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Modica Zacca, Casagrande & Willmott, genus novum. Type species — Euptychia confusa Staudinger, 1887, by present designation. Zoobank registration: https://zoobank.org/Nomenclatural Acts/D3CE81CE-5587-41D5-AFB6-934343C11D6F Systematic placement and diagnosis. Modica gen.n. is a member of the ‘ Splendeuptychia clade’ (Figure 10), in which its monophyly is strongly supported (FULL dataset SH-aLRT =100, UFB = 100). Its relationships to other members of the clade, however, are not clear; the genus is placed as sister to Parypthimoides, but support is inconclusive (FULL dataset SH-aLRT 100, UFB 43), and is further found within the same strongly supported clade that also contains Deltaya gen.n. (see under that genus), Scriptor, Emeryus, Colombeia and Malaveria (FULL dataset SH-aLRT 100, UFB 95). As with Deltaya gen.n., no morphological synapomorphies were identified for Modica gen.n., but these two genera share the same somewhat distinctive characters within Euptychiina (see under the former genus). Overall, the genus is best distinguished from phenotypically similar genera by the pale pupils of the HW ocellus in Cu 2 -Cu 1 being visible on the dorsal surface, by the VHW postdiscal ocelli in cells Cu 1 -M 3 and M 3 -M 2 being of similar size, round (not elongate) and each containing two enlarged, elongated silver pupils, and by the marginal line not widening in the VHW tornus (Figures 37 and 38). The genitalia of both sexes (Figure 39) are described below, and, as with Deltaya gen.n., they are broadly similar to those of a number of more or less distantly related euptychiine genera, in other words lacking obvious synapomorphies. The eighth abdominal segment of the female is only slightly pleated and expandable, similar to Scriptor and some related species but differing from others in which it is fully pleated and expandable, with this character varying within genera (e.g. within Paryphthimoides and Deltaya gen.n.). The female genitalia also lacks a sclerotized lamella antevaginalis and there is no sclerotized plate present on the ventral intersegmental membrane of the seventh and eighth abdominal segments, unlike Scriptor and some species of Deltaya gen.n. Characters that differ among the genera within the clade in which Modica gen.n. is placed are summarized in Table 1. Etymology. The generic name is a feminine Latin adjective treated as a noun in the nominative singular, meaning something that is modest, ordinary, average, in reference to the ‘typical’ euptychiine morphology of this genus and its lack of obvious distinguishing characters. Description (Figures 37 – 39). Some notable characters include: eyes setose; pterothoracic legs dorsally slightly darker, tibia with two principal longitudinal rows of spines ventrally, pair of spurs of similar length at distal end of tibia, first tarsomere with three principal longitudinal rows of spines ventrally, remaining tarsomeres with four principal longitudinal rows of spines ventrally. Medium-sized Euptychiina (FW length typically 21 – 28 mm), FW triangular and rather rounded at apex, HW rounded. No strong sexual dimorphism: Dorsal wings dark brown to greyish brown, pale pupils of ocellus in cell Cu 2 -Cu 1 visible on DHW, no androconial scales present. Ventral wings greyish brown to yellowish brown; relatively broad, dark brown to reddish brown discal and postdiscal lines traversing both wings; VFW with three postdiscal ocelli in cells Cu 1 -M 3, M 3 -M 2, M 2 -M 1, anterior ocellus more clearly marked than remainder, lying within a broad dark brown band (umbra), with this band and adjacent area basally in middle of wing tinged yellowish in M. myncea comb.n. and M. confusa comb.n.; VHW similar to VFW but with five postdiscal ocelli between Cu 2 and Rs, those in cells Cu 2 -Cu 1 and M 2 -M 1 typically slightly larger, black-centred with two silver dots in each ocellus as pupils, ocellus in M 1 -Rs similar but much smaller, and those in cell Cu 1 -M 2 with dark brown centres and elongate silver pupils; marginal line thin and even throughout, not thickening at tornus. Male 8th abdominal tergite reduced dorsally, leaving a sclerotized strip along anterior edge and usually an isolated sclerotized patch in posterior portion. Male genitalia with uncus longer than tegumen, brachia approximately parallel with uncus and about two-thirds its length; valvae elongate with dorsal edge straight or with slight projection; aedeagus adorned with scattered small spines in some species, and with or without cornuti. Female genitalia has 8th tergite reduced to a posterior sclerotized patch about two-thirds width of segment, intersegmental membrane between seventh and eighth abdominal segments only somewhat expandable with no strongly sclerotized plate ventrally, eighth segment with large irregular lateral sclerotized plate extending further dorsally at anterior edge, lamella antevaginalis and antrum unsclerotized, ductus bursae unsclerotized, corpus bursae small, oval and with two narrow sub-parallel signa. Distribution and natural history (Figure 40). Modica gen.n. contains five described species and several undescribed species (Zacca et al., unpublished data), which occur in rainforest from sea level to 1300 m, ranging from southern Mexico to western Ecuador and throughout the Amazon and Guianas to south-eastern Brazil. The genus reaches its peak diversity in the western Amazon, where both sexes may be common throughout the understory of both disturbed and undisturbed forest, with some species also occurring along forest edges and in overgrown, shady plantations. Males of some species perch from 1 to 3 m in the forest understory in the morning and late afternoon, sometimes on hilltops, and both sexes are attracted to rotting fruit (DeVries, 1987; Zacca et al., pers. obs.). Notes on the immature stages of M. myncea comb.n. and M. confusa comb.n. were provided by Singer et al. (1983), with hostplants (natural and in captivity) including Cyperaceae, Palmae, Poaceae and Marantaceae (see also Beccaloni et al., 2008; Singer & Ehrich, 1993). Discussion. The type species for this genus, Euptychia confusa, was described by Staudinger (1884) based on an unstated number of specimens from Chiriquí, Panama, and a lectotype at the MfN was designated by Singer et al. (1983). The female illustrated in Staudinger (1884: pl. 80) agrees with the description, although it was incorrectly labelled as ‘ Euptychia myncea ’. We chose to designate Euptychia confusa as the type species for this genus since DNA barcode data suggest the possibility of cryptic species within Modica myncea comb.n. that remain to be resolved. Although strongly supported by the molecular data, Modica gen.n. lacks any clear morphological synapomorphies and it is thus not surprising that a close relationship among all of its constituent species was unnoticed until recently; M. confusa comb.n., M. myncea comb.n. and M. maripa comb.n. were hitherto placed in Cissia (e.g. Forster, 1964 [under the name Argyreuptychia]; Singer et al., 1983; Lamas, 2004; Brévignon, 2005) and M. fugitiva comb.n. and M. kamel comb.n. were placed in Magneuptychia (Benmesbah et al., 2018; Forster, 1964; Lamas, 2004). As discussed by Zacca, Casagrande, et al. (2018), most species placed in Cissia prior to that paper were presumably considered to be related because of their possession of a yellowish patch on the VFW, but that character is clearly homoplasious, and the Cissia of Lamas (2004) are now placed in six genera in two clades: Cissia, Vanima, Megisto, Modica, Paryphthimoides, and Vareuptychia. Benmesbah et al. (2018) noted that M. myncea comb.n., M. maripa comb.n., M. fugitiva and M. kamel comb.n. were likely closely related, based on morphology and preliminary molecular data, and suggested that a new genus might be needed to accommodate these species. Our phylogenetic analysis failed to strongly resolve the relationships of Modica gen.n. to other clades. These species therefore cannot reasonably be accommodated in any described genus without combining at least five described genera, along with Modica gen.n. and Deltaya gen.n., into a single, large genus of morphologically and ecologically dissimilar species, which would not correspond to any author ’ s prior taxonomic hypothesis. Modica Zacca, Casagrande & Willmott, gen.n. confusa (Staudinger, 1887), comb.n., was Cissia fugitiva (Lamas [1997]), repl. name, comb.n., was Magneuptychia = helle (Cramer, 1779), preocc. (not [Denis & Schiffermüller], 1775), comb.n., was Magneuptychia kamel (Benmesbah & Zacca, 2018), comb.n., was Magneuptychia [Benmesbah et al. (2018, Zootaxa, 4425(1): 115-145)] maripa (Brévignon, 2005), comb.n., was Cissia [Brévignon (2005, Lambillionea, 105(3)(1): 393-404)] myncea (Cramer, 1780), comb.n., was Cissia = myncena (Stoll, 1782), missp., comb.n., was Cissia = crantor (Fabricius, 1793), comb.n., was Cissia = clerica (Herrich-Schäffer, 1865), repl. name, comb.n., was Cissia = pytheus (MÖschler, 1883), comb.n., was Cissia = isolata (Kaye, 1921), comb. n., was Cissia [Kaye (1921, Memoirs of the Department of Agriculture of Trinidad and Tobago, 2: i-xii, 13-163, 1 pl.)], Published as part of Espeland, Marianne, Nakahara, Shinichi, Zacca, Thamara, Barbosa, Eduardo P., Huertas, Blanca, Marín, Mario A., Lamas, Gerardo, Benmesbah, Mohamed, Brévignon, Christian, Casagrande, Mirna M., Fåhraeus, Christer, Grishin, Nick, Kawahara, Akito Y., Mielke, Olaf H. H., Miller, Jacqueline Y., Nakamura, Ichiro, Navas, Vanessa, Patrusky, Brooke, Pyrcz, Tomasz W., Richards, Lindsay, Tan, Denise, Tyler, Stephanie, Viloria, Angel, Warren, Andrew D., Xiao, Lei, Freitas, André V. L. & Willmott, Keith R., 2023, Combining target enrichment and Sanger sequencing data to clarify the systematics of the diverse Neotropical butterfly subtribe Euptychiina (Nymphalidae, Satyrinae), pp. 1-73 in Zoological Research 2023 on pages 42-67, DOI: 10.1111/syen.12590, http://zenodo.org/record/7909395, {"references":["DeVries, P. J. (1987) The butterflies of Costa Rica and their natural history: papilionidae, Pieridae, Nymphalidae. Princeton: Princeton University Press, p. 288.","Singer, M. C., DeVries, P. J. & Ehrlich, P. R. (1983) The Cissia confusa species-group in Costa Rica and Trinidad (Lepidoptera: Satyrinae). Zoological Journal of the Linnean Society, 79, 101 - 119.","Beccaloni, G. W., Viloria, A. L., Hall, S. K. & Robinson, G. S. (2008) Catalogue of the hostplants of the Neotropical butterflies. Monografias Tercer Milenio. Sociedad Entomologica´Aragonesa (SEA) / Red Iberoamericana de Biogeografia y Entomologia Sistematica (RIBES) / Ciencia y Tecnologia para el Desarrollo (CYTED) / Natural History Museum, London, U. K. (NHM) / Instituto Venezolano de Investigaciones Cientificas, Venezuela (IVIC), Zaragoza, p. 536.","Singer, M. & Ehrich, P. (1993) Host specialization of satyrine butterflies, and their responses to habitat fragmentation in Trinidad. Journal of Research on the Lepidoptera, 30, 248 - 256.","Staudinger, O. (1884) I. Theil. Exotische Tagfalter in systematischer Reihenfolge mit Berucksichtigung neuer Arten. In: Staudinger, O. & Schatz, E. (Eds.) Exotische Schmetterlinge. Furth: G. Lowensohn, pp. 1 - 333.","Forster, W. (1964) Beitrage zur Kenntnis der Insektenfauna Boliviens XIX. Lepidoptera III. Satyridae. Veroffentlichungen der Zoologischen Staatssammlung Munchen, 8, 51 - 188.","Lamas, G. (2004) Atlas of Neotropical Lepidoptera. Checklist: Part 4 A Hesperioidea - Papilionoidea. Gainesville: Scientific Publishers / Association of Tropical Lepidoptera, p. 439.","Brevignon, C. (2005) Description de nouveaux Satyrinae provenant de Guyane franCaise (Lepidoptera, Nymphalidae). Lambillionea, 105, 393 - 404.","Benmesbah, M., Zacca, T., Casagrande, M. M., Mielke, O. H. H., Lamas, G. & Willmott, K. R. (2018) Taxonomic notes on Papilio ocypete Fabricius, 1776 and Papilio helle Cramer, 1779 with description of two new similar species from South America (Lepidoptera: Nymphalidae: Satyrinae). Zootaxa, 4425, 115 - 145."]}

