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Specialised transformers-based models (such as BioBERT and BioMegatron) are adapted for the biomedical domain based on publicly available biomedical corpora. As such, they have the potential to encode large-scale biological knowledge. We investigate the encoding and representation of biological knowledge in these models, and its potential utility to support inference in cancer precision medicine - namely, the interpretation of the clinical significance of genomic alterations. We compare the performance of different transformer baselines; we use probing to determine the consistency of encodings for distinct entities; and we use clustering methods to compare and contrast the internal properties of the embeddings for genes, variants, drugs and diseases. We show that these models do indeed encode biological knowledge, although some of this is lost in fine-tuning for specific tasks. Finally, we analyse how the models behave with regard to biases and imbalances in the dataset., Comment: 25 pages, 12 figures, supplementary methods, tables and figures at the end of the manuscript

A brief historical review regarding the type locality of Euptychia stigmatica Godman, 1905 was conducted, which suggests that its type locality is actually Rio de Janeiro, Brazil, rather than northeastern Argentina, as previously purported. Consequently, E. stigmatica and its senior synonym E. cyanites Butler, 1871, are regarded to be two species-group names representing a taxon in the euptychiine genus Caeruleuptychia Forster, 1964 known from the Brazilian Atlantic Forest. A lectotype is designated for E. cyanites. Additionally, two closely related species are named and described using an integrative approach with morphological and molecular evidence. Caeruleuptychia harrisi Nakahara & Freitas, sp. nov. and C. aemulatio Nakahara & Willmott, sp. nov. both occur in Amazonia and COI barcode data recovered these taxa as part of the caerulea clade of Caeruleuptychia.

Regenerating natural language explanations in the scientific domain has been proposed as a benchmark to evaluate complex multi-hop and explainable inference. In this context, large language models can achieve state-of-the-art performance when employed as cross-encoder architectures and fine-tuned on human-annotated explanations. However, while much attention has been devoted to the quality of the explanations, the problem of performing inference efficiently is largely under-studied. Cross-encoders, in fact, are intrinsically not scalable, possessing limited applicability to real-world scenarios that require inference on massive facts banks. To enable complex multi-hop reasoning at scale, this paper focuses on bi-encoder architectures, investigating the problem of scientific explanation regeneration at the intersection of dense and sparse models. Specifically, we present SCAR (for Scalable Autoregressive Inference), a hybrid framework that iteratively combines a Transformer-based bi-encoder with a sparse model of explanatory power, designed to leverage explicit inference patterns in the explanations. Our experiments demonstrate that the hybrid framework significantly outperforms previous sparse models, achieving performance comparable with that of state-of-the-art cross-encoders while being approx 50 times faster and scalable to corpora of millions of facts. Further analyses on semantic drift and multi-hop question answering reveal that the proposed hybridisation boosts the quality of the most challenging explanations, contributing to improved performance on downstream inference tasks., Comment: To appear at the 36th AAAI Conference on Artificial Intelligence (AAAI-22)

Article published in the journal Tropical Lepidoptera Research.

