal additional insights, some of which are unexpected. First, the nominate subspecies of O. napa (Fig. 19 yellow circles) is restricted to the mountainous region of Colorado and its immediate neighborhood in southeastern Wyoming, eastern Utah, and northern New Mexico. Second, Ochlodes (Ochluma) napa kaibab Grishin, 2023 (type locality in USA: Arizona, Coconino Co.) (Fig. 19, blue circles) has a much wider distribution than anticipated and spreads as a narrow strip north from the type locality through central Utah, reaching northwestern Wyoming. This subspecies of O. napa, genetically differentiated from the nominate, represents populations between it and neighboring O. sylvanoides while being phylogenetically associated with the former. It is also possible that the populations of O. napa in Wyoming, Utah, and Arizona are best partitioned into several subspecies. They have different, albeit more closely related, mitogenome haplotypes (Fig. 20 b) and show different levels of introgression with other taxa. For instance, specimens of O. napa kaibab from Emery County, Utah, introgress stronger with O. napa napa and thus are placed near the base of the nuclear genome tree (Fig. 20 a) and possess mitochondrial genomes of O. napa napa. Nevertheless, specimens shown as blue circles in Fig. 19 are in the clade with O. napa kaibab, and we presently treat them as this subspecies. Furthermore, the populations of O. napa kaibab in Utah are coming close to and may even be sympatric with O. sylvanoides, a question to be addressed in future studies. Third, paler-colored populations to the west of the main Rocky Mountains chain north of Central Wyoming (Fig. 19 magenta and violet squares) traditionally associated with O. napa do not belong to this species and are O. sylvanoides instead. Due to their wing pattern differences that resulted in this misidentification, these populations are described below as two new subspecies that are somewhat differentiated genetically from other O. sylvanoides populations. These populations of O. sylvanoides are geographically close to O. napa kaibab in northwestern Wyoming and south-central Montana, another region to study interactions between the two species O. napa and O. sylvanoides. Fourth, phylogenetic analysis did not reveal prominent genetic differences of previously described O. sylvanoides subspecies: Ochlodes sylvanoides orecoasta J. Scott, 1981 (type locality in USA: Oregon, Clatsop Co.), Ochlodes sylvanoides bonnevilla J. Scott, 1981 (type locality in USA: Nevada, Elko Co.), and Ochlodes sylvanoides omnigena Austin, 1998 (type locality in USA: Nevada, Lander Co.). While we do not propose synonymizing these subspecies, studies of their distinction are beyond the scope of this work and will be addressed in future research. Here, all these populations combined are shown as green squares in Fig. 19.	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
42116960602FB325FE9B245159E2BD29.taxon	description	http: // zoobank. org / ADC 1 FE 05 - 7 B 71 - 4 C 9 D-B 788 - 64 E 7 A 2 F 82 FE 2 (Figs. 19 part, 20 part, 21 b – d)	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
42116960602FB325FE9B245159E2BD29.taxon	materials_examined	Type material. Holotype: ♂ deposited in the McGuire Center for Lepidoptera and Biodiversity Collection, Gainesville, FL, USA (MGCL), illustrated in Fig. 21 b, bears the following five printed (handwritten text in italics) rectangular labels, four white: [WY WASHAKIE Co. | T 47 N R 86 W S 5 | Tensleep Reserve | 6400 ' 11 - 12. viii. 98], [Allyn Museum | Acc. 1998 - 15], [Ochlodes sylvanoides | (Boisduval, 1852) ♂ | Det. S. R. Steinhauser], [DNA sample ID: | NVG- 24126 C 06 | c / o Nick V. Grishin], and one red [HOLOTYPE ♂ | Ochlodes sylvanoides | paranapa Grishin]. Paratypes: 4 ♂♂ and 1 ♀ from USA in MGCL: Montana, Big Horn Co.: 1 ♂ NVG- 24126 B 10 foothills of Pryor Mts., along Sage Creek, 5550 ’, 45.227 2, − 108.585 6, 13 - Aug- 1997, Chuck & Chris Harp leg. (Fig. 21 c) and 1 ♀ NVG- 24126 B 12 25 mi SW of Lodge Grass, 6 - Sep- 1971, Jack Harry leg.; 1 ♂ NVG- 24126 C 01 Wyoming, Big Horn Co., Red Grade Spring, 5000 ’, 13 - Aug- 1954; and South Dakota: 1 ♂ NVG- 24127 A 08 Butte Co., USH 212 at mi 17, ca. 2 mi E of Belle Fourche, 3 - Aug- 1983, D. L. Eiler leg. (Fig. 21 d) and 1 ♂ NVG- 24127 A 07 Lawrence Co., no locality details, 27 - Aug- 1969, M. L. May. Type locality. USA: Wyoming, Washakie Co., Bighorn Mountains, Tensleep Preserve, elevation 6400 ft.	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
42116960602FB325FE9B245159E2BD29.taxon	etymology	Etymology. The name reflects a superficial resemblance between O. napa and this new subspecies of O. sylvanoides, as it parallels O. napa but occurs at more northern latitudes. The name is treated as a noun in apposition.	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