Occulta Nakahara & Willmott, genus novum. Type species: Euptychia ocnus Butler, 1867, by present designation. Zoobank registration: https://zoobank.org/Nomenclatural Acts/7B2BBC71-C2FC-40B8-A24A-14A986151D85 Systematic placement and diagnosis. Occulta gen.n. is a member of the ‘ Splendeuptychia clade’, and it is rather distantly related to the remainder of the clade. Currently, Occulta gen.n. is modestly supported (SH-aLRT 100, UFB 75) as sister to a large clade including species-rich genera such as Caeruleuptychia, Paryphthimoides, and Splendeuptychia, among others (Figure 10). Based on hybrid enrichment data, Espeland et al. (2019a) recovered Occulta gen.n. (as ‘ Magneuptychia ocnus ’) as sister to ‘ Erichthodes ’ narapa (now Capronnieria narapa comb.n.) and Capronnieria galesus, although with only moderate support (BS and PP>0.75 Occulta gen.n. (Figure 30) resembles some species of Deltaya gen.n. (see Figures 33 and 34 below), but may be distinguished by possessing elongate, ‘smudge-like’ orangish spots with a silver marking in the middle in VHW cells M 2 and M 3, whereas the ocelli in these cells in Deltaya gen.n. are more rounded and have double silver spots encircled by a better defined yellowish ring. Furthermore, the dark ventral discal and postdiscal bands are typically narrower in Occulta gen.n. compared to Deltaya gen.n. species, perhaps with the exception of D. ocypete, whose ventral bands are often narrower in comparison with other closely related species. In particular, in Occulta gen.n. the VHW marginal band is of even width throughout, whereas this band forms a ‘wedge-shaped’ swelling at the tornus in Deltaya gen.n. In addition, Occulta gen.n. possesses a tiny, rather incomplete ocellus in VHW cell Cu 2, which is absent in Deltaya gen.n. The male genitalia also differ between Occulta gen.n. (Figure 31) and the type species of Deltaya gen.n. and close relatives (referred to as ‘core’ Deltaya gen.n.) (Figure 34), at least, by the lack of a developed ‘hump’ on the dorsal margin of the valva, and instead only have a slightly serrated region at the dorsal margin distal of the costa. In lateral view, the costa appears as a narrow plate in Occulta gen.n., whereas the costa appears as a somewhat trapezoidal plate in lateral view in ‘core’ Deltaya gen.n. See description of Deltaya gen.n. below for details about ‘core’ Deltaya gen.n. Etymology. The generic name is a Latin feminine adjective treated as a noun in the nominative singular, ‘occulta’, meaning a ‘hidden’ or ‘secret’ thing, in reference to the former concealment of this taxon within Magneuptychia. The generic name is also coined in alliteration with the species-group name. Description (Figures 30 and 31). Wing pattern and shape as illustrated (Figure 30). Other notable characters include eyes setose; pterothoracic legs dorsally darker, tibia with two principal longitudinal rows of spines ventrally, in addition to some spines laterally, pair of spurs of similar length at distal end of tibia, first tarsomere with three principal longitudinal rows of spines ventrally, remaining distal tarsomeres with four principal longitudinal rows of spines ventrally. Medium-sized Euptychiina (FW length typically 23 – 26 mm), DFW and DHW of males without obvious androconial scales, but long setiform scales visible in discal cell and adjacent areas, especially extending along inner margin. Male with 8th tergite a narrow stripe at basal side of the eighth abdominal segment, broad weakly sclerotized patch present at posterior side of eighth abdominal segment; genitalia as illustrated (Figure 31); costa appearing as a narrow plate in lateral view; cornuti present, otherwise genitalia as illustrated (Figure 31). Female genitalia (Figure 31) with intersegmental membrane of between seventh and eighth abdominal segment somewhat pleated and expandable with very weak sclerotization ventrally; lamella antevaginalis membranous; eighth segment with irregular lateral sclerotized plate narrowing slightly dorsally; pair of well-defined signa present. Occulta Nakahara & Willmott, gen.n. ocnus (Butler, 1867), comb.n., was Magneuptychia = gracilis (Weymer, 1911), comb.n., was Magneuptychia, Published as part of Espeland, Marianne, Nakahara, Shinichi, Zacca, Thamara, Barbosa, Eduardo P., Huertas, Blanca, Marín, Mario A., Lamas, Gerardo, Benmesbah, Mohamed, Brévignon, Christian, Casagrande, Mirna M., Fåhraeus, Christer, Grishin, Nick, Kawahara, Akito Y., Mielke, Olaf H. H., Miller, Jacqueline Y., Nakamura, Ichiro, Navas, Vanessa, Patrusky, Brooke, Pyrcz, Tomasz W., Richards, Lindsay, Tan, Denise, Tyler, Stephanie, Viloria, Angel, Warren, Andrew D., Xiao, Lei, Freitas, André V. L. & Willmott, Keith R., 2023, Combining target enrichment and Sanger sequencing data to clarify the systematics of the diverse Neotropical butterfly subtribe Euptychiina (Nymphalidae, Satyrinae), pp. 1-73 in Zoological Research 2023 on pages 35-67, DOI: 10.1111/syen.12590, http://zenodo.org/record/7909395, {"references":["Butler, A. G. (1867) Descriptions of some new species of Satyridae belonging to the genus Euptychia. Proceedings of the Zoological Society of London, 1867, 104 - 110.","Espeland, M., Breinholt, J., Barbosa, E. P., Casagrande, M., Huertas, B., Lamas, G. et al. (2019 a) Four hundred shades of brown: higher level phylogeny of the problematic Euptychiina (Lepidoptera, Nymphalidae, Satyrinae) based on hybrid enrichment data. Molecular Phylogenetics and Evolution, 131, 116 - 124.","Weymer, G. (1911) 4. Familie: Satyridae. In: Seitz, A. (Ed.) Die GrossSchmetterlinge der Erde. Stuttgart: A. Kernen, pp. 173 - 280."]}

Cissia Doubleday, 1848 = Argyreuptychia Forster, 1964 anabelae (L.D. Miller, 1976) [Zhang et al. (2022, The Taxonomic Report of the International Lepidoptera Survey, 10(7): 1-60)] cheneyorum (Chermock, 1949) [Zhang et al. (2022, The Taxonomic Report of the International Lepidoptera Survey, 10(7): 1-60)] cleophes (Godman & Salvin, 1889) [Zhang et al. (2020, The Taxonomic Report of the International Lepidoptera Survey, 8(7): 1-40)] eous (Butler, 1867) [Zacca et al. (2018, Arthropod Systematics and Phylogeny, 76(2): 349-376)] = eoüs (Butler, 1867), missp. [Zacca et al. (2018, Arthropod Systematics and Phylogeny, 76(2): 349-376)] = kiliani (Anken, 1999) [Zacca et al. (2018, Arthropod Systematics and Phylogeny, 76(2): 349-376)] penelope (Fabricius, 1775) = clarissa (Cramer, 1780) = moneta (Weymer, 1911) phronius (Godart, [1824]) [Zacca et al. (2018, Arthropod Systematics and Phylogeny, 76(2): 349-376)] = punctatolineata (J. Zikán & W. Zikán, 1968), nom. nud. [Lamas, unpublished data] pompilia (C. Felder & R. Felder, 1867) = usitata (Butler, 1867) = pieria (Butler, 1867) = austera (Butler, 1867) [Zacca et al. (2018, Arthropod Systematics and Phylogeny, 76(2): 349-376)] = thelete (Snellen, 1887) proba (Weymer, 1911) = mariameliae (Hayward, 1957) pseudocleophes (L.D. Miller, 1976) [Zhang et al. (2022, The Taxonomic Report of the International Lepidoptera Survey, 10(7): 1-60)] rubricata (Edwards, 1871) [Zacca et al. (2018, Arthropod Systematics and Phylogeny, 76(2): 349-376)] - smithorum (Wind, 1946) [Zacca et al. (2018, Arthropod Systematics and Phylogeny, 76(2): 349-376)] wahala Grishin, 2022 [Zhang et al. (2022, The Taxonomic Report of the International Lepidoptera Survey, 10(7): 1-60)], Published as part of Espeland, Marianne, Nakahara, Shinichi, Zacca, Thamara, Barbosa, Eduardo P., Huertas, Blanca, Marín, Mario A., Lamas, Gerardo, Benmesbah, Mohamed, Brévignon, Christian, Casagrande, Mirna M., Fåhraeus, Christer, Grishin, Nick, Kawahara, Akito Y., Mielke, Olaf H. H., Miller, Jacqueline Y., Nakamura, Ichiro, Navas, Vanessa, Patrusky, Brooke, Pyrcz, Tomasz W., Richards, Lindsay, Tan, Denise, Tyler, Stephanie, Viloria, Angel, Warren, Andrew D., Xiao, Lei, Freitas, André V. L. & Willmott, Keith R., 2023, Combining target enrichment and Sanger sequencing data to clarify the systematics of the diverse Neotropical butterfly subtribe Euptychiina (Nymphalidae, Satyrinae), pp. 1-73 in Zoological Research 2023 on page 56, DOI: 10.1111/syen.12590, http://zenodo.org/record/7909395

Argentaria Huertas & Willmott, genus novum. Type species — Euptychia itonis Hewitson, 1862, by present designation. Zoobank registration: https://zoobank.org/Nomenclatural Acts/F5A5D4DD-82D3-40AA-BD88-978B9A55AB46 Systematic placement and diagnosis. Argentaria gen.n. is a member of the ‘ Amphidecta clade’, in which its monophyly is strongly supported in all datasets (FULL dataset, SH-aLRT 100, UFB 100, Figure 6). However, its relationships to other members of the clade, which include (among described species) Zischkaia, Amphidecta, ‘ Euptychia ’ ordinata / insignis, and ‘ Pharneuptychia ’ innocentia, are not strongly resolved. The new genus can be distinguished from other Euptychiina by the combination of the following three characters: (a) the VFW has a postdiscal series of ocelli extending from cell 2A-Cu 2 to M 2 -M 1, with these ocelli similar in form and elongate parallel to the wing margin, often with either a linear silvery pupil or double pupils in each cell (Figure 17). In some individuals of A. salvini comb.n., the ocelli are fused into a continuous band and the pupils absent, thus superficially resembling the dark postdiscal band, which surrounds the postdiscal ocelli in other species (e.g. Magneuptychia libye). (b) the VHW has the postdiscal ocellus in cell M 3 -M 2 elongated parallel to the veins, with the silvery pupil divided into two distinct spots, except in A. clementia comb.n., in which the two spots touch or are partly fused (Figure 17). A few other species have an elongated pupil similar to that in A. clementia comb.n., such as Splendeutychia purusana comb.rev. or Nhambikuara furina comb.n. (c) the VHW tornus in cell 2A-Cu 2 has several silver streaks, which appear to represent the pupils of ocelli in both the anterior and posterior half of the cell, with the former often being split into two or elongated and pinched in the middle, while the latter is visible as an additional silver spot in some species (e.g. A. itonis comb.n.) (Figures 18 and 19). Other genera with an elongate ocellus and silver pupil in cell 2A-Cu 2, such as Nhambikuara, have only a single pupil that is not as elongate as in Argentaria gen.n. The genitalia of both sexes (Figure 19) are described below and are broadly like those of a number of other relatively distantly related euptychiine genera (e.g. Amiga, Pseudeuptychia, Scriptor). Notable features include, in the male, the lack of cornuti in the aedeagus (similar to many other euptychiine genera), and in the female, the pleated, expandable intersegmental membrane between the seventh and eighth abdominal segments, antrum and lamella antevaginalis membranous, and small, round corpus bursae with slender, elongate, converging signa. The other two described genera within the ‘ Amphidecta clade’, namely Amphidecta and Zischkaia, each have distinctive male and female genitalia differing in numerous respects from Argentaria gen.n., with these distinctive characters apparently representing generic autapomorphies (Marín et al., 2017; Nakahara, Zacca, et al., 2019; Nakahara pers. obs.). Etymology. The generic name is a feminine Latin noun in the nominative singular, meaning a silver-mine, in reference to the distinctive silver scales arranged as spots on the ventral hindwing and forewing of species in this genus. Description (Figures 17 – 19). Some notable characters include: eyes naked; pterothoracic legs dorsally slightly darker, tibia with two principal longitudinal rows of spines ventrally, pair of spurs of similar length at distal end of tibia, first tarsomere with three principal longitudinal rows of spines ventrally, remaining tarsomeres with four principal longitudinal rows of spines ventrally. Small to medium-sized Euptychiina (FW length typically 15 – 25 mm), FW triangular and varying from being rather pointed (e.g. A. itonis comb.n.) to more rounded (e.g. A. quadrina comb.n.), HW rounded with margin variably undulating. No strong sexual dimorphism in most species: Dorsal wings dark grey brown, some species with white patches in discal part of DFW, and white patches in basal or discal part of DHW and yellowish or orangish distally, no androconial scales present except in male A. libitina comb.n., which has dark androconial scales lining the DFW cubital vein and basal part of vein Cu 2, and basal part of vein 2A. Ventral wings showing relatively large inter-specific variation in pattern across the genus, with ground colour ranging from dark grey-brown to paler yellowish brown; a dark discal and postdiscal line traversing both wings, ranging from a thin, irregular line to a straight, broad band, which may be absent or scarcely visible in some species; discal area of one or both wings often white to pale yellowish brown, sometimes extending basally or distally beyond postdiscal line; VFW with series of narrow postdiscal ocelli in cells 2A-Cu 2 to M 2 -M 1, elongated parallel to distal margin, usually with pupil present as single silver line or double silver spots, often fused into a broad band, variably surrounded by orange or yellow and in some species a further black border, ocelli variably distinct across species; VHW with a complex array of postdiscal ocelli in cells 2A-Cu 2 to M 1 -Rs, with that in Cu 2 -Cu 1 typically the largest, and, along with that in M 2 -M 1, often the only ocellus to be present as a black spot with a single, double, or further modified silver pupil, remaining ocelli typically lacking black with their silver pupils enlarged into spots or streaks, orange rings around ocelli expanded and fused in all species to form an orange background around the ocelli; silver pupil in cell M 3 -M 2 elongated and pinched in the middle (A. clementia comb.n.) or split into two spots (all other species), and that in cell 2A-Cu 2 similarly pinched or split, often forming three distinct spots in combination with a silver spot presumed to represent a posterior ocellus in that cell; two black submarginal lines parallel to wing margin with pale scaling between them, sometimes pierced by elongate silver pupils of the postdiscal ocelli. Male 8th abdominal tergite reduced dorsally, leaving a sclerotized strip along anterior edge and usually an isolated sclerotized patch in posterior portion, that can be broader (e.g. A. jadea comb.n.) or absent (e.g. A. pagyris comb.n.) in some species. Male genitalia with uncus longer than tegumen, brachia pointing dorsally of uncus or parallel with uncus and ranging from slightly longer than uncus (e.g. A. quadrina comb.n.) to very short/vestigial (e.g. A. kendalli comb.n., A. jadea comb.n., A. clorimena comb.n.); valva usually broadening at distal tip in lateral view, with distal tip sometimes with an inwardly pointing dorsal lobe (e.g. A. itonis, A. clementia, A. quadrina) and sometimes without (e.g. A. libitina comb.n., A. kendalli comb.n., A. clorimena comb.n.); vinculum in most species (but not all, e.g., A. pagyris) elongated above articulation with dorsal base of valva; aedeagus relatively straight and lacking cornuti. Female genitalia showing relatively little inter-specific variation, eighth tergite with small sclerotized posterior patch and thin strip at ventral anterior edge, intersegmental membrane between segments A7 and A8 pleated and expandable with a circular to quadrate sclerotized plate ventrally, eighth segment with large irregular oval lateral sclerotized plate, lamella antevaginalis and antrum unsclerotized, ductus bursae narrow and unsclerotized, corpus bursae small, circular and with two narrow, posteriorly converging signa. Distribution and natural history (Figure 20). Argentaria gen.n. species are found in rainforest and cloudforest from sea level to 2100 m, although the great majority of species are found below 1200 m. Similarly, although the genus occurs throughout the Neotropical region, only a couple of species occur west of the Andes, and the known centre of diversity is in the bamboo-rich forests of the south-western Amazon (Madre de Dios, Peru, and Acre, Brazil), where up to 10 species can be found in sympatry, and to a lesser extent in south-eastern Brazil (Atlantic region). All species occur in close association with bamboos (Poaceae: Bambusoideae: Bambuseae), including Guadua and various species of weeping bamboos (unpublished data), and many species are known from only a handful of widely scattered localities, with a number of species still undescribed. The immature stages were recently described for A. quadrina comb.n. by See et al. (2018), which was recorded feeding on the climbing bamboo Rhipidocladum racemiflorum with several other species having been reared in south-eastern Brazil (Freitas, 2022) and Ecuador (Willmott & Hall, unpublished data). Discussion. The type species for this genus, Euptychia itonis, was described by Hewitson (1862) from the ‘Amazon’ based on specimens in his collection and the collection of W. W. Saunders. The underside of a specimen was illustrated in his Figure 3 and it closely matches the three syntypes examined in the NHMUK, in particular in having a thin, broken dark brown postdiscal line in the white area of the VHW, which is unique within the genus to A. itonis comb.n. Butler (1867) placed all of the species included here in Argentaria gen.n. that were described at that time in his Division V, Subdivision 2, which he characterized as having the VHW ocelli represented as silver streaks with a large black tornal ocellus, with his correct recognition of the close relationships of the species in this group reflecting the distinctive VHW pattern of the genus. Weymer (1911) likewise successfully grouped described Argentaria gen.n. species in his ‘Clorimene [sic] group’, which he placed alongside his ‘Doxes group’, which corresponds to Nhambikuara. Forster ’ s (1964) decision to move Argentaria gen.n. species from Euptychia into his newly described genus Splendeuptychia, with type species Euptychia ashna, seems somewhat surprising, since Weymer (1911) evidently did not regard E. ashna as being closely related to the former two groups. Equally puzzling are Forster ’ s (1964) drawings of the male genitalia of A. clementia comb.n. and A. pagyris comb.n. (p. 129, figure 163, 164), which are rather different to specimens we have examined, most notably in showing an aedeagus with cornuti, which was not observed in any dissected Argentaria gen.n. In general, Forster (1964) separated numerous taxa from the genus Euptychia into different genera, based on wing pattern and male genitalia, but he also frequently grouped species by overall appearance. Lamas (2004) followed Forster ’ s arrangement, and retained in Splendeuptychia the species placed here in Argentaria gen.n., but shortly afterwards Murray and Prowell (2005) showed that, based on DNA sequence data, A. itonis comb.n. and S. ashna were distantly related. Subsequent molecular (Espeland et al., 2019a; Peña et al., 2010) and morphological (Huertas, 2014; Marín et al., 2017) phylogenetic studies have corroborated that discovery, including this study, which contains sequence data for all but three of the described species. The relationships of Argentaria gen.n. to other members of the ‘ Amphidecta clade’ are not strongly resolved, but the numerous morphological differences that separate the genus from other genera within the clade, and the fact that its constituent species have universally been recognized as closely related for more than 150 years, support its recognition as a distinct genus. Argentaria Huertas & Willmott, gen.n. argyropsacas (Bryk, 1953), comb.n., stat.rev., was Splendeuptychia [treated as a synonym of " Splendeuptychia " telesphora by Lamas (2004), but ventral wing pattern is distinct] clementia (Butler, 1877), comb.n., was Splendeuptychia clorimena (Stoll, 1790), comb.n., was Splendeuptychia = boliviensis (Forster, 1964), comb.n., syn.n., was Splendeuptychia [wing pattern variation and DNA barcodes suggests this is a southern form of clorimena] cosmophila (Hübner, 1823), comb.n., was Splendeuptychia = argenteus (Swainson, 1823), comb.n., was Splendeuptychia hygina (Butler, 1877), comb.n., was Splendeuptychia itonis (Hewitson, 1862), comb.n., was Splendeuptychia jadea (Brévignon & Benmesbah, 2012), comb.n., was Splendeuptychia [Brévignon & Benmesbah (2012, Complément à l'inventaire des Satyrinae de Guyane (Lepidoptera: Nymphalidae), pp. 36-52, 4 pls., 1 tab. In: Lacomme, D. & L. Manil (Eds.), Lépidoptères de Guyane. Tome 7. Nymphalidae)] kendalli (L.D. Miller, 1978), comb.n., was Splendeuptychia libitina (Butler, 1870), comb.n., was Splendeuptychia pagyris (Godart, [1824]), comb.n., was Splendeuptychia = ava (Anken, 1998), comb.n., was Splendeuptychia quadrina (Butler, 1869), comb.n., was Splendeuptychia salvini (Butler, 1867), comb.n., was Splendeuptychia telesphora (Butler, 1877), comb.n., was Splendeuptychia zischkai (Forster, 1964), comb.n., was Splendeuptychia