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

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

Pareuptychia clade The ‘ Pareuptychia clade’ was proposed first by Murray and Prowell (2005) and corroborated by increasingly comprehensive subsequent studies (Espeland et al., 2019a; Marín et al., 2017; Peña et al., 2006, 2010). The clade contains 11 described genera, Neonympha (2 species), Pareuptychia (9 species), Erichthodes (monotypic), Megeuptychia (3 species), Satyrotaygetis (2 species), Euptychoides (4 species), Taydebis (3 species), Nhambikuara (11 species), Optimandes (2 species), Amiga (2 species), Xenovena gen. n. (monotypic) and an undescribed genus. The ‘ Pareuptychia clade’ is embedded within a larger clade also containing the ‘ Taygetis clade’, ‘ Splendeuptychia clade’ and ‘ Archeuptychia clade’, and we also include the unplaced genus Chloreuptychia in the discussions here (Figures 1, S 1 – S 3). In both FULL trees, Chloreuptychia is sister to the ‘ Pareuptychia clade’, while in both 4GENES trees the genus is sister to the ‘ Archeuptychia clade’. In the backbone tree (Figure S 1) the ‘ Pareuptychia clade’ is sister to the Taygetis clade, and here Chloreuptychia is found to be sister to a clade including the ‘ Taygetis clade’, ‘ Splendeuptychia clade’ and ‘ Archeuptychia clade’. These relationships are not well supported and need to be further investigated. The small genus Amiga is strongly supported as sister group to the remainder of the clade in all datasets. Most generic relationships are well-supported (Figures 9, S 2 and S 3). The clade is divided into two well-supported subclades, the Optimandes subclade (Optimandes, Nhambikuara) and the Pareuptychia subclade (Xenovena gen.n., Pareuptychia, Erichthodes, Neonympha, Euptychoides, Satyrotaygetis, and an undescribed generic-level clade containing some species currently in Erichthodes), consistent with previous molecular studies (Murray & Prowell, 2005; Peña et al., 2010). The placement of Taydebis needs to be further investigated; although it is found as sister to Optimandes + Nhambikuara in all trees, the support for this branch is always low. Since the checklist Lamas (2004), a number of papers have helped revise the generic classification of the ‘ Pareuptychia clade’, with the description of new genera such as Taydebis (Freitas, 2003), Nhambikuara (Freitas, Barbosa, et al., 2018), Amiga (Nakahara, Lamas, et al., 2019) and Optimandes (Willmott et al., 2019), the sorting of species in previously described genera (Andrade-Correa et al., 2019) and descriptions of 11 new species (Benmesbah, 2015; Brévignon, 2005; Freitas et al., 2011; Freitas, Barbosa, et al., 2018; Huertas, 2011; Huertas et al., 2009; Nakahara, Marin, & Neild, 2016; Willmott et al., 2018, 2019). We found strong support for a clade containing the type species of Nhambikuara, N. cerradensis, along with nine other species formerly placed in Splendeuptychia (Lamas, 2004), a relationship partly indicated in Freitas, Barbosa, et al. (2018). Morphological characters also support this as a relatively compact group of species (Huertas, 2014) and we thus transfer the nine former Splendeuptychia species into Nhambikuara (comb.n.). We found strong support for the recently described genus Macrocissia (Andrade-Correa et al., 2019) not being monophyletic, and the two species previously included in this genus are transferred to Satyrotaygetis (Satyrotaygetis iris [C. Felder & R. Felder, 1867]) and Euptychoides (Euptychoides inani [Staudinger, 1886]) in concordance with the phylogeny. The combination of morphological characters used by Andrade-Correa et al. (2019) to support Macrocissia, ‘vestigial recurrent vein into the discal cell’ in the forewing, and ‘valvae with a hook-like distal process’ and a ‘finely serrate cucullus in the male genitalia’, is a combination of characters present in previously described genera such as Satyrotaygetis and Euptychoides, and we thus synonymize Macrocissia syn.n. with Satyrotaygetis. The ‘ Pareuptychia clade’ contains 44 described species, with the genera Pareuptychia and Nhambikuara, with nine and 11 species respectively, being the most diverse. Taxonomic revisions for these two genera are in progress, in addition to work on Taydebis, Euptychoides and Erichthodes. Members of the ‘ Pareuptychia clade’ have a wide distribution, ranging from southern Canada to northern Argentina, and from sea level to almost 2600 m, with the highest diversity in the Amazon basin and the Andes (Marín et al. in preparation). According to Marín et al. (2017), possible synapomorphies (not unique) and/or distinctive characters for the clade include the following: (a) obvious sclerotized cornuti in aedeagus; (b) in the valva, posterior ending of the cucullus acute. Hostplants for members of this clade are mostly bamboos or grasses in the family Poaceae, with records of Cyperaceae in Pareuptychia (Janzen & Hallwachs, 2022) and Neonympha (Hamm et al., 2013), and Marantaceae in Megeuptychia (Janzen & Hallwachs, 2022). Immature stages have been described or illustrated for Amiga, Taydebis, Pareuptychia, Optimandes, Satyrotaygetis, Megeuptychia and Nhambikuara (Corahua-Espinoza, Nakahara, Kabir, et al., 2022; Corahua-Espinoza, Nakahara, Shellman, et al., 2022; Freitas, 2003; Freitas, Barbosa, & Marín, 2016; Janzen & Hallwachs, 2022; Murray, 2001b; Willmott et al., 2019). These are morphologically simple and similar to those of several other species of Neotropical Euptychiina larvae, lacking body scoli, and having short- to medium-sized head horns and caudal filaments, and the pupae are short and smooth., 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 18, DOI: 10.1111/syen.12590, http://zenodo.org/record/7909395, {"references":["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.","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.","Marin, M. A., Pena, C., Uribe, S. I. & Freitas, A. V. L. (2017) Morphology agrees with molecular data: phylogenetic affinities of Euptychiina butterflies (Nymphalidae: Satyrinae). Systematic Entomology, 42, 768 - 785.","Pena, C., Wahlberg, N., Weingartner, E., Kodandaramaiah, U., Nylin, S., Freitas, A. V. L. et al. (2006) Higher level phylogeny of Satyrinae butterflies (Lepidoptera: Nymphalidae) based on DNA sequence data. Molecular Phylogenetics and Evolution, 40, 29 - 49.","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.","Lamas, G. (2004) Atlas of Neotropical Lepidoptera. Checklist: Part 4 A Hesperioidea - Papilionoidea. Gainesville: Scientific Publishers / Association of Tropical Lepidoptera, p. 439.","Freitas, A. V. L. (2003) Description of a new genus for \" Euptychia peculiaris \" (Nymphalidae: Satyrinae): immature stages and systematic position. Journal of the Lepidopterists' Society, 57, 100 - 106.","Willmott, K. R., Marin, M. A., Nakahara, S., Pomerantz, T., Lamas, G., Huertas, B. et al. (2019) A revision of the new Andean butterfly genus Optimandes Marin, Nakahara & Willmott, n. gen., with the description of a new species (Nymphalidae: Satyrinae: Euptychiina). Tropical Lepidoptera Research, 29, 29 - 44.","Andrade-Correa, G., Viloria, A´., Henao-Banol, E. R. & Le Crom, J. F. (2019) Generic reassignments of satyrine butterflies from Colombia and Venezuela (Lepidoptera: Nymphalidae). Anartia, 28, 26 - 46.","Benmesbah, M. (2015) Description de deux nouvelles especes de Satyrinae de Guyane franCaise (Lepidoptera, Nymphalidae). Bulletin de la Societe Entomologique de France, 120, 147 - 156.","Brevignon, C. (2005) Description de nouveaux Satyrinae provenant de Guyane franCaise (Lepidoptera, Nymphalidae). Lambillionea, 105, 393 - 404.","Freitas, A., Mielke, O., Moser, A., Silva-Brandao t, K. & Iserhard, C. (2011) A new genus and species of Euptychiina (Lepidoptera: Nymphalidae: Satyrinae) from southern Brazil. Neotropical Entomology, 40, 231 - 237.","Huertas, B. (2011) A new species of satyrid butterfly from Peru (Lepidoptera: Nymphalidae: Satyrinae: Euptychiini). Zootaxa, 2802, 63 - 68.","Huertas, B., Rios-Malaver, C. & Le Crom, J. F. (2009) A new species of Splendeuptychia from the Magdalena Valley in Colombia (Lepidoptera: Nymphalidae: Satyrinae). Zootaxa, 2014, 51 - 58.","Willmott, K. R., Lamas, G., Radford, J. T., Marin, M. A., Nakahara, S., Espeland, M. et al. (2018) A distinctive new species of cloud forest Euptychiina (Lepidoptera: Nymphalidae: Satyrinae) from Ecuador and Peru. Tropical Lepidoptera Research, 28, 39 - 45.","Chazot, N., Willmott, K. R., Lamas, G., Freitas, A. V. L., Piron-Prunier, F., Arias, C. F. et al. (2019) Renewed diversification following Miocene landscape turnover in a Neotropical butterfly radiation. Global Ecology and Biogeography, 28, 1118 - 1132.","Huertas, B. (2014) Systematics of the genus Splendeuptychia (Forster, 1964) and implications of taxonomy in conservation assessments. Chapter 4. In Evaluating the conservation status of Neotropical butterflies and the impact of systematics on threat assessments (PhD thesis). University College London.","Janzen, D. H. & Hallwachs, W. (2022) Area de Conservacion´Guanacaste (ACG), northwestern Costa Rica - caterpillars, pupae, butterflies & moths [WWW document]. Available from: http: // janzen. sas. upenn. edu / caterpillars / database. lasso [Accessed on 1 st January 2022]","Hamm, C. A., Williams, B. L. & Landis, D. A. (2013) Natural history and conservation status of the endangered Mitchell ' s satyr butterfly: synthesis and expansion of our knowledge regarding Neonympha mitchellii mitchellii French 1889. Journal of the Lepidopterists' Society, 67, 15 - 28.","Murray, D. L. (2001 b) Systematics of euptychiine butterflies (Nymphalidae: Satyrinae: Euptychiina) based on larval morphology and DNA sequence data and the evolution of life history traits (PhD thesis). Louisiana State University. 367 pp."]}

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."]}

Scriptor Nakhara & Espeland, 2020 sphenophorus Lamas & Nakahara, 2020 [Nakahara et al. (2020, PeerJ, DOI: 10.7717/peerj.10324)]

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."]}

Erichthodes Forster, 1964 antonina (C. Felder & R. Felder, 1867) = erichtho (Butler, 1867), 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

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

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

Chloreuptychia Forster, 1964 agatha (Butler, 1867) = agathina (Weymer, 1911) amethysta 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)] chlorimene (Hübner, [1819]), repl. name = chloris (Cramer, 1780), preocc. (not Fabricius, 1775) gordonsmalli Thurman, MacDonald & Nakahara, 2021 [Nakahara et al. (2021, Tropical Lepidoptera Research, 31(3): 145-150)], 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 63, 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

Stephenympha Viloria, 2022 arius (Weymer, 1911) [Viloria (2022, Anartia, 34: 28-42)] eriphule (Butler, 1867) [Viloria (2022, Anartia, 34: 28-42)] pauliana Viloria, 2022 [Viloria (2022, Anartia, 34: 28-42)], 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 63, DOI: 10.1111/syen.12590, http://zenodo.org/record/7909395