42116960602FB325FE9B245159E2BD29.taxon	distribution	Distribution. Currently known from east of the main Rocky Mountain chain in Montana and Wyoming, and from South Dakota.	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
42116960602DB31BFEA2261D5A08BEBC.taxon	description	http: // zoobank. org / F 27 AE 8 D 7 - F 5 DA- 4 B 7 F-B 444 - A 3343470021 A (Figs. 19 part, 20 part, 21 e – f)	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
42116960602DB31BFEA2261D5A08BEBC.taxon	materials_examined	Type material. Holotype: ♂ deposited in the McGuire Center for Lepidoptera and Biodiversity Collection, Gainesville, FL, USA (MGCL), illustrated in Fig. 21 e, bears the following five printed rectangular labels, four white: [ND: Slope Co. elev 2560 ' | East River Road, approximately | 1.75 miles from Burning Coal | Vein Campground 46 ° 35 ' 51.1 " | N, 103 ° 27 ' 24.5 " W August 23, | 2023 Leg: W. R. Dempwolf], [Ochlodes sylvanoides | napa | ♂ | Coll of: W R Dempwolf], [DNA sample ID: | NVG- 23032 D 05 | c / o Nick V. Grishin], [WRD 23,554], and one red [HOLOTYPE ♂ | Ochlodes sylvanoides | dempwolfi Grishin]. Paratypes: 1 ♂ and 3 ♀♀: from USA, North Dakota, Slope Co. data as the holotype except as indicated, [WRD]: 1 ♂ NVG- 23032 D 06, WRD 23555 22 - Aug- 2023, 1 ♀ NVG- 23032 D 02, WRD 23551; 1 ♀ NVG- 23032 D 03; WRD 23552 (Fig. 21 f); and 1 ♀ NVG- 23032 D 04, WRD 23553 East River Road, ca. 18 mi south of Medora, 2647 ’, 46.698 861, − 103.487 417, 22 - Aug- 2023. Other specimens: Due to genetic similarity (Fig. 20), we currently attribute the following three sequenced specimens from Canada in the CNC to this subspecies but exclude them from the type series because they exhibit some differences compared to the population at the type locality: Alberta: 1 ♂ NVG- 24014 A 07, CNCLEP 00169968 Taber, 49.787 3, − 112.149 0, 16 - Aug- 1996, T. Pike leg. and 1 ♀ NVG- 24014 A 06, CNCLEP 00169953 Hwy 880 & US border, 49.000 0, − 111.266 7, 16 - Aug- 1998, R. A. Layberry leg. and 1 ♂ NVG- 24014 A 05, CNCLEP 00169950 Saskatchewan, Val Marie, 49.246 4, − 107.728 3, 10 - Aug- 1983, R. Hooper leg. Type locality. USA: North Dakota, Slope Co., East River Road, ca. 1.75 mi from Burning Coal Vein Campground, elevation 2560 ’, GPS 46.597 5, − 103.4568.	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
42116960602DB31BFEA2261D5A08BEBC.taxon	etymology	Etymology. The name, a noun in the genitive case, honors Bill Dempwolf, the collector of the type series and a friend of the author. An exceptional lepidopterist, Bill is recognized for meticulously curating and assembling one of the finest collections. His generosity has been instrumental in our genomic studies, manifested through both dedicated specimen collection for our lab research and open access to his entire collection for leg sampling and sequencing. His profound contributions to our projects are highly significant and greatly appreciated.	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
42116960602DB31BFEA2261D5A08BEBC.taxon	distribution	Distribution. Northeastern parts of the range, east of the Rockies from Alberta and Saskatchewan in Canada to North Dakota in the U. S.	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606012B31AFE50247C5D67BC8F.taxon	description	http: // zoobank. org / 8 E 47 C 852 - 82 A 8 - 4841 - B 2 E 0 - 85 DE 20 E 52512 (Figs. 22 part, 23)	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606012B31AFE50247C5D67BC8F.taxon	materials_examined	Type material. Holotype: ♂ deposited in the Carnegie Museum of Natural History, Pittsburgh, PA, USA (CMNH), illustrated in Fig. 23, bears the following five printed rectangular labels, four white: [Rio Grande | do Sul], [692.], [Lindsay Collection | C. M. Acc. No. 8584], [DNA sample ID: | NVG- 23109 G 03 | c / o Nick V. Grishin], and one red [HOLOTYPE ♂ | Thespieus | grandosul Grishin]. Type locality. Brazil: Rio Grande do Sul.	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606012B31AFE50247C5D67BC8F.taxon	etymology	Etymology. The name is given for the type locality, [Rio] Gran [de] + do + Sul, and is treated as a noun in apposition.	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
421169606012B31AFE50247C5D67BC8F.taxon	distribution	Distribution. Currently known only from the holotype collected in Rio Grande do Sul, Brazil.	en	Zhang, Jing, Cong, Qian, Shen, Jinhui, Song, Leina, Grishin, Nick V. (2025): Update to: Advancing butterfly systematics through genomic analysis. The Taxonomic Report of the International Lepidoptera Survey 12 (6): 1-36, DOI: 10.5281/zenodo.16570612