Amiga Nakahara, Willmott & Espeland, 2019 arnaca (Fabricius, [1777]) [Nakahara et al. (2019, ZooKeys, 821: 85-152)] = ebusa (Cramer, 1780) [Nakahara et al. (2019, ZooKeys, 821: 85-152)] = arnaea (Fabricius, 1781), missp. = aranea (Fabricius, 1793), missp. = priamis (D’Almeida, 1922) [Nakahara et al. (2019, ZooKeys, 821: 85-152)] - adela (Nakahara & Espeland, 2019) [Nakahara et al. (2019, ZooKeys, 821: 85-152)] - indianacristoi (Nakahara & Marín, 2019), stat.rev. [raised from subspecies of A. arnaca to species by Orellana et al. (2020) based on unspecified morphological differences)/Orellana et al. (2020, Anartia, 29: 54-67)/Nakahara et al. (2019, ZooKeys, 821: 85-152)] sericeella (Bates, 1854) [Nakahara & Gallardo (2020, Tropical Lepidoptera Research, 29(2): 111-114)], Published as part of Espeland, Marianne, Nakahara, Shinichi, Zacca, Thamara, Barbosa, Eduardo P., Huertas, Blanca, Marín, Mario A., Lamas, Gerardo, Benmesbah, Mohamed, Brévignon, Christian, Casagrande, Mirna M., Fåhraeus, Christer, Grishin, Nick, Kawahara, Akito Y., Mielke, Olaf H. H., Miller, Jacqueline Y., Nakamura, Ichiro, Navas, Vanessa, Patrusky, Brooke, Pyrcz, Tomasz W., Richards, Lindsay, Tan, Denise, Tyler, Stephanie, Viloria, Angel, Warren, Andrew D., Xiao, Lei, Freitas, André V. L. & Willmott, Keith R., 2023, Combining target enrichment and Sanger sequencing data to clarify the systematics of the diverse Neotropical butterfly subtribe Euptychiina (Nymphalidae, Satyrinae), pp. 1-73 in Zoological Research 2023 on page 64, DOI: 10.1111/syen.12590, http://zenodo.org/record/7909395

Orotaygetis Nakahara & Zacca, 2018 surui Nakahara, Zacca & Lamas, 2018 [Nakahara et al. (2018, Insecta Mundi, 0639: 1-38)]

Taydebis Freitas, 2003 = Prenda Freitas & Mielke, 2011 clarissa (Freitas & Mielke, 2011) [Zacca et al. (2021, Zootaxa, 5023(4): 555-570)] guria Zacca, Casagrande & Mielke, 2021 [Zacca et al. (2021, Zootaxa, 5023(4): 555-570)] melobosis (Capronnier, 1874) [Zacca et al. (2021, Zootaxa, 5023(4): 555-570)] = peculiaris (Butler, 1874) [Zacca et al. (2021, Zootaxa, 5023(4): 555-570)] = fractifasciata (J. Zikán & W. Zikán, 1998), nom. nud., Published as part of Espeland, Marianne, Nakahara, Shinichi, Zacca, Thamara, Barbosa, Eduardo P., Huertas, Blanca, Marín, Mario A., Lamas, Gerardo, Benmesbah, Mohamed, Brévignon, Christian, Casagrande, Mirna M., Fåhraeus, Christer, Grishin, Nick, Kawahara, Akito Y., Mielke, Olaf H. H., Miller, Jacqueline Y., Nakamura, Ichiro, Navas, Vanessa, Patrusky, Brooke, Pyrcz, Tomasz W., Richards, Lindsay, Tan, Denise, Tyler, Stephanie, Viloria, Angel, Warren, Andrew D., Xiao, Lei, Freitas, André V. L. & Willmott, Keith R., 2023, Combining target enrichment and Sanger sequencing data to clarify the systematics of the diverse Neotropical butterfly subtribe Euptychiina (Nymphalidae, Satyrinae), pp. 1-73 in Zoological Research 2023 on page 65, DOI: 10.1111/syen.12590, http://zenodo.org/record/7909395

Atlanteuptychia Freitas, Barbosa & Mielke, 2013 ernestina (Weymer, 1911) [Freitas et al. (2013, Zoologia, 30: 661-668)]

Paramacera Butler, 1868 = Paramecera Butler, 1868, missp. allyni L.D. Miller, 1972 chinanteca L.D. Miller, 1972 copiosa L.D. Miller, 1972 xicaque (Reakirt [1867]) = epinephele (C. Felder & R. Felder, 1867) = conhiera Butler, 1868, nom. nud. rubrosuffusa L.D. Miller, 1972 [Zhang et al. (2022, The Taxonomic Report of the International Lepidoptera Survey, 10(7): 1-60)], Published as part of Espeland, Marianne, Nakahara, Shinichi, Zacca, Thamara, Barbosa, Eduardo P., Huertas, Blanca, Marín, Mario A., Lamas, Gerardo, Benmesbah, Mohamed, Brévignon, Christian, Casagrande, Mirna M., Fåhraeus, Christer, Grishin, Nick, Kawahara, Akito Y., Mielke, Olaf H. H., Miller, Jacqueline Y., Nakamura, Ichiro, Navas, Vanessa, Patrusky, Brooke, Pyrcz, Tomasz W., Richards, Lindsay, Tan, Denise, Tyler, Stephanie, Viloria, Angel, Warren, Andrew D., Xiao, Lei, Freitas, André V. L. & Willmott, Keith R., 2023, Combining target enrichment and Sanger sequencing data to clarify the systematics of the diverse Neotropical butterfly subtribe Euptychiina (Nymphalidae, Satyrinae), pp. 1-73 in Zoological Research 2023 on page 55, DOI: 10.1111/syen.12590, http://zenodo.org/record/7909395

Pseudeuptychia Forster, 1964 callichloris (Butler, 1867), comb.n., was Chloreuptychia [forms a well-supported clade with Pseudeuptychia languida, type species of Pseudeuptychia] cuzquenya Nakahara & Lamas, 2018 [Nakahara et al. (2018, Insecta Mundi, 0639: 1-38)] hemileuca (Staudinger, [1886]) herseis (Godart, [1824]), repl. name, comb.n., was Chloreuptychia [forms a well-supported clade with Pseudeuptychia languida, type species of Pseudeuptychia] = herse (Cramer, 1775), preocc. (not Hufnagel, 1766), comb.n., was Chloreuptychia = peruviana (J. Prüffer, 1922), comb.n., was Chloreuptychia = bellatula (D’Almeida, 1922), comb.n., was Chloreuptychia languida (Butler, 1871) - austrina Nakahara & Lamas, 2018 [Nakahara et al. (2018, Insecta Mundi, 0639: 1-38)] marica (Weymer, 1911), comb.n., was Chloreuptychia [forms a well-supported clade with Pseudeuptychia languida, type species of Pseudeuptychia] rectilinea (Brévignon, Rosant, Lamas & Willmott, 2019), comb.n., was Chloreuptychia [forms a well-supported clade with Pseudeuptychia languida, type species of Pseudeuptychia /Brévignon et al. (2019, Bulletin de la Société entomologique de France, 124(2): 127-138)], Published as part of Espeland, Marianne, Nakahara, Shinichi, Zacca, Thamara, Barbosa, Eduardo P., Huertas, Blanca, Marín, Mario A., Lamas, Gerardo, Benmesbah, Mohamed, Brévignon, Christian, Casagrande, Mirna M., Fåhraeus, Christer, Grishin, Nick, Kawahara, Akito Y., Mielke, Olaf H. H., Miller, Jacqueline Y., Nakamura, Ichiro, Navas, Vanessa, Patrusky, Brooke, Pyrcz, Tomasz W., Richards, Lindsay, Tan, Denise, Tyler, Stephanie, Viloria, Angel, Warren, Andrew D., Xiao, Lei, Freitas, André V. L. & Willmott, Keith R., 2023, Combining target enrichment and Sanger sequencing data to clarify the systematics of the diverse Neotropical butterfly subtribe Euptychiina (Nymphalidae, Satyrinae), pp. 1-73 in Zoological Research 2023 on pages 63-64, DOI: 10.1111/syen.12590, http://zenodo.org/record/7909395