Taguaiba Freitas, Zacca & Siewert genus novum. Type species — Taygetis drogoni Siewert, Zacca et al., 2013, by present designation. Zoobank registration: https://zoobank.org/Nomenclatural Acts/A1AE37F3-5212-453C-AADD-A87F452EFA5D Systematic placement and diagnosis. Taguaiba gen.n. is a mem-ber of the ‘ Taygetis clade’, where it is sister to a clade containing Pseudodebis + Taygetis (Figure 7, FULL dataset, SH-aLRT 100, UFB 98). Species of Taguaiba gen.n. resemble species of Pseudodebis in wing pattern, but can be distinguished from the latter by: (a) the acute apex of the FW in cell R 3 -R 4 (Figure 21), which is rounded in Pseudodebis, although this character is somewhat variable in some species, for example, T. servius comb.n.; (b) the male genitalia of Taguaiba gen. n. species have elongated brachia directed upwards and no cornuti on the vesica (brachia are short and cornuti are present in Pseudodebis — see illustrations in Siewert et al. [2013] and Forster [1964, pp. 76 – 77]); (c) the female genitalia have the lamella antevaginalis fused to the lamella post-vaginalis, forming a sclerotized sterigma that can exhibit lateral projections (in T. drogoni comb.n., T. ypthima comb.n. and T. fulginia comb.n.), a ductus bursae that is sclerotized in the posterior region, and a well-developed corpus bursae with the paired signa occupying the entire bursae. Etymology. The generic name is based on that of the Taguaîba, an evil spirit from the mythology of the Brazilian indigenous people Tupinambá (Métraux, 1950), whose original distribution includes the core distribution of the species of this new genus. Taguaiba should be treated as a neuter noun in the nominative singular. Description (Figure 21). Some notable characters include the following: 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. Large Euptychiina (FW length typically 32 – 38 mm), FW with apex acute in cell R 3 -R 4. No strong sexual dimorphism: VFW with four small circular ocelli from R 4 -Cu 1, each with a well-marked white pupil, and wide brown sub-basal line inside the discal cell; VHW with five ocelli from Rs-Cu 2, each with a well-marked white pupil, and a narrow brown sub-basal line from costal to anal margin. Males with androconial scales on DFW and DHW. Male genitalia with a pair of elongated brachia directed upwards with or without ventral projection at base, apex of valvae serrated or hook-like, saccus cylindrical and elongated, vesica without cornuti (see illustrations in Siewert et al., 2013, p. 23, figure 6). Female genitalia with a sclerotized sterigma with or without lateral projections, ductus bursae sclerotized at the posterior region and corpus bursae well developed with the paired signa occupying the entire bursae (see illustrations in Siewert et al., 2013, p. 24, figure 7). Distribution and natural history (Figure 22). Siewert et al. (2013) summarized the distribution (also shown here in Figure 22), biology and taxonomic history of members of this genus. Discussion. Using DNA barcodes and up to three nuclear genes of 52 species within the ‘ Taygetis clade’ plus seven euptychiine out-groups, Matos-Maraví et al. (2013) proposed the exclusion of T. ypthima and T. rectifascia from Taygetis and indicated that a new genus should be described to contain both species; that genus is described here as Taguaiba gen.n. The morphology of this group of species was subsequently described (Siewert et al., 2013). Taguaiba ypthima comb.n. and Taguaiba rectifascia comb.n. also share morphological similarities in their wing pattern elements, venation and male and female genitalia (see illustrations in Siewert et al., 2013), which are shared with Taguaiba drogoni comb.n., Taguaiba fulginia comb.n. and Taguaiba servius comb.n, all placed in the ‘ Taygetis ypthima species group’ by Siewert et al. (2013). In our study, the ‘ T. ypthima species group’ and respective subgroups as proposed in Siewert et al. (2013) were recovered as monophyletic with the following relationships: ([T. drogoni comb.n. + T. ypthima comb.n.] [T. rectifascia comb. n. + T. fulginia comb.n.]). Currently, T. servius comb.n. is only known from old collection specimens, which has so far precluded the obtaining of DNA sequences. The inclusion of T. servius comb.n. in Taguaiba gen.n. is thus based on morphology and should be tested in future studies. In the study of Matos-Maraví et al. (2013) study, T. ypthima comb.n. and T. rectifascia comb.n. were sister to Pseudodebis. This relationship was not recovered in our study, in which species of Taguaiba gen.n. appear stably as sister to a clade containing Pseudodebis + Taygetis (Figure 7, FULL dataset, SH-aLRT 100, UFB 98). Taguaiba Freitas, Zacca & Siewert, gen.n. drogoni (Siewert, Zacca, Dias & Freitas, 2013), comb.n., was Taygetis [Siewert et al. (2013, ZooKeys, 356: 11-29)] fulginia (D’Almeida, 1922), comb.n., was Taygetis [Siewert et al. (2013, ZooKeys, 356: 11-29)] rectifascia (Weymer, 1907), comb.n., was Taygetis = stigma (Weymer, 1907), comb.n., was Taygetis = latifascia (Weymer, 1907), comb.n., was Taygetis = epithyma (Forster, 1964), nom. nud., comb.n., was Taygetis servius (Weymer, 1910), comb.n., was Taygetis [Siewert et al. (2013, ZooKeys, 356: 11-29)] ypthima (Hübner, [1821]), comb.n., was Taygetis = xantippe (Butler, 1870), comb.n., was Taygetis = ophelia (Butler, 1870), comb.n., was Taygetis = semibrunnea (Weymer, 1910), comb.n., was Taygetis = lineata (Kivirikko, 1936), comb.n., was Taygetis, 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 28-61, DOI: 10.1111/syen.12590, http://zenodo.org/record/7909395, {"references":["Siewert, R. R., Zacca, T., Dias, F. M. S., Freitas, A. V. L., Mielke, O. H. H. & Casagrande, M. M. (2013) The \" Taygetis ypthima species group \" (Lepidoptera, Nymphalidae, Satyrinae): taxonomy, variation and description of a new species. ZooKeys, 356, 11 - 29.","Forster, W. (1964) Beitrage zur Kenntnis der Insektenfauna Boliviens XIX. Lepidoptera III. Satyridae. Veroffentlichungen der Zoologischen Staatssammlung Munchen, 8, 51 - 188.","Metraux, A. (1950) A religiao tdos Tupinambas e suas relaCOes com das demais tribus Tupi-Guaranis. Brasiliana, 267, 1 - 421.","Matos-Maravi, P. F., Pena, C., Willmott, K. R., Freitas, A. V. L. & Wahlberg, N. (2013) Systematics and evolutionary history of butterflies in the \" Taygetis clade \" (Nymphalidae: Satyrinae: Euptychiina): towards a better understanding of Neotropical biogeography. Molecular Phylogenetics and Evolution, 66, 54 - 68."]}

Harjesia Forster, 1964 argentata Nakahara, Zacca & Lamas, 2018 [Nakahara et al. (2018, Insecta Mundi, 0639: 1-38)] blanda (MÖschler, 1877) obscura (Butler, 1867) = eremita (Weymer, 1911), 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 61, DOI: 10.1111/syen.12590, http://zenodo.org/record/7909395

Pseudodebis Forster, 1964 = Taygetomorpha L.D. Miller, 2004 celia (Cramer, 1779) [Matos-Maraví et al. (2012, Molecular Phylogenetics and Evolution, 66: 54-68)] = phorcys (Fabricius, 1793) [Matos-Maraví et al. (2012, Molecular Phylogenetics and Evolution, 66: 54-68)] = keneza (Butler, 1869) [Matos-Maraví et al. (2012, Molecular Phylogenetics and Evolution, 66: 54-68)] darrenthroopi Nakahara & Willmott, 2022 [Nakahara et al. (2022, Neotropical Entomology, 51: 536-556)] dubiosa Forster, 1964 euptychidia (Butler, 1868) = modesta (Hayward, 1949) = theresa Forster, 1964, nom. nud. hartmanni Nakahara & MacDonald, 2021 [Nakahara et al. (2021, Tropical Lepidoptera Research, 31(1): 35-41)] marpessa (Hewitson, 1862) = arpi Forster, 1964 nakamurai Nakahara & Willmott, 2021 [Nakahara et al. (2021, Invertebrate Systematics, 35: 158-180)] pieti Nakahara & Willmott, 2021 [Nakahara et al. (2021, Invertebrate Systematics, 35: 158-180)] puritana (Weeks, 1902) [Matos-Maraví et al. (2012, Molecular Phylogenetics and Evolution, 66: 54-68)] = haenschi (Weymer, 1910) [Matos-Maraví et al. (2012, Molecular Phylogenetics and Evolution, 66: 54-68)] = oliva (Forster, 1964), nom. nud. [Matos-Maraví et al. (2012, Molecular Phylogenetics and Evolution, 66: 54-68)] tigrillo Nakahara & Willmott, 2022 [Nakahara et al. (2022, Neotropical Entomology, 51: 536-556)] valentina (Cramer, 1779) = tarmensis (Prüffer, 1922) vrazi (Kheil, 1896) [Nakahara et al. (2018, Insecta Mundi, 0639: 1-38)] zimri (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 page 61, DOI: 10.1111/syen.12590, http://zenodo.org/record/7909395

Pindis R. Felder, 1869 squamistriga R. Felder, 1869 = zabdi (Butler, 1869) = obnixa Draudt, 1931, 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 59, DOI: 10.1111/syen.12590, http://zenodo.org/record/7909395