Koutalina Viloria & Murienne, 2021 pamela (Hayward, 1957) [Benmesbah et al. (2021, Anartia, 31: 7-62)]

Trico tricolor tricolor (Hewitson, 1850), comb.n. Euptychia tricolor Hewitson (1850, p. 440). TL: river Amazon [Pará, Brazil]. Types: LT male (here designated): ‘ BMNH (E)-1,266,952//Type//B. M. TYPE No. Rh. 3183 Euptychia tricolor ♂ Hew// Para. Hewitson Coll. 79 – 69. Euptychia Tricolor. Hew. 1’. (NHMUK) [figured on www.butterfliesofamerica.com, examined]. BRAZIL: Amapá: Serra do Navio (ICOMI), [0°59'N, 52°3'W], 14 Sep 1963, 1 ♀, (DZUP); Amazonas: 78 km W Manaus, Rio Preto de Eva, 100 m, (Büche, M.), Dec 1996, 1 ♀ [MUSM-LEP-103501], 1 ♀ [MUSM-LEP-103502], (MUSM); Camuna, (Le Moult, E.), 1 ♁ [FLMNH-MGCL-266051], (FLMNH); Manaus, [3°7'S, 60°2'W], (Le Moult, E.), 1 ♁ [FLMNH-MGCL-266043], 1 ♁ [FLMNH-MGCL-266047], 1 ♁ [FLMNH-MGCL-266048], 1 ♁ [FLMNH- MGCL-266049], (FLMNH), (Moss, A. M.), 1 ♀ [BMNH (E)-525460], (NHMUK); Manaus, Reserva Ducke, [2°53'S, 59°57'W], (Hutchings, R. W., Sullivan, J. B.), 13 Dec 1993, 1 ♁ [FLMNH-MGCL-266065], (FLMNH); Maués, [3°22'S, 57°43'W], Nov 1998, 1 ♁ [FLMNH-MGCL-266061], (FLMNH), (Balint), 1 ♁, (DZUP), (Le Moult, E.), 1 ♁ [FLMNH-MGCL-266044], 1 ♁ [FLMNH-MGCL-266045], 1 ♁ [FLMNH-MGCL-266050], 1 ♁ [FLMNH-MGCL-266062], 1 ♁ [FLMNH-MGCL-266063], 1 ♁ [FLMNH-MGCL-266064], (FLMNH); ' São Paulo de Olivença' - (error), (Moss, A. M.), 1 ♀ [BMNH (E)- 525449], 1 ♀ [BMNH (E)- 525451], (NHMUK); Pará: [Rio] Tapajós, [4°16'8''S, 55°59'10''W], 25 m, 1 ♀ [BMNH (E)- 1668489], (NHMUK), (Bates, H. W.), 1 ♁ [BMNH (E)- 1668486], (NHMUK), 3 Oct 2012, 1 ♁, (ZUEC), 4 Oct 2012, 1 ♁, (ZUEC), 8 Dec 2012, 1 ♁, (ZUEC); 15 km S Itaituba, [4°17'S, 56°5'W], (Callaghan, C. J.), 23 Jul 1978, 1 ♁ [FLMNH-MGCL-266054], (FLMNH); Amazonas, (Fassl, A. H.), (ZSM); Belém, Utinga, [1°27'S, 48°25'W], Feb 1957, 1 ♁, (DZUP); Rio Cumina, Igarapé dos Indios, 13 Oct 1936, 1 ♁, (DZUP); Itaituba on Rio Tapajós, [4°16'8''S, 55°59'10''W], (Le Moult, E.), 1 ♁ [FLMNHMGCL-266052], 1 ♁ [FLMNH-MGCL-266060], (FLMNH); Óbidos, [1°54'S, 55°31'W], (Le Moult, E.), 1 ♁ [FLMNH-MGCL-266053], 1 ♁ FLMNHMGCL-266055], 1 ♁ [FLMNH-MGCL-266056], 1 ♀ [FLMNH-MGCL-266059], (FLMNH); Pará, [1°0'S, 51°11'W], 1 ♁ [BMNH (E)- 1266952], 1 ♁ [BMNH (E)- 1668482], 1 ♁ [BMNH (E)- 1668483], 1 ♁ [BMNH (E)- 1668485], (NHMUK), 15 Mar 1893, (ZSM), 1 ♀, (ZSM), (Bates, H. W.), 1 ♁ [BMNH (E)- 1668484], (NHMUK), (Erhardt), (ZSM), (Mathan, M. de), 1 ♁ [BMNH (E)- 1668480], 1 ♁ [BMNH (E)- 1668481], (NHMUK), (Moss, A. M.), 1 ♁ [BMNH (E)- 1499285], 1 ♁ [BMNH (E)- 525595], 1 ♁ [BMNH (E)- 525596], 1 ♁ [BMNH (E)- 525603], 1 ♁ [BMNH (E)- 525605], (NHMUK); Rio Tapajós, [4°16'8''S, 55°59' 10''W], 25 m, (Bates, H. W.), 1 ♁ [BMNH (E)- 1668487], (NHMUK), (Klug, G. G.), Sep 1931, 1 ♁ [BMNH (E)- 1499283], 1 ♁ [BMNH (E)- 1499284], (NHMUK); Santarém, [2°26'S, 54°43'W], (Le Moult, E.), 1 ♀ [FLMNH-MGCL-266057], 1 ♀ [FLMNH-MGCL-266058], (FLMNH); Santa Bárbara do Pará, (Carneiro, E., Dolibaina, D., Dias, F., Moreira), 2-3 Feb 2010, 1 ♁, (DZUP), 6 Feb 2010, 1 ♁, (DZUP); Tapará, [2°2'S, 49°36'W], (Le Moult, E.), 1 ♁ [FLMNH-MGCL-266249], (FLMNH); Taperinha, [0°52' S,47°49'W], (Le Moult, E.), 1 ♁ [FLMNH-MGCL-266046], (FLMNH). FRENCH GUIANA: Cayenne: Approuage River, 10 mi W Regina, vicinity of St Anthanase Lodge, [4°11'10''N, 52°20'8''W], 120 m, (Brock, J.), 14-17 Oct 1990, 2 ♁, (BME); Cayenne: Cacao, [4°35' 51''N, 52°26'11''W], 0-150 m, (Harris, L. N.), 6 May 1998, 1 ♁ [FLMNH-MGCL-266088; Streams-falls-forest], (FLMNH); Cayenne, [4°56'N, 52°20'W], 1 ♁ [BMNH (E)- 1499279], 1 ♁ [BMNH (E)- 1499280], 1 ♁ [BMNH (E)- 1499281], 1 ♁ [BMNH (E)- 1668505], 1 ♁ [BMNH (E)- 1668507], 1 ♀ [BMNH (E)- 1668513], (NHMUK), (Bar, C.), 1 ♁ [BMNH (E)- 1668508], (NHMUK); St-Laurent du Maroni: Aloike-Maripasoula, [3°39'22''N, 53°59' 4''W], 110 m, (Brévignon, C.), 6 Apr 1987, 1 ♁ [MUSM-LEP-103500], (MUSM); Maroni River, (Le Moult, E.), 1 ♁ [FLMNH-MGCL-266082; dissection, KW-14-015], 1 ♁ [FLMNH-MGCL-266083], 1 ♁ [FLMNH-MGCL-266084], 1 ♁ [FLMNH-MGCL-266085], 1 ♁ [FLMNH-MGCL-266086], 1 ♁ [FLMNH-MGCL-266087], 1 ♀ [FLMNH-MGCL-266089], 1 ♀ [FLMNH-MGCL-266090], (FLMNH); Maroni river, Maripasoula, [3°41'N, 54°2'W], (Brévignon, C.), 3 Mar 1987, 1 ♁ [MUSM-LEP-103499], (MUSM); Plateau Mines, Moutouchi Lodge, [5°19'47''N, 54°4'1''W], 45 m, (Sourakov, A.), 1-5 Mar 2013, 1 ♀ [FLMNH-MGCL-209668], (FLMNH); St. Laurent du Maroni, [5°30'N, 54°2'W], 1923, 1 ♁ [BMNH (E)- 1668510], (NHMUK), Jul-Sep 1915, 1 ♁ [BMNH (E)- 1668509], (NHMUK), Jun, 1 ♁ [BMNH (E)-1499282], (NHMUK), (Moult), Nov, 1 ♁ [BMNH (E)- 1668511], (NHMUK); Not located: ‘ French Guiana’, (Bar, C.), 1 ♁ [BMNH (E)- 1668506], (NHMUK). GUYANA: Cuyuni-Mazaruni: Bartica, [6°24'N, 58°37'W], Mar 1939, 1 ♁, (BMB); junction of Cuyuni and Arimu rivers, 30 m, (Fratello, S.), 2 Oct 1991, 1 ♁ [FLMNH-MGCL-266076], (FLMNH); junction of Cuyuni and Arimu rivers, 46 m, (Fratello, S.), 28 Sep 1991, 1 ♀ [FLMNH-MGCL-266072], (FLMNH); East Berbice-Corentyne: Confluence of Oronoque and New River, [2°45'8''N, 57°26'5''W], (Hudson, C. A.), 20 Aug-20 Sep 1937, 1 ♁ [BMNH (E)- 1499271], (NHMUK); ' Kutari [River] Sources', [1°58'N, 56°36'W], (Hudson, C. A.), Jan-Feb 1936, 1 ♁ [BMNH (E)- 1668492], (NHMUK); Upper Demerara-Berbice: Essequibo River, 35 mi SW of Georgetown, Shanklands Resort, [6°28'44''N, 58°34' 54''W], 20 m, (Douglas, M. G.), 20-28 Sep 2006, 1 ♁ [FLMNH-MGCL-266074], 1 ♁ [FLMNH-MGCL-266075], 1 ♀ [FLMNHMGCL-266073], 1 ♀ [FLMNH-MGCL-266078], 1 ♀ [FLMNH-MGCL-266080], 1 ♀ [FLMNH-MGCL-266081], (FLMNH); Not located: ‘ Guyana’, 1 ♁ [BMNH (E)- 1499272], 1 ♁ [BMNH (E)- 1668493], 1 ♁ [BMNH (E)- 1668495], (NHMUK); Demerara, [6°47'N, 58°10'W], (Castell), 1 ♁ [BMNH (E)- 1668491], 1 ♁ [BMNH (E)- 1668494], (NHMUK); Essequibo River, Moraballi Creek, 10 Nov 1929, 1 ♁ [BMNH (E)- 1499270], (NHMUK); Potaro River, 1 ♀ [FLMNH-MGCL-266070], (FLMNH), (Roberts, C. B.), 12 Jul 1902, 1 ♁ [FLMNH-MGCL-266067], (FLMNH), 17 Jul 1902, 1 ♁ [FLMNH-MGCL-266068], 1 ♀ [FLMNH-MGCL-266069], (FLMNH), Jul 1902, 1 ♀ [FLMNH-MGCL-266079], (FLMNH), Oct-Nov 1902, 1 ♁ [FLMNH-MGCL-266066], (FLMNH); Río Demerara, 1 ♁ [BMNH (E)- 1499267], 1 ♁ [BMNH (E)- 1499268], 1 ♁ [BMNH (E)- 1499269], 1 ♁ [BMNH (E)- 1668490], (NHMUK); Suruwabaru CK, Mt. Wokamung, 680-690 m, (Fratello, S.), Nov, 1 ♀ [FLMNH-MGCL-266071], (FLMNH); Suruwabaru CK, Mt. Wokamung, 686 m, (Fratello, S.), Nov, 1 ♀ [FLMNH-MGCL-266077], (FLMNH). SURINAME: Brokopondo: Berg-en-Dal, [5°9'N, 55°4'W], (Ellacombe, C. W.), May 1892, 1 ♁ [BMNH (E)- 1668496], (NHMUK), (Ellacombe, G. W.), May 1892, 1 ♁ [BMNH (E)- 1499273], 1 ♁ [BMNH (E)- 1499275], 1 ♁ [BMNH (E)- 1499277], (NHMUK); Marowijne: Río Lawa, Anapaika, [3°25'N, 54°1'W], (Malkin, B.), 27 Nov 1963, 1 ♁ [MUSM-LEP-103498], (MUSM); Not located: ‘ Surinam’, 1 ♁ [BMNH (E)- 1668497], 1 ♁ [BMNH (E)- 1668498], 1 ♁ [BMNH (E)- 1668499], 1 ♁ [BMNH (E)- 1668500], 1 ♁ [BMNH (E)- 1668502], 1 ♁ [BMNH (E)- 1668503], 1 ♁ [BMNH (E)- 1668504], 1 ♁ [BMNH (E)- 1668514], (NHMUK), (Ellacombe, G. W.), 1 ♁ [BMNH (E)- 1499276], (NHMUK); 'interior Surinam', Sep 1892, 1 ♁ [BMNH (E)- 1499278], (NHMUK), (Ellacombe, C. W.), Aug 1892, 1 ♁ [BMNH (E)- 1668501], (NHMUK), (Ellacombe, G. W.), Sep 1892, 1 ♁ [BMNH (E)- 1499274], (NHMUK). TRINIDAD & TOBAGO: Not located: ‘Trinidad’, [10°26'17''N, 61°15'12''w], 1 ♁ [BMNH (E)- 1668512], (NHMUK). VENEZUELA: Bolívar: El Dorado - Sta Elena, km 83, [6°9'59''N, 61°25' 37''W], 160 m, (Harris, L. & H.), 27 Jun 1984, 1 ♀ [FLMNH-MGCL-266042; km. 82], (FLMNH), (Lichy, R.), 18 Feb 1966, 1 ♁ [FLMNH-MGCL-266041; km. 82], (FLMNH). Country unknown: Not located: ' Amazon', (Bates, H. W.), 1 ♁ [BMNH (E)- 1668488], (NHMUK); 'Amazonas', (Fassl, A. H.), 1 ♁ [BMNH (E)- 1499286], (NHMUK); no data, 1 ♁ [BMNH (E)- 1499287], 1 ♁ [BMNH (E)- 1499288], 1 ♁ [BMNH (E)- 1499289], 1 ♁ [BMNH (E)- 1499290], (NHMUK) Other records: VENEZUELA: Bolívar: Río Grande, 90 km N El Callao, [7°56'54''N, 62°18'6''W], 100 m, (Neild, A. F. E.), 23 Oct 2000, 1 ♁, (ANNE) (Neild, A. (12 Aug 2020, pers. comm. to KRW by email with images))., Published as part of Espeland, Marianne, Nakahara, Shinichi, Zacca, Thamara, Barbosa, Eduardo P., Huertas, Blanca, Marín, Mario A., Lamas, Gerardo, Benmesbah, Mohamed, Brévignon, Christian, Casagrande, Mirna M., Fåhraeus, Christer, Grishin, Nick, Kawahara, Akito Y., Mielke, Olaf H. H., Miller, Jacqueline Y., Nakamura, Ichiro, Navas, Vanessa, Patrusky, Brooke, Pyrcz, Tomasz W., Richards, Lindsay, Tan, Denise, Tyler, Stephanie, Viloria, Angel, Warren, Andrew D., Xiao, Lei, Freitas, André V. L. & Willmott, Keith R., 2023, Combining target enrichment and Sanger sequencing data to clarify the systematics of the diverse Neotropical butterfly subtribe Euptychiina (Nymphalidae, Satyrinae), pp. 1-73 in Zoological Research 2023 on pages 34-71, DOI: 10.1111/syen.12590, http://zenodo.org/record/7909395, {"references":["Hewitson, W. C. (1850) Descriptions of some new species of butterflies. Annals and Magazine of Natural History, Second Series, 6, 434 - 440."]}