Lazulina Willmott, Nakahara & Espeland, genus novum. Type species — Euptychia hewitsonii Butler, 1867, by present designation. Zoobank registration: https://zoobank.org/Nomenclatural Acts/2F9B7D53-51B3-4A62-B5F1-AE2312CD4163 Systematic placement and diagnosis. Lazulina gen.n. is wellsupported as monophyletic (SH-aLRT = 100, UFB = 100), but only moderately supported as sister to Cristalinaia (SH-aLRT = 89.2, UFB = 90), with the clade containing these two genera placed without strong support as sister to the Hermeuptychia clade. The new genus can be distinguished from all other Euptychiina by a rather large number of distinctive possible synapomorphies in the male, including: (a) a dark band of androconial scales just posterior of the cubital vein on the DFW extends from 2A-Cu 2 into Cu 1 -M 3, with associated modified scales (Figure 11a – c; see Description below for details). (b) an elongate cluster of long, hair-like scales (‘hair-pencil’) is located near the HW base in cell 3A-2A, the distal tip of which covers an elongate, oval patch of modified scales surrounding vein 2A (Figure 11a,c). A more elongate patch of somewhat similar androconial scales is present in HW cell 2A-Cu 2 in Atlanteuptychia (Freitas et al., 2013). (c) the basal half of HW vein 2A is bowed outwards, towards the adjacent ‘hair-pencil’ (Figure 11). (d) Fourth and fifth sternites of the male abdomen are reduced to a small, sclerotized band at the anterior edge of each segment, with the remainder of each segment forming a flexible pocket that contains dense black elongate rectangular androconial scales, which clearly stand out beyond the ventral abdominal edge in lateral view (Figure 11g). Similar scales are present dorsally on the terminal two abdominal segments of Atlanteuptychia (Freitas et al., 2013). A further possibly distinctive character is the reduction of the middle of FW vein M 2 -M 3, and numerous other characters distinguish the genus from its possible sister genus, the highly autapomorphic Cristalinaia (see Freitas, Mota, Zacca, & Barbosa, 2019). Overall, the iridescent blue dorsal wing colour of Lazulina gen.n. species readily distinguishes them from most euptychiines, except for females in comparison with several species of Chloreuptychia and Pseudeuptychia. Lazulina gen.n. can be distinguished by the presence of yellowish or reddish brown scaling within the dark band posterior of the VFW apical ocellus, and by the VHW ocellus in M 3 -M 2 being basally displaced in comparison with that in M 2 -M 1. Etymology. The generic name is a feminine noun in the nominative singular, derived from the Latin adjective ‘lazulinus’, meaning something that is blue, in reference to the distinctive blue dorsal wing markings of this genus. Description (Figures 11 and 12). Some notable characters include: eyes naked; pterothoracic legs dorsally 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 Euptychiina (FW length typically 17 – 21 mm), FW triangular and somewhat elongate, HW margin not scalloped, slightly angled at end of vein Cu 1 and slightly elongated towards tornus. Sexes dimorphic, dorsal wings dark brown: Male: HW with iridescent blue to purple scaling at least in posterior half, to covering most of wing, L. hewitsonii comb.n. partially translucent pale greyish in HW discal cell and with iridescent blue scaling along anal margin and distal margin of FW; middle of FW with complex configuration of androconial scales, comprising long hair-like scales in posterior third of discal cell, which appear to be ‘swept-back’ towards cubital vein, cubital vein bordered posteriorly by a band of broad, short, rectangular pale brown scales sparsely covered with long hair-like scales, then a band of dense, dark androconial scales that are attenuated towards tip then terminate in a curved, brush-like structure; 8th abdominal tergite reduced dorsally, leaving a sclerotized strip along anterior edge; genitalia with downwardly curving uncus longer than tegumen, brachia short and pointing ventrally of uncus, valva ventral edge distinctly angled in middle, terminating in blunt tip; saccus short (similar in length to tegumen); aedeagus short, anterior portion opening dorso-anteriorly (rather than dorsally) and slightly flared in dorsal view, with two narrow bands of cornuti. Female: lacking androconial scales, with faint purplish blue iridescence over both wings or HW only, not extending to distal margins. Ventral wings with ground colour ranging from brown to silvery grey; dark reddish brown discal and postdiscal lines traversing both wings, discal line angled inwards at HW anal margin, postdiscal line displaced towards tornus in cell 2A-Cu 2; VFW with a single subapical ocellus in cell M 2 -M 1, black with a yellow ring and 1 – 2 silvery elongate pupils (when 2 pupils are present they unite to form a ‘C’-shape), placed within a broad, dark brown postdiscal band (umbra); posterior of ocellus, VFW umbra has dark orange-brown scaling or yellowish brown scaling, in some species bordered basally by a pale purplish band; two dark marginal lines, straight and parallel to wing margin; VHW has five postdiscal ocelli as follows: In Cu 2 -Cu 1, largest ocellus, black with yellow border that extends into adjacent cells, with two elongate silver dashes as pupils; in Cu 1 -M 3, elongate brown ocellus with yellow border that may be absent distally, with large central elongate silver pupil; in M 3 -M 2, similar to preceding but smaller and displaced basally; in M 2 -M 1, black ocellus with yellow ring filling cell, two silver pupils united to form distally pointing ‘C’-shape; in M 1 -Rs, similar to preceding but smaller; ocelli bordered distally by dark brown, especially distal of smaller ocelli, then pale purplish surrounding two dark marginal lines which become gradually thicker and more reddish towards tornus; genitalia with each sternite reduced posteriorly; eighth tergite reduced to small sclerotized posterior patch, intersegmental membrane between seventh and eighth abdominal segments somewhat pleated and expandable with an oval sclerotized plate ventrally, eighth segment with large irregular lateral sclerotized plate, lamella antevaginalis and antrum unsclerotized, ductus bursae narrow and unsclerotized, corpus bursae small, circular and with two narrow signa. Distribution and natural history (Figure 13). Lazulina gen.n. species occur from sea level to 1200 m in the Amazon, Guianas and Trinidad, with the highest local diversity occurring in the upper Amazon. All species are found in the understory of disturbed to primary rainforest, often near forest edges at rivers, streams or palm swamps, where they typically occur as solitary individuals. There is no published information on the hostplants or immature stages, although some data on the immature stages of L. tolumnia comb.n. are provided in Murray ’ s (2001b) unpublished dissertation. Discussion. Butler (1867) described the type species for this genus, Euptychia hewitsonii, based on both male and female specimens, from Pará (Brazil), then in the collections of Bates, Salvin, Hewitson and the ‘B[ritish].M[useum].’ (now NHMUK), along with a male or males from Ega (now Tefé, Brazil), then in the last of these collections. The description is detailed, notably mentioning the somewhat translucent basal area of the wings that are distinctive to this species, and the ventral surface of a male was illustrated on pl. 40, Figure 4. A syntype in the NHMUK was examined and matches the original illustration. The three described species placed here in Lazulina gen.n were placed until now in Chloreuptychia (Forster, 1964; Lamas, 2004), a genus now split into four genera in four clades (e.g. Nakahara, Lamas, et al., 2019; this publication). However, Weymer (Weymer, 1911, p. 220) placed the three described Lazulina species in a distinct ‘Tolumnia Group’, correctly noting that males of these species shared a band of androconial scales just posterior of the discal cell on the DFW. Forster (1964, p. 121) used this character in his key to separate out these three species, but nevertheless grouped them with other unrelated ‘blue’ species in his new genus Chloreuptychia. The molecular analysis here shows that Lazulina gen.n. are not closely related to any species formerly included in Chloreuptychia, and indeed the relationships of this highly distinctive genus are still not confidently known. There are a number of remarkably similar structures in Lazulina gen.n. and Atlanteuptychia, including androconial scales in the middle of the DFW, a DHW androconial ‘hair-pencil’, and dense black androconial scales on the male abdomen. Nevertheless, differences in the morphology and position of the androconial structures support independent derivation, the female genitalia and wing patterns show no particular similarity, and molecular data, based on multiple individuals of each genus, indicate that these are not closely related genera, placing them in distinct clades. Lazulina Willmott, Nakahara & Espeland, gen.n. catharina (Staudinger, [1886]), comb.n., was Chloreuptychia hewitsonii (Butler, 1867), comb.n., was Chloreuptychia = polla (MÖschler, 1883), comb.n., was Chloreuptychia tolumnia (Cramer, 1777), comb.n., was Chloreuptychia, 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 20-58, 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.","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.","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.","Weymer, G. (1911) 4. Familie: Satyridae. In: Seitz, A. (Ed.) Die Gross- Schmetterlinge der Erde. Stuttgart: A. Kernen, pp. 173 - 280."]}