Optimandes Marín, Nakahara & Willmott, 2019 eugenia (C. Felder & R. Felder, 1867) [Willmott et al. (2019, Tropical Lepidoptera Research, 29(1): 29-44)] = phineus (Butler, 1867) [Willmott et al. (2019, Tropical Lepidoptera Research, 29(1): 29-44)] - transversa (Weymer, 1911) [Willmott et al. (2019, Tropical Lepidoptera Research, 29(1): 29-44)] mocha Willmott, J. Hall & Lamas, 2019 [Willmott et al. (2019, Tropical Lepidoptera Research, 29(1): 29-44)], Published as part of Espeland, Marianne, Nakahara, Shinichi, Zacca, Thamara, Barbosa, Eduardo P., Huertas, Blanca, Marín, Mario A., Lamas, Gerardo, Benmesbah, Mohamed, Brévignon, Christian, Casagrande, Mirna M., Fåhraeus, Christer, Grishin, Nick, Kawahara, Akito Y., Mielke, Olaf H. H., Miller, Jacqueline Y., Nakamura, Ichiro, Navas, Vanessa, Patrusky, Brooke, Pyrcz, Tomasz W., Richards, Lindsay, Tan, Denise, Tyler, Stephanie, Viloria, Angel, Warren, Andrew D., Xiao, Lei, Freitas, André V. L. & Willmott, Keith R., 2023, Combining target enrichment and Sanger sequencing data to clarify the systematics of the diverse Neotropical butterfly subtribe Euptychiina (Nymphalidae, Satyrinae), pp. 1-73 in Zoological Research 2023 on page 65, DOI: 10.1111/syen.12590, http://zenodo.org/record/7909395

Emeryus Zacca, Casagrande & Mielke, 2020 argulus (Godart, [1824]), repl. name [Zacca et al. (2020, Austral Entomology, 59: 505-523)] = argante (Cramer, 1779), preocc. (not Fabricius, 1775) [Zacca et al. (2020, Austral Entomology, 59: 505-523)] = huebneri (Butler, 1867) [Zacca et al. (2020, Austral Entomology, 59: 505-523)] - magnum Zacca, Casagrande & Mielke, 2020 [Zacca et al. (2020, Austral Entomology, 59: 505-523)] difficilis (Forster, 1964) [Zacca et al. (2020, Austral Entomology, 59: 505-523)] numeria (C. Felder & R. Felder, 1867) [Zacca et al. (2020, Austral Entomology, 59: 505-523)] = ambigua (Butler, 1867) [Zacca et al. (2020, Austral Entomology, 59: 505-523)] = historie (Weymer, 1911), nom. nud. [Zacca et al. (2020, Austral Entomology, 59: 505-523)]

Scriptor Nakhara & Espeland, 2020 sphenophorus Lamas & Nakahara, 2020 [Nakahara et al. (2020, PeerJ, DOI: 10.7717/peerj.10324)], Published as part of Espeland, Marianne, Nakahara, Shinichi, Zacca, Thamara, Barbosa, Eduardo P., Huertas, Blanca, Marín, Mario A., Lamas, Gerardo, Benmesbah, Mohamed, Brévignon, Christian, Casagrande, Mirna M., Fåhraeus, Christer, Grishin, Nick, Kawahara, Akito Y., Mielke, Olaf H. H., Miller, Jacqueline Y., Nakamura, Ichiro, Navas, Vanessa, Patrusky, Brooke, Pyrcz, Tomasz W., Richards, Lindsay, Tan, Denise, Tyler, Stephanie, Viloria, Angel, Warren, Andrew D., Xiao, Lei, Freitas, André V. L. & Willmott, Keith R., 2023, Combining target enrichment and Sanger sequencing data to clarify the systematics of the diverse Neotropical butterfly subtribe Euptychiina (Nymphalidae, Satyrinae), pp. 1-73 in Zoological Research 2023 on page 69, DOI: 10.1111/syen.12590, http://zenodo.org/record/7909395

Trico Nakahara & Espeland, genus novum. Type species — Euptychia tricolor Hewitson, 1850, by present designation. Zoobank registration: https://zoobank.org/Nomenclatural Acts/1EB03CDD-C7B3-4AB0-9920-CC59642A4353 Systematic placement and diagnosis. Trico gen.n. is a member of the so-called ‘ Splendeuptychia clade’, and it is weakly supported as sister to Omacha (SH-aLRT/UFBoot = 52.3/62, Figure 10), with these two taxa being rather distantly related to the remainder of the clade. This genus was not included in Espeland et al. (2019a). Nevertheless, both Trico gen.n. and Omacha are monotypic genera, and both accommodate distinctive euptychiine species that do not resemble each other, or other euptychiine genera. Males of Trico gen.n. are distinguished from virtually all euptychiine species by the combination of their iridescent blue scales covering DHW cells Cu 2 and 2A, which also traverse up along the outer margin towards apex, coupled with orangish area anterior of DHW M 2 (Figures 26 and 27). Although lilac-blue scales are present in a similar area in some male euptychiine species (e.g. Chloreuptychia chlorimene, Lazulina tolumnia comb.n.), these species never possess orangish colouration at the anterior region of DHW. On the other hand, female specimens of Trico gen.n. can closely resemble some euptychiine taxa, perhaps phenotypically being most similar to Satyrotaygetis iris comb.n., despite being distantly related. The male of S. iris is easily distinguished from female specimens of Trico gen.n. by its androconial scales at the distal side of DHW discal cell extending slightly along M 3 and Cu 1, as well as presence of greyish long setiform scales in the discal cell, which are both absent in the female of Trico gen.n. Female specimens of Trico gen.n. are distinguished from females of S. iris comb.n. by having a smoother VHW submarginal band in cell Cu 2, which usually bends inwards in S. iris comb.n. The females of these two taxa can be further distinguished by the membranous lamella antevaginalis of specimens of Trico gen.n. (Figure 28c; sclerotized in S. iris) and ductus seminalis exiting from the ductus bursae closer to the corpus bursae in specimens of Trico gen.n. (Figure 28d; origin of ductus bursae close to ostium bursae in S. iris comb.n.). Etymology. The generic name is based on the Latin word ‘trico’, which is a masculine noun in the nominative singular, meaning ‘mischief-maker’ or ‘trickster’, in reference to the remarkable sexual dimorphism of this species. Description (Figures 26 – 28). Some notable characters include: eyes naked; pterothoracic legs dorsally darker, tibia with two principal longitudinal rows of spines ventrally, pair of short spurs of similar length at distal end of tibia, first tarsomere with three principal longitudinal rows of spines ventrally until distal end, distal end of first tarsomere and remaining distal tarsomeres with four principal longitudinal rows of spines ventrally. Medium-sized Euptychiina (FW length typically 21 – 25 mm). Sexes dimorphic. Male: DHW with light yellowish long hair-like scales at base of discal cell, somewhat at posterior edge of orangish area; greyish long hair-like scales visible in cell Cu 2, area between postdiscal band and submarginal band, along 2A; greyish androconial patch present in adjacent area of cell 2A in male, otherwise wing shape and pattern as illustrated (Figures 26 and 27). Male genitalia with cornuti absent, otherwise as illustrated (Figure 28). Female: DFW and DHW with purple lilac sheen; otherwise as illustrated (Figures 26 and 27). Female abdomen with intersegmental membrane of 7th and 8th abdominal segment not pleated and expandable (Figure 28); genitalia with lamella antevaginalis membranous; ductus bursae membranous, pair of signa present on corpus bursae. Trico Nakahara & Espeland, gen.n. tricolor (Hewitson, 1850), comb.n., was Magneuptychia - fulgora (Butler, 1869), comb.n., was Magneuptychia, Published as part of Espeland, Marianne, Nakahara, Shinichi, Zacca, Thamara, Barbosa, Eduardo P., Huertas, Blanca, Marín, Mario A., Lamas, Gerardo, Benmesbah, Mohamed, Brévignon, Christian, Casagrande, Mirna M., Fåhraeus, Christer, Grishin, Nick, Kawahara, Akito Y., Mielke, Olaf H. H., Miller, Jacqueline Y., Nakamura, Ichiro, Navas, Vanessa, Patrusky, Brooke, Pyrcz, Tomasz W., Richards, Lindsay, Tan, Denise, Tyler, Stephanie, Viloria, Angel, Warren, Andrew D., Xiao, Lei, Freitas, André V. L. & Willmott, Keith R., 2023, Combining target enrichment and Sanger sequencing data to clarify the systematics of the diverse Neotropical butterfly subtribe Euptychiina (Nymphalidae, Satyrinae), pp. 1-73 in Zoological Research 2023 on pages 32-67, DOI: 10.1111/syen.12590, http://zenodo.org/record/7909395, {"references":["Hewitson, W. C. (1850) Descriptions of some new species of butterflies. Annals and Magazine of Natural History, Second Series, 6, 434 - 440.","Espeland, M., Breinholt, J., Barbosa, E. P., Casagrande, M., Huertas, B., Lamas, G. et al. (2019 a) Four hundred shades of brown: higher level phylogeny of the problematic Euptychiina (Lepidoptera, Nymphalidae, Satyrinae) based on hybrid enrichment data. Molecular Phylogenetics and Evolution, 131, 116 - 124.","Butler, A. G. (1869) Lepidoptera Exotica, or descriptions and illustrations of exotic Lepidoptera. London: E. W. Janson, p. 339."]}

Euptychoides Forster, 1964 albofasciata (Hewitson, 1869) inani (Staudinger, [1886]), comb.n., was Macrocissia [Andrade et al. (2019) placed this species in Macrocissia, but it does not cluster with the type species of that genus, instead being strongly supported by molecular data as a member of a clade that includes the type species of Euptychoides] = juani (Staudinger, 1887), missp., comb.n., was Macrocissia laccine (C. Felder & R. Felder, 1867) = saturnus (Butler, 1867), syn.n. [number of pupils in VHW ocelli is variable even between wings of the same individual] pseudosaturnus Forster, 1964, Published as part of Espeland, Marianne, Nakahara, Shinichi, Zacca, Thamara, Barbosa, Eduardo P., Huertas, Blanca, Marín, Mario A., Lamas, Gerardo, Benmesbah, Mohamed, Brévignon, Christian, Casagrande, Mirna M., Fåhraeus, Christer, Grishin, Nick, Kawahara, Akito Y., Mielke, Olaf H. H., Miller, Jacqueline Y., Nakamura, Ichiro, Navas, Vanessa, Patrusky, Brooke, Pyrcz, Tomasz W., Richards, Lindsay, Tan, Denise, Tyler, Stephanie, Viloria, Angel, Warren, Andrew D., Xiao, Lei, Freitas, André V. L. & Willmott, Keith R., 2023, Combining target enrichment and Sanger sequencing data to clarify the systematics of the diverse Neotropical butterfly subtribe Euptychiina (Nymphalidae, Satyrinae), pp. 1-73 in Zoological Research 2023 on page 64, DOI: 10.1111/syen.12590, http://zenodo.org/record/7909395