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)], 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 67, DOI: 10.1111/syen.12590, http://zenodo.org/record/7909395

Euptychia Euptychia is strongly supported as a sister to the remainder of the subtribe (Figures 2, S 2 and S 3) as also found in Espeland et al. (2019a). The systematic placement of Euptychia has been unstable in previous phylogenetic studies (Murray & Prowell, 2005; Peña et al., 2010, 2011). Currently, 37 valid species are recognized (Fratello et al., 2015; Freitas, Wahlberg, et al., 2012; Nakahara et al., 2014, 2017; Nakahara, Hall, et al., 2015; Nakahara, Janzen, et al., 2015; Nakahara, Llorente-Bousquets, et al., 2015; Nakahara, Vega, & Willmott, 2016; Neild et al., 2014, 2015). Only a subset of species was included here, and several taxonomic changes, in addition to the description of over a dozen species, will be made in a forthcoming revision of the genus, and thus the genus is estimated to include more than 50 species (Nakahara et al., in preparation). Members of Euptychia are relatively small for the subtribe, with the forewing length often less than 20 mm. Except for a few species, they have sexually monomorphic wing patterns often marked prominently with ventral bands and submarginal eyespots and many species exhibit translucence. Possible synapomorphies and/or distinctive characters for Euptychia include the following: (a) Third segment of labial palpi shorter than one-fourth of second segment in length; (b) absence of tibial spurs in midleg and hindleg; (c) absence of lateral spines on tibia; (d) presence of the forewing recurrent vein in the discal cell; (e) absence of basal swelling of the forewing cubital vein; (f) humeral vein relatively reduced in comparison with other Euptychiina; (g) developed male eighth tergite and sternite; (h) presence of the sclerotized region of the eighth abdominal segment in the female, located at the very basal side of the eighth abdominal segment; (i) absence of the lateral sclerotization of the 8th abdominal segment of the female; (j) origin of the ductus seminalis at the posterior end of the ductus bursae In fact, many of these characters are unique to Euptychia (Nakahara, unpublished data), highlighting the distinctiveness of this lineage within the subtribe. Members of Euptychia range from north-western Mexico to south-eastern Brazil, and somewhat unusually for the subtribe, the genus appears to be more diverse in lower montane regions and adjacent lowlands than in lowland forest far from mountains. Adult males of some species exhibit perching behaviour, often with several individuals in close proximity, mostly in light gaps or sunflecks and often on hilltops. Although most known host plants for Euptychiina are either bamboo or other grasses in the family Poaceae, larvae of Euptychia notably feed instead on some species of Selaginella (Lycopsida), as well as a single record utilizing Neckeropsis (Neckeraceae) (Brévignon, 2008; Freitas, Mota, Barbosa, & Carreira, 2019; Hamm & Fordyce, 2016; Singer et al., 1971; Singer & Mallet, 1986). FIGURE 13 Lazulina gen.n. Species diversity mapped on a FIGURE 16 Saurona gen.n. Species diversity mapped on a FIGURE 20 Argentaria gen.n. Species diversity mapped on a FIGURE 25 Xenovena murrayae comb.n. Locality records. FIGURE 29 Trico tricolor comb.n. Locality records. FIGURE 32 Occulta ocnus comb.n. Locality records. FIGURE 36 Deltaya gen.n. Species diversity mapped on a FIGURE 40 Modica gen.n. Species diversity mapped on a Euptychia Hübner, 1818 = Caenoptychia Le Cerf, 1919 = Ristia Gagarin, 1936 alacristata Neild, Nakahara & Fratello, 2014 [Neild et al. (2014, Tropical Lepidoptera Research, 24(1): 4-9)] aquila Fratello, Nakahara & Brévignon, 2015 [Fratello et al. (2015, Journal of the Lepidopterists' Society, 69(4): 293-306)] atlantica Nakahara & Freitas, 2017 [Nakahara et al. (2017, Neotropical Entomology, 46: 302-309)] attenboroughi Neild, Nakahara, Fratello & Le Crom, 2015 [Neild et al. (2015, ZooKeys, 541: 87-108)] audacia Brévignon, Fratello & Nakahara, 2015 [Fratello et al. (2015, Journal of the Lepidopterists' Society, 69(4): 293-306)] boulleti (Le Cerf, 1919) [Freitas et al. (2012, Neotropical Entomology, 41(6): 461-467)] = virgata Joicey & Talbot, 1924 [Freitas et al. (2012, Neotropical Entomology, 41(6): 461-467)] = tigrina (Gagarin, 1936) [Freitas et al. (2012, Neotropical Entomology, 41(6): 461-467)] cesarense Pulido, Andrade, Peña & Lamas, 2011 [Pulido et al. (2011, Zootaxa, 2906: 43-51)] - obtusa Nakahara, 2015 [Nakahara et al. (2015, Tropical Lepidoptera Research, 25(2): 63-79)] - viloriai Andrade, Pulido, Peña & Lamas, 2011 [Pulido et al. (2011, Zootaxa, 2906: 43-51)] efraini Ríos, 2019 [Ríos (2019, Anartia, 29: 49-53)] = similis Henao, 2019, preocc. (not Butler, 1867) [Ríos (2019, Anartia, 29: 49-53)] enyita Nakahara, Lamas & Willmott, 2015 [Nakahara et al. (2015, Tropical Lepidoptera Research, 25(2): 63-79)] enyo Butler, 1867 favonius Nakahara, Vega & Willmott, 2016 [Nakahara et al. (2016, Zootaxa, 4184: 358-366)] fernandae Nakahara & Willmott, 2015 [Nakahara et al. (2015, Tropical Lepidoptera Research, 25(2): 63-79)] fetna Butler, 1870 granatina Nakahara, Le Crom & Hall, 2015 [Nakahara et al. (2015, Tropical Lepidoptera Research, 25(2): 63-79)] hannemanni Forster, 1964 insolata Butler & Druce, 1872 = macrophthalma Staudinger, 1876 jesia Butler, 1869 juanjoi Le Crom, Nakahara & Lamas, 2015 [Nakahara et al. (2015, Journal of Research on the Lepidoptera, 48: 59-63)] lacandona Warren & Nakahara, 2015 [Nakahara et al. (2015, Journal of Research on the Lepidoptera, 48: 51-57)] marceli Brévignon, 2005 [Brévignon (2005, Lambillionea, 105(3)(1): 393-404)] - divisa Benmesbah, Costa, Attal & Viloria, 2021 [Costa et al. (2021, Antenor, 8(1): 2-28)] meta Weymer, 1911 mollina (Hübner, [1818]) = mollina (Hübner, 1808), nom. nud. - suzannae Brévignon, 2005 [Brévignon (2005, Lambillionea, 105(3)(1): 393-404)] mollis Staudinger, 1876 neblina Warren & Nakahara, 2015 [Nakahara et al. (2015, Journal of Research on the Lepidoptera, 48: 51-57)] neildi Brévignon, 2005 [Brévignon (2005, Lambillionea, 105(3)(1): 393-404)] padroni Nakahara, Lamas & Willmott, 2015 [Nakahara et al. (2015, Tropical Lepidoptera Research, 25(2): 63-79)] pegasus Nakahara & J. Hall, 2015 [Nakahara et al. (2015, Tropical Lepidoptera Research, 25(2): 63-79)] picea Butler, 1867 pillaca Nakahara & Willmott, 2015 [Nakahara et al. (2015, Tropical Lepidoptera Research, 25(2): 63-79)] roraima Nakahara, Fratello & Harvey, 2014 [Nakahara et al. (2014, Zootaxa, 3881(3): 291-300)] rubrofasciata L.D. Miller & J.Y. Miller, 1988 rufocincta Weymer, 1911 sarah Benmesbah & Viloria, 2019 [Costa et al. (2019, Anartia, 29: 20-48)] sophiae Zacca, Nakahara, Dolibaina & Dias, 2015 [Neild et al. (2015, ZooKeys, 541: 87-108)] truncata Nakahara & J. Hall, 2015 [Nakahara et al. (2015, Tropical Lepidoptera Research, 25(2): 63-79)] westwoodi Butler, 1867 - muli Brévignon, 2005 [Brévignon (2005, Lambillionea, 105(3)(1): 393-404)] woroina Viloria & Benmesbah, 2020 [Costa et al. (2020, Antenor, 7(1): 19-41)], 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 7-54, 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.","Pena, C., Nylin, S. & Wahlberg, N. (2011) The radiation of Satyrini butterflies (Nymphalidae: Satyrinae): a challenge for phylogenetic methods. Zoological Journal of the Linnean Society, 161, 64 - 87.","Fratello, S. A., Nakahara, S., Brevignon, C. R. & Harvey, D. J. (2015) Two new species of Euptychia Hubner, 1818 (Lepidoptera: Nymphalidae: Satyrinae) from the Guiana shield, with notes on E. marceli Brevignon, 2005 and E. rufocincta Weymer, 1911. Journal of the Lepidopterists' Society, 69, 293 - 306.","Nakahara, S., Fratello, S. A. & Harvey, D. J. (2014) A new species of Euptychia Hubner, 1818 (Lepidoptera: Nymphalidae: Satyrinae: Satyrini) from Mount Roraima, Guyana. Zootaxa, 3881, 291 - 300.","Nakahara, S., Barbosa, E. P. & Freitas, A. V. L. (2017) A potentially endangered new species of Euptychia Hubner, 1818 (Lepidoptera: Nymphalidae: Satyrinae) from the Atlantic coastal forest of Brazil. Neotropical Entomology, 46, 1 - 8.","Neild, A. F. E., Nakahara, S., Fratello, S. A. & Harvey, D. J. (2014) A new species of Euptychia Hubner, 1818 (Nymphalidae: Satyrinae: Satyrini) from the Amazon basin and the Guianas. Tropical Lepidoptera Research, 24, 4 - 9.","Kozak, K. M., Wahlberg, N., Neild, A. F. E., Dasmahapatra, K. K., Mallet, J. & Jiggins, C. D. (2015) Multilocus species trees show the recent adaptive radiation of the mimetic Heliconius butterflies. Systematic Biology, 64, 505 - 524.","Brevignon, C. (2008) Inventaire des Satyrinae de Guyane FranCaise (Lepidoptera: Nymphalidae). In: Lacomme, D. & Manil, L. (Eds.) Lepidopteres de Guyane. Paris: Lepidopteristes de France, pp. 62 - 94.","Hamm, C. A. & Fordyce, J. A. (2016) Selaginella and the satyr: Euptychia westwoodi (Lepidoptera: Nymphalidae) oviposition preference and larval performance. Journal of Insect Science, 16, 1 - 4.","Singer, M. C., Ehrlich, P. R. & Gilbert, L. E. (1971) Butterfly feeding on lycopsid. Science, 172, 1341 - 1342.","Singer, M. C. & Mallet, J. (1986) Moss-feeding by a satyrine butterfly. Journal of Research on the Lepidoptera, 24, 392.","Neild, A. F. E., Nakahara, S., Zacca, T., Fratello, S., Lamas, G., Le Crom, J. - F. et al. (2015) Two new species of Euptychia Hubner, 1818 from the upper Amazon basin (Lepidoptera, Nymphalidae, Satyrinae). ZooKeys, 541, 87 - 108.","Freitas, A. V. L., Kaminski, L. A., Mielke, O. H. H., Barbosa, E. P. & Silva-Brandao t, K. L. (2012) A new species of Yphthimoides (Lepidoptera: Nymphalidae: Satyrinae) from the southern Atlantic forest region. Zootaxa, 3526, 31 - 44.","Nakahara, S., Hall, J. P. W., Lamas, G. & Willmott, K. R. (2015) Seven new species and one new subspecies of Euptychia Hubner, 1818 (Lepidoptera: Nymphalidae: Satyrinae) from the tropical Andes. Tropical Lepidoptera Research, 25, 63 - 79.","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.","Nakahara, S., Vega, G. & Willmott, K. R. (2016) Description of a new species of Euptychia Hubner, 1818 (Lepidoptera: Nymphalidae: Satyrinae) from the Western Andes. Zootaxa, 4184, 358 - 366.","Nakahara, S., Barbosa, E. P., Marin, M. A., Freitas, A. V. L., Pomerantz, T. & Willmott, K. R. (2016) Graphita gen. nov., a new genus for Neonympha griphe C. Felder & R. Felder, 1867 (Lepidoptera, Nymphalidae, Satyrinae). Neotropical Entomology, 45, 1 - 17.","Forster, W. (1964) Beitrage zur Kenntnis der Insektenfauna Boliviens XIX. Lepidoptera III. Satyridae. Veroffentlichungen der Zoologischen Staatssammlung Munchen, 8, 51 - 188.","Butler, A. G. (1869) Lepidoptera Exotica, or descriptions and illustrations of exotic Lepidoptera. London: E. W. Janson, p. 339.","Brevignon, C. (2005) Description de nouveaux Satyrinae provenant de Guyane franCaise (Lepidoptera, Nymphalidae). Lambillionea, 105, 393 - 404.","Viloria, A´. L. (2021) The Hermeuptychia papers. Anartia, 32, 26 - 52.","Weymer, G. (1911) 4. Familie: Satyridae. In: Seitz, A. (Ed.) Die GrossSchmetterlinge der Erde. Stuttgart: A. Kernen, pp. 173 - 280.","Viloria, A´. L. & Luis-Martinez, A. (2019) Llorenteana, a new butterfly genus from the American continent (Lepidoptera: Nymphalidae: Satyrinae). Dugesiana, 26, 167 - 172."]}