Cyllopsis clade The ‘ Cyllopsis clade’ was recognized in Espeland et al. (2019a), where it was placed as a strongly supported sister group to a much larger clade including major, species-rich clades such as the ‘ Splendeuptychia clade’, ‘ Taygetis clade’, ‘ Megisto clade’, ‘ Pareuptychia clade’. The clade was similarly recovered in previous molecular phylogenetic studies (Murray & Prowell, 2005; Nakahara, Janzen, et al., 2015; Peña et al., 2010), and in the recent study of Rosa et al. (2021). Currently, four genera are recognized in the ‘ Cyllopsis clade’, namely Cyllopsis (30 species), Paramacera (5 species), Inbio (monotypic) and Atlanteuptychia (monotypic), and our molecular data suggest further taxonomic changes at the generic level are not needed (Figures 3, S 2 and S 3). The relationships between all genera are stable and well-supported in all analyses, with Atlanteuptychia being sister to the remainder of the clade, Inbio being sister to Paramacera, and these two again being sister to Cyllopsis. Thus, the generic classification is perhaps the most stable in comparison with other clades in the subtribe that contain multiple genera. Based on our unpublished data, the only known undescribed species within this clade are in Cyllopsis. Only two Cyllopsis species have been described more recently, including the unique Cyllopsis emilia, which appears to be one of the few euptychiine species that may be involved in mimicry with Pieridae butterflies (Chacon & Nishida,2002; Warren et al., 2018). Conversely, there are likely several currently recognized Cyllopsis species that may represent seasonal forms, so some revision to the species taxonomy is likely needed (Warren, unpublished data). Members of the ‘ Cyllopsis clade’ are relatively small butterflies, with the forewing length of most species not exceeding 25 mm. Possible synapomorphies (not unique) and distinctive characters for the clade include the following: (a) brachia short (about half of uncus in length or shorter),described as ‘not freely articulated’ in Miller (1974); (b) smooth vesica lacking obvious sclerotized cornuti. Unlike other clades in the subtribe, members of the ‘ Cyllopsis clade’ are most diverse in Central America, especially in southern Mexico and Guatemala. In fact, the monotypic genus Atlanteuptychia is the only taxon in this clade found east of the Andes, namely from the Atlantic Forest of Brazil (Freitas et al., 2013), resulting in a disjunct distribution for the clade. Habitat preferences of species in this clade range from lowland rainforest to cloud forest at over 2000 m, with Cyllopsis being one of only three Euptychiina genera to have diversified extensively in montane regions (the others are Forsterinaria and Moneuptychia). The immature stages have been described for Inbio hilara feeding on Rhipidocladum (Poaceae: Bambusoideae: Bambuseae) (Nakahara,Janzen, et al., 2015), and various Cyllopsis species have been reported feeding on bamboo and other grass genera (Beccaloni et al., 2008), but otherwise there are few comprehensive, detailed descriptions or hostplant records for most species in the clade., Published as part of Espeland, Marianne, Nakahara, Shinichi, Zacca, Thamara, Barbosa, Eduardo P., Huertas, Blanca, Marín, Mario A., Lamas, Gerardo, Benmesbah, Mohamed, Brévignon, Christian, Casagrande, Mirna M., Fåhraeus, Christer, Grishin, Nick, Kawahara, Akito Y., Mielke, Olaf H. H., Miller, Jacqueline Y., Nakamura, Ichiro, Navas, Vanessa, Patrusky, Brooke, Pyrcz, Tomasz W., Richards, Lindsay, Tan, Denise, Tyler, Stephanie, Viloria, Angel, Warren, Andrew D., Xiao, Lei, Freitas, André V. L. & Willmott, Keith R., 2023, Combining target enrichment and Sanger sequencing data to clarify the systematics of the diverse Neotropical butterfly subtribe Euptychiina (Nymphalidae, Satyrinae), pp. 1-73 in Zoological Research 2023 on page 8, DOI: 10.1111/syen.12590, http://zenodo.org/record/7909395, {"references":["Espeland, M., Breinholt, J., Barbosa, E. P., Casagrande, M., Huertas, B., Lamas, G. et al. (2019 a) Four hundred shades of brown: higher level phylogeny of the problematic Euptychiina (Lepidoptera, Nymphalidae, Satyrinae) based on hybrid enrichment data. Molecular Phylogenetics and Evolution, 131, 116 - 124.","Murray, D. & Prowell, D. P. (2005) Molecular phylogenetics and evolutionary history of the neotropical Satyrine subtribe Euptychiina (Nymphalidae: Satyrinae). Molecular Phylogenetics and Evolution, 34, 67 - 80.","Pena, C., Nylin, S., Freitas, A. V. L. & Wahlberg, N. (2010) Biogeographic history of the butterfly subtribe Euptychiina (Lepidoptera, Nymphalidae, Satyrinae). Zoologica Scripta, 39, 243 - 258.","Rosa, A. H. B., Huertas, B., Willmott, K. R., Barbosa, E. P., Machado, P. A., Mielke, O. H. H. et al. (2021) Fifty years without a name: a new species of Splendeuptychia Forster (Lepidoptera: Nymphalidae: Satyrinae). Zootaxa, 5061, 95 - 114.","Chacon, I. & Nishida, K. (2002) A new species of Cyllopsis (Lepidoptera: Nymphalidae: Satyrinae) from Costa Rica. Revista de Biologia Tropical, 50, 679 - 683.","Warren, A. D., Nakahara, S., Llorente-Bousquets, J., Luis-Martinez, A. & Miller, J. Y. (2018) A new species of Cyllopsis R. Felder, 1869 from the highlands of Chiapas, Mexico (Lepidoptera: Nymphalidae: Satyrinae). Zootaxa, 4403, 570 - 577.","Miller, L. D. (1974) Revision of the Euptychiini (Satyridae) 2. Cyllopsis R. Felder. Bulletin of the Allyn Museum, 20, 1 - 98.","Freitas, A. V. L., Barbosa, E. P., Santos, J. P. & Mielke, O. H. H. (2013) A new genus, Atlanteuptychia gen. Nov., for Euptychia Ernestina (Lepidoptera: Nymphalidae: Satyrinae). Zoologia (Curitiba), 30, 661 - 668.","Beccaloni, G. W., Viloria, A. L., Hall, S. K. & Robinson, G. S. (2008) Catalogue of the hostplants of the Neotropical butterflies. Monografias Tercer Milenio. Sociedad Entomologica´Aragonesa (SEA) / Red Iberoamericana de Biogeografia y Entomologia Sistematica (RIBES) / Ciencia y Tecnologia para el Desarrollo (CYTED) / Natural History Museum, London, U. K. (NHM) / Instituto Venezolano de Investigaciones Cientificas, Venezuela (IVIC), Zaragoza, p. 536."]}

Caeruleuptychia Forster, 1964 = Weymerana Forster, 1964 aegrota (Butler, 1867) aetherialis (Butler, 1877) [Nakahara et al. (2017, Insect Systematics and Evolution, 49(2): 130-182)] coelestis (Butler, 1867) coelica (Hewitson, 1869) cyanites (Butler, 1871) = stigmatica (Godman, 1905) divina (Weymer, 1911) francisca (Butler, 1870) [Nakahara et al. (2020, Tijdschrift voor Entomologie, 163: 51-61)] glauca (Weymer, 1911) helena (Anken, 1994) helios (Weymer, 1911) = caelestis Brévignon, 2008 [Brévignon (2008, Inventaire des Satyrinae de Guyane française (Lepidoptera: Nymphalidae), pp. 62- 94, 7 pls., 2 figs. In: Lacomme, D. & L. Manil (Eds.), Lépidoptères de Guyane. Tome 3. Rhopalocères 2)] = caelestissima Brévignon, 2010, repl. name [Nakahara et al. (2017, Insect Systematics and Evolution, 49(2): 130-182)] = keltoumae (Brévignon & Benmesbah, 2012) [Nakahara et al. (2017, Insect Systematics and Evolution, 49(2): 130-182)] lobelia (Butler, 1870) mare (Butler, 1869) maryzenderae Lamas & Nakahara, 2017 [Nakahara et al. (2017, Insect Systematics and Evolution, 49(2): 130-182)] penicillata (Godman, 1905) pilata (Butler, 1867) romani (Aurivillius, 1929), comb.n., was Cepheuptychia [preliminary DNA sequence data suggest that this species should provisionally be placed in Caeruleuptychia (Barbosa, unpublished data)] saul Brévignon & Benmesbah, 2012 [Brévignon & Benmesbah (2012, Complément à l'inventaire des Satyrinae de Guyane (Lepidoptera: Nymphalidae), pp. 36-52, 4 pls., 1 tab. In: Lacomme, D. & L. Manil (Eds.), Lépidoptères de Guyane. Tome 7. Nymphalidae)] scopulata (Godman, 1905) = marisea (Dyar, 1914) = susanna (Hayward, 1957) scripta Nakahara, Zacca & Huertas, 2017 [Nakahara et al. (2017, Insect Systematics and Evolution, 49(2): 130-182)] tenera (Weymer, 1911) thaliana Nakahara & Piovesan, 2022 [Nakahara et al. (2023, Tropical Lepidoptera Research, 33(1): 23-36)] trembathi Willmott, Nakahara, J. Hall & Neild, 2017 [Nakahara et al. (2017, Insect Systematics and Evolution, 49(2): 130-182)] twalela Brévignon, 2005 [Brévignon (2005, Lambillionea, 105(3)(1): 393-404)] umbrosa (Butler, 1870) = viridicans (Weymer, 1911) urania (Butler, 1867) = caerulea (Butler, 1869) [Ríos (2019, Anartia, 29: 49-53)] = sinchi Henao, 2019 [Ríos (2019, Anartia, 29: 49-53)] ziza (Butler, 1869), Published as part of Espeland, Marianne, Nakahara, Shinichi, Zacca, Thamara, Barbosa, Eduardo P., Huertas, Blanca, Marín, Mario A., Lamas, Gerardo, Benmesbah, Mohamed, Brévignon, Christian, Casagrande, Mirna M., Fåhraeus, Christer, Grishin, Nick, Kawahara, Akito Y., Mielke, Olaf H. H., Miller, Jacqueline Y., Nakamura, Ichiro, Navas, Vanessa, Patrusky, Brooke, Pyrcz, Tomasz W., Richards, Lindsay, Tan, Denise, Tyler, Stephanie, Viloria, Angel, Warren, Andrew D., Xiao, Lei, Freitas, André V. L. & Willmott, Keith R., 2023, Combining target enrichment and Sanger sequencing data to clarify the systematics of the diverse Neotropical butterfly subtribe Euptychiina (Nymphalidae, Satyrinae), pp. 1-73 in Zoological Research 2023 on pages 65-66, DOI: 10.1111/syen.12590, http://zenodo.org/record/7909395

Graphita Nakahara, Marín & Barbosa, 2016 griphe (C. Felder & R. Felder, 1867) [Nakahara et al. (2016, Neotropical Entomology, 45: 675-691)] = vesta (Butler, 1867) [Nakahara et al. (2016, Neotropical Entomology, 45: 675-691)] = ucumariensis (Andrade, 1991) [Nakahara et al. (2016, Neotropical Entomology, 45: 675-691)], Published as part of Espeland, Marianne, Nakahara, Shinichi, Zacca, Thamara, Barbosa, Eduardo P., Huertas, Blanca, Marín, Mario A., Lamas, Gerardo, Benmesbah, Mohamed, Brévignon, Christian, Casagrande, Mirna M., Fåhraeus, Christer, Grishin, Nick, Kawahara, Akito Y., Mielke, Olaf H. H., Miller, Jacqueline Y., Nakamura, Ichiro, Navas, Vanessa, Patrusky, Brooke, Pyrcz, Tomasz W., Richards, Lindsay, Tan, Denise, Tyler, Stephanie, Viloria, Angel, Warren, Andrew D., Xiao, Lei, Freitas, André V. L. & Willmott, Keith R., 2023, Combining target enrichment and Sanger sequencing data to clarify the systematics of the diverse Neotropical butterfly subtribe Euptychiina (Nymphalidae, Satyrinae), pp. 1-73 in Zoological Research 2023 on page 56, DOI: 10.1111/syen.12590, http://zenodo.org/record/7909395