Carminda Ebert & Dias, 1998 = Carminda Ebert & Dias, 1997, unavailable (no type species designated) griseldis (Weymer, 1911) [Dias (2011, Brazilian Journal of Biology, 71(2): 537-540)] paeon (Godart, [1824]) [Dias (2011, Brazilian Journal of Biology, 71(2): 537-540)] = marmorata (Butler, 1867) [Dias (2011, Brazilian Journal of Biology, 71(2): 537-540)] = albiplaga (J. Zikán & W. Zikán, 1968), nom. nud. [Lamas, unpublished data] = simillima (J. Zikán & W. Zikán, 1968), nom. nud. [Lamas, unpublished data] surpresa Barbosa, Aguiar, Rosa, Zacca & Freitas, 2020 [Barbosa et al. (2020, Zootaxa, 4819(1): 128-142)] umuarama Ebert & Dias, 1997 [Dias (2011, Brazilian Journal of Biology, 71(2): 537-540)] = elisae (J. Zikán & W. Zikán, 1968), nom. nud. [Lamas, unpublished data], 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 55-56, DOI: 10.1111/syen.12590, http://zenodo.org/record/7909395

Sepona Freitas & Barbosa, 2016 punctata (Weymer, 1911) [Freitas et al. (2016, Revista Brasileira de Entomologia, 60(2): 157-165)] = griseola (Weymer, 1911) [Freitas et al. (2016, Revista Brasileira de Entomologia, 60(2): 157-165)] = indecisa (Ribeiro, 1931) [Freitas et al. (2016, Revista Brasileira de Entomologia, 60(2): 157-165)], 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 61, DOI: 10.1111/syen.12590, http://zenodo.org/record/7909395