Deltaya Willmott, Nakahara & Espeland, genus novum. Type species — Papilio ocypete Fabricius, [1777], by pre- sent designation. Zoobank registration: https://zoobank.org/Nomenclatural Acts/C5441C63-9631-4E9D-A37F-6ED27676D185 Systematic placement and diagnosis. Deltaya gen.n. is a member of the ‘ Splendeuptychia clade’ (Figure 10), and three of its four ‘core’ described species, Deltaya ocypete comb.n, Deltaya louisammour comb.n and Deltaya opima comb.n. (Figures 33 and 34), form a clade with low support (FULL dataset SH-aLRT = 67.9, UFB = 53). Deltaya pallema comb.n. is closely related to D. ocypete and is also considered a ‘core’ species (Benmesbah et al., 2018). Two species are placed with some support as forming a grade sister to this clade (SHaLRT = 85.4, UFB = 83) and are included here within the genus (Deltaya andrei comb.n. and Deltaya probata comb.n, see Discussion below). Within the ‘ Splendeuptychia clade’, Deltaya gen.n. is a member of the same clade as Modica gen.n., Paryphthimoides, Colombeia, Scriptor, and Malaveria, with it being placed as sister to Scriptor (SHaLRT = 98.3, UFB = 61), but none of the nodes connecting these genera have strong support. Deltaya gen.n. lacks obvious morphological synapomorphies, but the following are somewhat distinctive characters within Euptychiina (shared variously with other genera) on the ventral wings (Figures 33 and 34): relatively broad dark transverse discal bands; the presence of only five relatively large postdiscal VHW ocelli, with each ocellus having two pupils, which are small dots (in cells Cu 2 -Cu 1, M 2 - M 1) or elongate and large (in cells Cu 1 - M 3, M 3 - M 2, M 1 -Rs), and the postdiscal ocelli on both wings lying within a broad darker band (umbra). Overall, the genus is best distinguished from phenotypically similar genera by the pale pupils of the HW ocellus in cell Cu 2 -Cu 1 not being visible on the dorsal surface, by the VHW postdiscal ocelli in cells Cu 1 -M 3 and M 3 -M 2 being of similar size, round (not elongate) and each containing two enlarged, elongated silver pupils (when the ocelli are large enough for the pupils to be visible), and by the marginal line widening in the VHW tornus. This last character also occurs in Scriptor, some Paryphthimoides, and some other Euptychiina (e.g. Vanima labe, Vanima palladia), but is otherwise a relatively uncommon character. Characters that differ among the genera within the clade in which Deltaya gen.n. is placed are summarized in Table 1. The genitalia of both sexes are described below and there are no obvious synapomorphies for the genus as a whole. However, ‘core’ Deltaya gen.n. (Figure 35) share several distinctive characters, including: (a) the aedeagus has two relatively long and broad rectangular patches of cornuti (smaller or no patches in other species). (b) in the female, the anterior half of the 8th tergite is weakly sclerotized and the intersegmental membrane between the 7th and 8th segments is pleated and expandable; (c) the ventral portion of the intersegmental membrane between the 7th and 8th segments bears a sclerotized plate similar to the 7th sternite; (d) a sclerotized rectangular lamella antevaginalis extending horizontally and fused with the lateral plate of the 8th abdominal segment is present (as noted by Nakahara, Kleckner, et al., 2020). Etymology. The generic name is derived from the name of the fourth letter of the Greek alphabet, delta, whose triangular upper-case symbol is the source of the word used in multiple languages to describe the broadening of a river at its mouth. The name is treated as a feminine noun in the nominative singular, and refers to the broadening VHW marginal line at the tornus that distinguishes Deltaya species from a number of other similar species. Description (Figures 33 – 35). Some notable characters include: eyes setose; pterothoracic legs dorsally slightly darker, tibia with two principal longitudinal rows of spines ventrally, pair of spurs of similar length at distal end of tibia, first tarsomere with three principal longitudinal rows of spines ventrally, remaining tarsomeres with four principal longitudinal rows of spines ventrally. Medium-sized Euptychiina (FW length typically 19 – 25 mm), FW triangular and rather rounded at apex, HW rounded. No strong sexual dimorphism: Dorsal wings dark brown to blue-grey, pale pupils of ocellus in Cu 2 -Cu 1 not visible on DHW, no androconial scales present. Ventral wings blue-grey to brownish grey; relatively broad, dark brown to reddish brown discal and postdiscal lines traversing both wings; VFW with three postdiscal ocelli in cells Cu 1 -M 3, M 3 -M 2, M 2 -M 1, anterior ocellus more clearly marked than remainder, lying within a broad dark brown band (umbra); VHW similar to VFW but with five postdiscal ocelli between veins Cu 2 and Rs, those in Cu 2 -Cu 1 and M 2 -M 1 typically slightly larger, black-centred with two silver dots in each ocellus as pupils, ocelli in M 1 -Rs similar but much smaller, and those in Cu 1 -M 2 with dark brown centres and elongate silver pupils; marginal line thin and even throughout (except rather broad throughout in D. probata comb.n.), thickening noticeably at tornus. Male eighth abdominal tergite reduced dorsally, leaving a sclerotized strip along anterior edge and no or an isolated weakly sclerotized patch in posterior portion. Male genitalia with uncus longer than tegumen, brachia pointing slightly dorsally of uncus and about two-thirds its length; valvae elongate with dorsal edge straight or (‘core’ Deltaya gen.n.) with projection near distal tip covered with tiny spines; aedeagus lacking cornuti but with tiny spines scattered dorsally in D. probata comb.n. and D. andrei comb.n., and with cornuti in ‘core’ Deltaya gen.n. forming two relatively broad and long rectangular patches of tiny spines; intersegmental membrane between eighth segment and genitalic capsule in D. andrei comb.n. forms a ventro-lateral, eversible ‘pocket’ containing dense, long, black scales. Female genitalia has eighth tergite reduced to a posterior sclerotized patch about one- to two-thirds width of segment, eighth segment with large irregular lateral sclerotized plate extending broadening dorsally and with indentation at dorsal edge, antrum unsclerotized, ductus bursae unsclerotized, corpus bursae small, oval and with two narrow sub-parallel signa. ‘Core’ Deltaya gen. n. have additional characters described above under Diagnosis. Distribution and natural history (Figure 36). Deltaya gen.n. contains six described species and probably several undescribed species, which occur in rainforest from sea level to 1400 m and are confined to South America east of the Andes. The genus reaches its peak diversity in the central and western Amazon, where both sexes are typically uncommon in the understory of both disturbed and undisturbed forest, except for D. ocypete comb.n., which may also be locally common in secondary habitats, including in the drier interandean valleys and extra-Amazonian regions from Bolivia across to south-eastern Brazil. Deltaya gen.n. species have also been recorded in fruit-baited traps (Benmesbah et al., 2018; Oliveira et al., 2021; Zacca, Casagrande, et al., 2017). The immature stages have not been described for any species to date, and while there are records of Cyperaceae and Poaceae as hostplants of D. ocypete comb.n. (Singer & Ehrich, 1993), these require confirmation given prior confusion over the identification of this species. Discussion. Benmesbah et al. (2018) discussed in detail the taxonomic history of the type species of this genus, Papilio ocypete, and designated a specimen with a DNA barcode as the neotype. This species and three other described species, D. opima comb.n., D. louisammour comb.n. and D. pallema comb.n., form a clade based on DNA sequences (‘core’ Deltaya gen. n. ), and share several morphological synapomorphies. All these species were previously placed in Magneuptychia by Lamas (2004), but, for the same reasons as discussed under Modica gen.n., they cannot reasonably be accommodated in any described genus, except perhaps Scriptor. Nevertheless, monophyly with Scriptor is only weakly supported in our results, and we thus believe that description of a new genus is the best solution. As mentioned above under Systematic Placement and Diagnosis, we also provisionally include two species in Deltaya that are weakly supported based on molecular data as members of the same clade as ‘core’ Deltaya gen.n., namely D. probata comb.n. (Figure 34d) and D. andrei comb.n. (Figure 34c). Although both species share some wing pattern and genitalic similarities with ‘core’ Deltaya gen. n. , none of these are convincing synapomorphies for Deltaya gen.n. Deltaya andrei comb.n. has a very distinctive autapomorphy, namely a pocket of dense black scales adjacent to the valvae in the male genitalia, while the wing pattern of D. probata comb.n. is rather different from the remaining Deltaya species, with a VHW marginal line that is thickened throughout. Only COI sequences are available for D. probata comb.n., and COI + RPS5 for D. andrei comb.n., and more complete molecular data for both species and other members of the ‘ Splendeuptychia clade’ are therefore needed to confirm the generic classification of D. andrei and D. probata. Deltaya Willmott, Nakahara & Espeland, gen.n. andrei (Zacca, Casagrande & Mielke, 2017), comb.n., was Magneuptychia [Zacca et al. (2017, Zootaxa, 4231(3): 442-450)] louisammour (Benmesbah & Zacca, 2018), comb.n., was Magneuptychia [Benmesbah et al. (2018, Zootaxa, 4425(1): 115-145)] ocypete (Fabricius, [1777]), comb.n., was Magneuptychia = sabina (C. Felder & R. Felder, 1867), comb.n., was Magneuptychia = olivacea (Aurivillius, 1929), comb.n., was Magneuptychia opima (Weymer, 1911), comb.n., was Magneuptychia pallema (Schaus, 1902), comb.n., was Magneuptychia probata (Weymer, 1911), comb.n., was Magneuptychia = aliciae (Hayward, 1957), comb.n., was Magneuptychia, Published as part of Espeland, Marianne, Nakahara, Shinichi, Zacca, Thamara, Barbosa, Eduardo P., Huertas, Blanca, Marín, Mario A., Lamas, Gerardo, Benmesbah, Mohamed, Brévignon, Christian, Casagrande, Mirna M., Fåhraeus, Christer, Grishin, Nick, Kawahara, Akito Y., Mielke, Olaf H. H., Miller, Jacqueline Y., Nakamura, Ichiro, Navas, Vanessa, Patrusky, Brooke, Pyrcz, Tomasz W., Richards, Lindsay, Tan, Denise, Tyler, Stephanie, Viloria, Angel, Warren, Andrew D., Xiao, Lei, Freitas, André V. L. & Willmott, Keith R., 2023, Combining target enrichment and Sanger sequencing data to clarify the systematics of the diverse Neotropical butterfly subtribe Euptychiina (Nymphalidae, Satyrinae), pp. 1-73 in Zoological Research 2023 on pages 38-67, DOI: 10.1111/syen.12590, http://zenodo.org/record/7909395, {"references":["Benmesbah, M., Zacca, T., Casagrande, M. M., Mielke, O. H. H., Lamas, G. & Willmott, K. R. (2018) Taxonomic notes on Papilio ocypete Fabricius, 1776 and Papilio helle Cramer, 1779 with description of two new similar species from South America (Lepidoptera: Nymphalidae: Satyrinae). Zootaxa, 4425, 115 - 145.","Oliveira, I. F., Baccaro, F. B., Werneck, F. P., Zacca, T. & Haugaasen, T. (2021) Marked differences in butterfly assemblage composition between forest types in Central Amazonia, Brazil. Forests, 12, 942.","Singer, M. & Ehrich, P. (1993) Host specialization of satyrine butterflies, and their responses to habitat fragmentation in Trinidad. Journal of Research on the Lepidoptera, 30, 248 - 256.","Lamas, G. (2004) Atlas of Neotropical Lepidoptera. Checklist: Part 4 A Hesperioidea - Papilionoidea. Gainesville: Scientific Publishers / Association of Tropical Lepidoptera, p. 439.","Zacca, T., Casagrande, M. M., Mielke, O. H. H., Huertas, B., Neild, A. F. E. & Benmesbah, M. (2017) Description of a new species of Euptychiina (Lepidoptera: Nymphalidae: Satyrinae) from South America. Zootaxa, 4231, 442 - 450.","Aurivillius, P. O. C. (1929) Wissenschaftliche Ergebnisse der schwedischen Entomologischen Reisen des Herrn Dr. A. Roman in Amazonas 1914 - 1915 und 1923 - 1924. 13. Rhopalocera. Entomologisk Tidskrift, 50, 153 - 168.","Weymer, G. (1911) 4. Familie: Satyridae. In: Seitz, A. (Ed.) Die GrossSchmetterlinge der Erde. Stuttgart: A. Kernen, pp. 173 - 280."]}

Cisandina Nakahara & Espeland, 2022 castanya Lamas & Nakahara, 2022 [Nakahara et al. (2022, Insect Systematics and Diversity, 6(1): 2; 1-30)] esmeralda Nakahara & Barbosa, 2022 [Nakahara et al. (2022, Insect Systematics and Diversity, 6(1): 2; 1-30)] fida (Weymer, 1911) [Nakahara et al. (2022, Insect Systematics and Diversity, 6(1): 2; 1-30)] - directa Nakahara & Willmott, 2022 [Nakahara et al. (2022, Insect Systematics and Diversity, 6(1): 2; 1-30)] lea (Cramer, 1777) [Nakahara et al. (2022, Insect Systematics and Diversity, 6(1): 2; 1-30)] = junia (Cramer, 1780) [Nakahara et al. (2022, Insect Systematics and Diversity, 6(1): 2; 1-30)] philippa (Butler, 1867) [Nakahara et al. (2022, Insect Systematics and Diversity, 6(1): 2; 1-30)] = tersa (Weymer, 1911) [Nakahara et al. (2022, Insect Systematics and Diversity, 6(1): 2; 1-30)] sanmarcos (Nakahara & Lamas, 2018) [Nakahara et al. (2022, Insect Systematics and Diversity, 6(1): 2; 1-30)] trinitensis (Brévignon & Benmesbah, 2012) [Nakahara et al. (2022, Insect Systematics and Diversity, 6(1): 2; 1-30)]

Godartiana Forster, 1964 = Praefaunula Forster, 1964 = Cercyeuptychia L.D. Miller & Emmel, 1971 amadoi Paluch, Zacca & Freitas, 2017 [Zacca et al. (2017, Austral Entomology 56(3): 349)] = amadoi Paluch, Zacca & Freitas, 2016, unavailable armilla (Butler, 1867) [Zacca et al. (2017, Austral Entomology, 56: 169-190)] = vesper (Butler, 1867) [Zacca et al. (2017, Austral Entomology, 56: 169-190)] = vesper (Butler, 1867), nom. nud. [Zacca et al. (2017, Austral Entomology, 56: 169-190)] = liturata (Butler, 1867) [Zacca et al. (2017, Austral Entomology, 56: 169-190)] = liturata (Butler, 1867), nom. nud. [Zacca et al. (2017, Austral Entomology, 56: 169-190)] = armilla (Butler, 1867), nom. nud. [Zacca et al. (2017, Austral Entomology, 56: 169-190)] = strigillata (Weymer, 1911) [Zacca et al. (2017, Austral Entomology, 56: 169-190)] astronesthes Lamas & Nakahara, 2018 [Nakahara et al. (2018, Insecta Mundi, 0639: 1-38)] byses (Godart, [1824]) = bimaculata (D’Almeida, 1922) luederwaldti (Spitz, 1931) [Zacca et al. (2017, Austral Entomology, 56: 169-190)] muscosa (Butler, 1870), Published as part of Espeland, Marianne, Nakahara, Shinichi, Zacca, Thamara, Barbosa, Eduardo P., Huertas, Blanca, Marín, Mario A., Lamas, Gerardo, Benmesbah, Mohamed, Brévignon, Christian, Casagrande, Mirna M., Fåhraeus, Christer, Grishin, Nick, Kawahara, Akito Y., Mielke, Olaf H. H., Miller, Jacqueline Y., Nakamura, Ichiro, Navas, Vanessa, Patrusky, Brooke, Pyrcz, Tomasz W., Richards, Lindsay, Tan, Denise, Tyler, Stephanie, Viloria, Angel, Warren, Andrew D., Xiao, Lei, Freitas, André V. L. & Willmott, Keith R., 2023, Combining target enrichment and Sanger sequencing data to clarify the systematics of the diverse Neotropical butterfly subtribe Euptychiina (Nymphalidae, Satyrinae), pp. 1-73 in Zoological Research 2023 on pages 59-60, DOI: 10.1111/syen.12590, http://zenodo.org/record/7909395