Forsterinaria Gray, 1973 = Haywardina Forster, 1964, preocc. (not Aczél, 1952 (Diptera)) = Forsterina Lamas, 1977, missp. = Forsterinonia Brown, 1992, missp. = Guaianaza Freitas & Peña, 2006 anophthalma (C. Felder & R. Felder, 1867) antje Peña & Lamas, 2005 [Peña & Lamas (2005, Revista peruana de Biología, 12(1): 5-48)] boliviana (Godman, 1905) = vetula (Forster, 1964), nom. nud. chaniorum Viloria & Costa, 2019 [Costa et al. (2019, Anartia, 29: 20-48)] coipa Peña & Lamas, 2005 [Peña & Lamas (2005, Revista peruana de Biología, 12(1): 5-48)] difficilis (Forster, 1964) = anachoreta Pulido & Andrade, 2008 [Zubek & Pyrcz (2011, Genus, 22(2): 217-225)] emo Zubek, Pyrcz & Boyer, 2014 [Zubek et al. (2014, Neotropical Entomology, 43(1): 68-77)] enjuerma Peña & Lamas, 2005 [Peña & Lamas (2005, Revista peruana de Biología, 12(1): 5-48)] falcata Peña & Lamas, 2005 [Peña & Lamas (2005, Revista peruana de Biología, 12(1): 5-48)] guaniloi Peña & Lamas, 2005 [Peña & Lamas (2005, Revista peruana de Biología, 12(1): 5-48)] hannieri Zubek & Pyrcz, 2011 [Zubek & Pyrcz (2011, Genus, 22(3): 523-530)] inornata (C. Felder & R. Felder, 1867) = eusebia (Butler, 1877) - magdalena (Hayward, 1957) [Zubek & Pyrcz (2015, Zootaxa, 4028(4): 579-582)] itatiaia Peña & Lamas, 2005 [Peña & Lamas (2005, Revista peruana de Biología, 12(1): 5-48)] = otomara (J. Zikán & W. Zikán, 1968), nom. nud. [Lamas, unpublished data] necys (Godart, [1824]) = vastata (Butler, 1867) = pseudonecys (Strand, 1916) [Peña & Lamas (2005, Revista peruana de Biología, 12(1): 5-48)] = stellifera (J. Zikán & W, Zikán, 1968), nom. nud. [Lamas, unpublished data] neonympha (C. Felder & R. Felder, 1867) = polyphemus (Butler, 1867) = umbracea (Butler & Druce, 1872) = cyclops (Butler, 1877) pallida Peña & Lamas, 2005 [Peña & Lamas (2005, Revista peruana de Biología, 12(1): 5-48)] - aurita Peña & Lamas, 2005 [Peña & Lamas (2005, Revista peruana de Biología, 12(1): 5-48)] pichita Peña & Lamas, 2005 [Peña & Lamas (2005, Revista peruana de Biología, 12(1): 5-48)] pilosa Peña & Lamas, 2005 [Peña & Lamas (2005, Revista peruana de Biología, 12(1): 5-48)] pronophila (Butler, 1867) [Matos-Maraví et al. (2012, Molecular Phylogenetics and Evolution, 66: 54-68)] = pronophila (Butler, 1867), nom. nud. [Matos-Maraví et al. (2012, Molecular Phylogenetics and Evolution, 66: 54-68)] proxima (Hayward, 1957) pseudinornata (Forster, 1964) punctata Peña & Lamas, 2005 [Peña & Lamas (2005, Revista peruana de Biología, 12(1): 5-48)] pyrczi Peña & Lamas, 2005 [Peña & Lamas (2005, Revista peruana de Biología, 12(1): 5-48)] quantius (Godart, [1824]) = stelligera (Butler, 1874) [Peña & Lamas (2005, Revista peruana de Biología, 12(1): 5-48)] = fabiana (Butler, 1877) [Peña & Lamas (2005, Revista peruana de Biología, 12(1): 5-48)] = morima (Schaus, 1902) = wordhilda (J. Zikán & W. Zikán, 1968), nom. nud. [Lamas, unpublished data] rotunda Peña & Lamas, 2005 [Peña & Lamas (2005, Revista peruana de Biología, 12(1): 5-48)] rustica (Butler, 1868) - villarresi (Dognin, 1887) = weyrauchi (Hayward, 1964) - glendita Peña & Lamas, 2005 [Peña & Lamas (2005, Revista peruana de Biología, 12(1): 5-48)] stella (Hayward, 1957) = howarthi (Hayward, 1962), 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 60-61, 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)], 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 61, DOI: 10.1111/syen.12590, http://zenodo.org/record/7909395

Parataygetis Forster, 1964 albinotata (Butler, 1867) lineata (Godman & Salvin, 1880) = melania (Weymer, 1911) = interrupta (Weymer, 1911, 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 61, DOI: 10.1111/syen.12590, http://zenodo.org/record/7909395

Archeuptychia clade The ‘ Archeuptychia clade’ is a relatively small clade recognized in Espeland et al. (2019a), and that work showed its placement as a weakly supported sister group to the much larger ‘ Splendeuptychia clade’, which is also found in the extended backbone tree in this study (Figure S 1). In the FULL dataset it is, however, weakly supported as sister to a clade including Chloreuptychia and the Pareuptychia clade (Figure S 2), while in the 4GENES dataset (Figure S 3) it is sister to Chloreuptychia, and these two again are sister to the ‘ Splendeuptychia clade’. Previous molecular phylogenetic studies only sampled a few representatives and thus did not result in recognition of this clade (Marín et al., 2017; Murray & Prowell, 2005) or subsumed it within the ‘ Pareuptychia clade’ (Peña et al., 2010), but close relationships among Archeuptychia, species placed here in Pseudeuptychia, Chloreuptychia and Cisandina were variously implied in these studies. The clade (Figure 8) currently includes 16 described species and at least three undescribed species. Currently, four genera, Pseudeuptychia (seven species), Archeuptychia (monotypic), Cisandina (seven species) and Stephenympha (three species) are recognized in the clade. The two sampled Stephenympha species are here found to be sister groups, with some support in the best trees (FULL 90/91, 4 GENES 90.2/88), and are well-supported as members of the ‘ Archeuptychia clade’, but their exact placement within the clade needs to be investigated, since support of relationships within the group are rather low (Figure 8). Here, we provisionally move four species to Pseudeuptychia (comb.n.) that were placed in Chloreuptychia prior to this study, since these species form a strongly supported clade that contains Euptychia languida Butler, 1871, the type of Pseudeuptychia, but that excludes Papilio chloris Cramer, 1780, the type species of Chloreuptychia (Marín et al., 2017; Nakahara et al., in preparation). We do not know of any taxonomic issues surrounding Archeuptychia, and thus this monotypic genus does not require study at present. Members of the ‘ Archeuptychia clade’ are small to medium-sized butterflies, all rather typical of euptychiines in terms of size. However, unlike most other euptychiine groups, the clade harbours some of the more ‘showy’ euptychiines, with many species bearing iridescent blue or green dorsal and/or ventral coloration. Possible synapomorphies and distinctive characters for the clade include the following: (a) lamella antevaginalis at least somewhat sclerotized (the degree of sclerotization is apparently variable, ranging from a rather weakly sclerotized plate in species such as Pseudeuptychia herseis comb.n., to a more heavily sclerotized plate in species such as Archeuptychia cluena). The male genitalia of species in Cisandina show some interesting features, such as a sclerotized portion of the manica extending from the phallobase, a character not observed in other euptychiines. Stephenympha eriphule has a uniformly sclerotized eighth tergite, which is an unusual feature for euptychiines otherwise known only for species in the genus Euptychia. However, a similar form of the eighth tergite is observed in Cristalinaia and species in Amphidecta, but the eighth tergite of these species appears to have a membranous portion in the middle, thus not being uniformly sclerotized. Members of the ‘ Archeuptychia clade’ are known exclusively from east of the Andes, except for Pseudeuptychia hemileuca, which is still known only from a single syntype, that was reportedly collected in western Colombia. Some species are confined to cloud forest along the slopes of the eastern Andes, while the majority of species occur in lowland rainforest from the Amazon basin and/or Guianas, with a few species found in the Atlantic coastal forest of Brazil. Information on the early stage biology for species in the ‘ Archeuptychia clade’ is limited, like many other major euptychiine clades, but the immature stages of Pseudeuptychia marica comb.n. and two species of Cisandina were recently described, with caterpillars feeding on the herbaceous bamboo genera Pariana, Olyra and Taquara (Poaceae: Bambusoideae: Olyreae) (Nakahara, Rodríguez-Melgarejo, et al., 2022; Tejeira et al., 2021), and additional data will soon be published (Corahua-Espinoza et al., in preparation)., 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 17, 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.","Marin, M. A., Pena, C., Uribe, S. I. & Freitas, A. V. L. (2017) Morphology agrees with molecular data: phylogenetic affinities of Euptychiina butterflies (Nymphalidae: Satyrinae). Systematic Entomology, 42, 768 - 785.","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.","Tejeira, R., Ccahuana, R., Hurtado, T., Nakahara, S., See, J., Rodriguez-Melgarejo, M. et al. (2021) Immature stages of Chloreuptychia marica (Weymer, 1911) (Lepidoptera: Nymphalidae: Satyrinae: Satyrini). Tropical Lepidoptera Research, 31, 96 - 100."]}