Occulta ocnus (Butler, 1867) combinatio nova. Euptychia ocnus Butler (1867, p. 467). TL: Tapajos [Pará, Brazil]; Villa Nova (=Parintins) [Amazonas, Brazil]. Types: ST female: ‘ BMNH (E)-1,266,945//Tapajos, Amazons. H. W. Bates// Type of Species// Type H. T.// ♀ // ♀ Tapajos ocnus Butler Type //B. M. Type No. Rh 3179 Euptychia ocnus ♀ Butl.//Godman-Salvin Coll. 1904:1. Euptychia ocnus, Butl. ’, (NHMUK) [Figure 30d,e, examined]. ST female: ‘Villa Nova, Amazons. Coll. by Bates. 56 – 44.//Villa Nova// BMNH (E) 1497830’, (NHMUK) [examined]. = Euptychia libye f. gracilis (Weymer, 1911; 212, figure 48f) TL: Pebas, [Loreto], Peru. Types: ST male, ‘ LECTOTYPE ♂ Euptychia libye f. gracilis Weymer, 1911 By G. Lamas (‘94//Pebas//Libye - Forma// libyoidea Btlr’), (SMTD) (figured in Warren et al., 2022, examined). Taxonomy. Butler (1867, p. 467) described this species from Brazil, based on an unspecified number of specimens from Tapaj́os, in Bates ’ s collection, and ‘Villa Nova’, in the NHMUK. The original description closely matches two female syntypes in the NHMUK from Tapajos ́and Villa Nova, in particular in describing the five main ocelli in the VHW as comprising three that are encircled with yellow and each having two pupils, and the remaining two as being yellow with a central silver line. Weymer (1911, p. 212, pl. 48f) described Euptychia libye f. gracilis based on an unspecified number of specimens from Pebas, Peru, in the collection of Bang-Haas, and the figure of the ventral surface corresponds with the specimen labelled as lectotype by Lamas in the SMTD. In comparison with the two syntypes of ocnus, this specimen is somewhat paler, and it has a distinct rather than vestigial ocellus in 2A-Cu2 on the VHW, but the remaining VHW ocelli and other wing pattern characters are otherwise typical of this species as here defined. Distribution and natural history (Figure 32). Occulta ocnus gen.n. and comb.n. is known to date from the central Amazon basin, with all recorded localities being in the vicinity of large rivers below 250 m. A list of specimens examined is provided in Table S3. The only two records from Ecuador are from the region of black-water rivers and lakes north of the Río Napo; at Zancudococha, a male was collected flying near to the ground at the edge of forest and small field, on a low ridge, at 13:00 hrs. Immature stages have not been described. A list of specimens examined is provided in Appendix B. Discussion. Occulta gen.n. is described as a monotypic genus for Euptychia ocnus based on molecular data presented here. As previously indicated by Espeland et al. (2019a), Euptychia ocnus is distantly related to the type species of Magneuptychia, Papilio libye Linnaeus, 1767. Although Papilio libye is a member of the clade to which Occulta gen.n. is weakly supported as belonging, expanding the concept of Magneuptychia to include Euptychia ocnus would be unreasonable, for a number of reasons. First, the support for its relationship with the clade containing Papilio libye is low. Second, and perhaps more importantly, such a broad generic classification would be inconsistent with current approaches to building a stable and informative generic classification for the Euptychiina, where monophyly, phenotype and ecological information are incorporated in deciding on the most appropriate taxonomy (e.g. Nakahara, Lamas, et al., 2019). This clade not only includes both widespread lowland taxa (e.g. Magneuptychia libye) and rather narrowly distributed montane species (e.g. undescribed Splendeuptychia species), but also some of the most colourful euptychiine species in the genus Caeruleuptychia. Given that generic names should ideally be informative for the recognition of clades by including species with similar phenotypes, in addition to having potential value in predicting unknown biological traits, we refute the idea of a broad generic classification to include all these taxa contained within the clade containing E. ocnus, and instead establish a monotypic genus for this species. BRAZIL: Amazonas: Carauari, RESEX Médio Juruá, [5°8'57"S,67°18'45"W], (Oliveira, F.), 1 Dec 2018, 1 ♁, 2 ♀ [MN-LEP 0004444, 0004446, 0004453], 2 Dec 2018, 1 ♁, 4 ♀ [MN-LEP 0004448 to 0004452]; Ega, [3°22'S,64°42'W], (Bates, H. W.), 1 ♁ [BMNH (E)-1421814], (NHMUK); Igarapé Massauari, [2°54'17''S,57°8'23''W], 1 ♁ [BMNH (E)-1421845], (NHMUK); São Paulo de Olivença, [3°28'S,68°57'W], (Mathan, M. de), 1 ♁ [BMNH (E)-1497827], (NHMUK), Mar 1883, 1 ♀ [BMNH (E)-1497829], (NHMUK); Villa Nova, (Bates, H. W.), 1 ♀ [BMNH (E)-1497830], (NHMUK); Pará: [Rio] Tapájos, [4°16'8''S,55°59'10''W], 25 m, 25 Oct 2012, 1 ♁, (ZUEC). COLOMBIA: Amazonas: Leticia, [4°12'S,69°56'W], 82 m, (Marín, M. A.), 11 Oct 2012, 1 ♁, (ZUEC), 18 Oct 2012, 1 ♁, (ZUEC), 28 Sep 2012, 1 ♁, (ZUEC). ECUADOR: Orellana: Río Aguarico, Zancudococha, [0°34'23''S,75°26'13''W], 240 m, (Willmott, K. R., J. C. R., J. I. R., Aldaz, R.), 3,5- 8 Jul 2017, 1 ♁ [FLMNH-MGCL-288721], (FLMNH); Sucumbíos: Cuyabeno Lodge, across lagoon, [0°0'18''S,76°10'23''W], 224 m, (Turner, J. D.), 6 Dec 2010, 1 ♁ [FLMNH-MGCL-150971], (FLMNH), 7 Dec 2010, 1 ♁ [FLMNH-MGCL-150970], (FLMNH); Cuyabeno Lodge, salt trail, [0°0'4''S,76°10'50''W], 238 m, (Turner, J. D.), 8 Dec 2010, 1 ♁ [FLMNH-MGCL-150972], (FLMNH). PERU: Loreto: ‘Cavallo Cocha’ [= Caballococha], [3°55'S,70°31'W], 90 m, (Mathan, M. de), May-Jul 1884, 1 ♁ [BMNH (E)-1421876], 1 ♁ [BMNH (E)-1497832], 1 ♁ [BMNH (E)-1497833], 1 ♁ [BMNH (E)-1497835], 1 ♀ [BMNH (E)-1497834], (NHMUK); 55 km NE Iquitos, Explorama [Lodge], [3°27'S,72°51'W], 120 m, (Harvey, D. J.), 24 Sep 1995, 1 ♁ [MUSM-LEP-103309], (MUSM), (Lamas, G.), 24 Sep 1995, 1 ♁ [MUSM-LEP-103306], 1 ♁ [MUSM-LEP-103308], 1 ♀ [MUSM-LEP-103311], (MUSM), (Robbins, R. K.), 24 Sep 1995, 1 ♁ [MUSM-LEP-103310], (MUSM); 55 km NE Iquitos, Explorama [Lodge], [3°27'S,72°51'W], 140 m, (Harvey, D. J.), 23 Sep 1995, 1 ♀ [MUSM-LEP-103312], (MUSM); Boca Río Samíria, 130 m, (Lamas, G.), 14 May 1990, 1 ♁ [MUSM-LEP-103307], (MUSM); ' Explorama 50 mi NE of Iquitos' [=Explorama Lodge 55 km NE Iquitos], [3°27'S,72°51'W], (Brown, H.& B.), 30 Nov 1982, 1 ♁ [FLMNH-MGCL-296549], (FLMNH); Iquitos, [3°45'S,73°15' W], 100 m, (Whitely, H.), 1 ♁ [BMNH (E)-1497836], 1 ♁ [BMNH (E)-1497837], 1 ♀ [BMNH (E)-1497828], (NHMUK); nr. Iquitos, Explornapo Camp, [3°15'S,72°55'W], 120 m, (Brown, H.& B.), 9 Feb 1982, 1 ♁ [FLMNH-MGCL-296551], (FLMNH); Sarayacu, [6°43'S,75°8'W], 160 m, (Garlepp, G.), 1885, 1 ♀, (MNHU). SURINAME: Not located: ‘Surinam’, 1 ♁ [BMNH (E)-1421815], (NHMUK), 1 ♀ [BMNH (E)-1497831], (NHMUK). Country unknown: Not located: no data, 20-22 Jun 2000, 1 ♁ ['RTPRS'], (ZUEC), Published as part of Espeland, Marianne, Nakahara, Shinichi, Zacca, Thamara, Barbosa, Eduardo P., Huertas, Blanca, Marín, Mario A., Lamas, Gerardo, Benmesbah, Mohamed, Brévignon, Christian, Casagrande, Mirna M., Fåhraeus, Christer, Grishin, Nick, Kawahara, Akito Y., Mielke, Olaf H. H., Miller, Jacqueline Y., Nakamura, Ichiro, Navas, Vanessa, Patrusky, Brooke, Pyrcz, Tomasz W., Richards, Lindsay, Tan, Denise, Tyler, Stephanie, Viloria, Angel, Warren, Andrew D., Xiao, Lei, Freitas, André V. L. & Willmott, Keith R., 2023, Combining target enrichment and Sanger sequencing data to clarify the systematics of the diverse Neotropical butterfly subtribe Euptychiina (Nymphalidae, Satyrinae), pp. 1-73 in Zoological Research 2023 on pages 37-73, DOI: 10.1111/syen.12590, http://zenodo.org/record/7909395, {"references":["Butler, A. G. (1867) Descriptions of some new species of Satyridae belonging to the genus Euptychia. Proceedings of the Zoological Society of London, 1867, 104 - 110.","Weymer, G. (1911) 4. Familie: Satyridae. In: Seitz, A. (Ed.) Die Gross- Schmetterlinge der Erde. Stuttgart: A. Kernen, pp. 173 - 280.","Warren, A. D., Davis, K., Stangeland, M., Pelham, J. & Grishin, N. V. (2022) Butterflies of America [WWW document]. Available from: http: // butterfliesofamerica. com [Accessed 25 th June 2022]","Espeland, M., Breinholt, J., Barbosa, E. P., Casagrande, M., Huertas, B., Lamas, G. et al. (2019 a) Four hundred shades of brown: higher level phylogeny of the problematic Euptychiina (Lepidoptera, Nymphalidae, Satyrinae) based on hybrid enrichment data. Molecular Phylogenetics and Evolution, 131, 116 - 124."]}

Satyrotaygetis Forster, 1964 = Macrocissia Viloria, Le Crom & Andrade, 2019, syn.n. iris (C. Felder & R. Felder, 1867), comb.n., was Macrocissia [Andrade et al. (2019) designated Neonympha iris as the type species of a new genus, Macrocissia, which they also noted was "close to Satyrotaygetis ". Molecular data strongly support the two species originally included in Macrocissia by Andrade et al. (2019) as not being sister species, with N. iris sister to S. satyrina (type species of Satyrotaygetis) + S. tiessa. We therefore transfer N. iris to Satyrotaygetis and synonymize Macrocissia] = ayaya (Butler, 1867), comb.n., was Macrocissia = azulina (Ribeiro, 1931), comb.n., was Macrocissia satyrina (Bates, 1865) = gigas (Butler, 1867), stat.rev. [restored from synonymy with satyrina by Andrade et al. (2019) without any justification, we return this name as a synonym pending further study] = incerta (Butler & Druce, 1872), stat.rev. [restored from synonymy with satyrina by Andrade et al. (2019) without any justification, we return this name as a synonym pending further study] tiessa (Hewitson, 1869) [Andrade et al. (2019, Anartia, 28: 26-46)], Published as part of Espeland, Marianne, Nakahara, Shinichi, Zacca, Thamara, Barbosa, Eduardo P., Huertas, Blanca, Marín, Mario A., Lamas, Gerardo, Benmesbah, Mohamed, Brévignon, Christian, Casagrande, Mirna M., Fåhraeus, Christer, Grishin, Nick, Kawahara, Akito Y., Mielke, Olaf H. H., Miller, Jacqueline Y., Nakamura, Ichiro, Navas, Vanessa, Patrusky, Brooke, Pyrcz, Tomasz W., Richards, Lindsay, Tan, Denise, Tyler, Stephanie, Viloria, Angel, Warren, Andrew D., Xiao, Lei, Freitas, André V. L. & Willmott, Keith R., 2023, Combining target enrichment and Sanger sequencing data to clarify the systematics of the diverse Neotropical butterfly subtribe Euptychiina (Nymphalidae, Satyrinae), pp. 1-73 in Zoological Research 2023 on page 65, DOI: 10.1111/syen.12590, http://zenodo.org/record/7909395