Megisto clade The ‘Megisto clade’ currently comprises 10 genera, Megisto (with 2 species), Pharneuptychia (5 species, in addition to ‘ Pharneuptychia ’ innocentia in the ‘ Amphidecta clade’), Moneuptychia (11 species), Graphita (1 species), Stegosatyrus (4 species), Koutalina (1 species), Cissia (10 species), Carminda (4 species), Vanima (3 species) and Yphthimoides (21 species). The relationships between genera in our analyses (Figures 4, S 2 and S 3) match those of previous molecular studies (Murray & Prowell, 2005; Peña et al., 2010) and are stable, with the exception of the placement of the genus Koutalina. In the best FULL tree Koutalina is found as sister to the remainder of the clade, except Megisto, but is placed deeper within the clade as sister to Graphita in the second best FULL tree, and as sister to Moneuptychia + Pharneuptychia in the best 4GENES tree. This genus was not included in the backbone tree, and only COI sequences are available for the included samples, which likely explains the instability of this taxon. The recently described Yphthimoides kinyoni (Nakahara, Barbosa, Nakamura, et al., 2021) is found within the genus Yphthimoide s, as expected, in the best FULL tree, but in the second best FULL tree and in the best two 4GENES trees it is placed as sister to Stegosatyrus within Cissia. This instability is likely also caused by its lack of very close relatives within the genus and only short COI sequences being available, but a placement within Yphthimoides is also justified morphologically (Nakahara, Barbosa, Nakamura, et al., 2021). It is estimated that the diversity of this clade will increase with the description of new species for Moneuptychia and Pharneuptychia (Barbosa et al., 2022; Freitas et al., in preparation) in forthcoming papers. Some recent papers provided generic taxonomic revisions and description of new taxa in this clade (Zacca et al., 2013 [Stegosatyrus], Zacca, Casagrande, et al., 2018 [Cissia], Zacca, Casagrande, Mielke, Huertas, Espeland, Freitas, et al., 2020 [Vanima]; Barbosa et al., 2015 [Yphthimoides], Barbosa et al., 2016 [Yphthimoides], Barbosa et al., 2020 [Yphthimoides]; Freitas et al., 2015 [Moneuptychia]; Nakahara, Barbosa, et al., 2016 [Graphita], Nakahara, Barbosa, Nakamura, et al., 2021 [Yphthimoides]; Henao-Bañol & Meneses, 2018 [Pharneuptychia]; Zhang et al., 2020 [Cissia], Zhang et al., 2022 [Cissia]; Benmesbah et al., 2021 [Koutalina]). Barbosa et al. (2022) provided a molecular phylogeny of Yphthimoides and relatives, and investigated the biogeography of the group. Possible synapomorphies and relatively distinctive characters for the clade include the following: (a) bifid distal region of the aedeagus, (b) saccus relatively short compared with other euptychiines; (c) lamella antevaginalis well developed (but somewhat variable,may be less developed in some species); (d) spiracle on the lateral plate of the 8th abdominal segment in females. Most of the species of the clade occur in Central and South America, except the Nearctic Megisto cymela and Oriental Megisto opalina. Several species of the clade (e.g. Moneuptychia, Stegosatyrus and Yphthimoides) are typical of montane natural grasslands in southern Brazil with remarkable examples including Stegosatyrus hemiclara in high elevation grasslands in the Bolivian and Peruvian Andes (Zacca et al., 2013) and Graphita griphe (Nakahara, Barbosa, et al., 2016) in the Andean cloud forests in Venezuela, Colombia, Peru and Ecuador. Adults vary from small to medium size and most species are characterized by their dull brown coloration on the dorsal and ventral wings, except for the remarkable marbled ventral pattern of some species of Carminda. Adults of both sexes are commonly attracted to traps baited with rotting fruit.Some species of Moneuptychia exhibit territorial behaviour and two or more males may make a clicking noise when they fly together, produced by the callus on the forewing subcostal vein (Freitas et al., 2015; Freitas, Rosa, et al., 2018; Murillo-Hiller, 2006). The larvae feed on various genera of Poaceae and Cyperaceae (Ackery, 1988; Beccaloni et al., 2008; DeVries, 1987; Freitas et al., 2015; Freitas, Barbosa, & Carreira, 2021; Freitas, Rosa, et al., 2018; Freitas, Rosa, & Kaminski, 2021; Janzen & Hallwachs,2022; Piovesan et al., 2022; Singer et al., 1983), and the last instars of Moneuptychia, Pharneuptychia, Yphthimoides, Carminda, and Cissia show a common body pattern with thin longitudinal zigzag stripes (Barbosa et al., 2018; Freitas et al., 2015; Freitas, 2007; Freitas, Barbosa, & Carreira, 2021; Freitas, Kaminski, et al., 2012; Freitas,Rosa,et al., 2018; Freitas pers.obs.)., 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 11, DOI: 10.1111/syen.12590, http://zenodo.org/record/7909395, {"references":["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.","Barbosa, E. P., Seraphim, N., Valencia, G., Azeredo-Espin, A. M. L. & Freitas, A. V. L. (2022) Phylogenetic systematics of Yphthimoides Forster, 1964 and related taxa, with notes on the biogeographical history of Yphthimoides species. Molecular Phylogenetics and Evolution, 168, 107390.","Zacca, T., Mielke, O. H. H., Pyrcz, T. W., Casagrande, M. M., Freitas, A. V. L. & Boyer, P. (2013) Stegosatyrus, a new genus of Euptychiina from the grasslands of neotropical realm (Lepidoptera: Nymphalidae: Satyrinae). Zootaxa, 3682, 331 - 350.","Barbosa, E. P., Silva, A. K., Paluch, M., Azeredo-Espin, A. M. L. & Freitas, A. V. L. (2015) Uncovering the hidden diversity of the Neotropical butterfly genus Yphthimoides Forster (Nymphalidae: Satyrinae): description of three new species based on morphological and molecular data. Organisms, Diversity and Evolution, 15, 577 - 589.","Barbosa, E., Marin, M., Giraldo, C. E., Uribe, S. & Freitas, A. (2016) Description of two new species of the Neotropical genus Yphthimoides Forster, 1964 (Lepidoptera: Nymphalidae: Satyrinae) from the ' renata clade. '. Neotropical Biodiversity, 2, 87 - 98.","Barbosa, E. P., Aguiar, T. M. C., Rosa, A. H. B., Zacca, T. & Freitas, A. V. L. (2020) A new species of butterfly (Lepidoptera: Nymphalidae) from the highlands of southeastern Brazil. Zootaxa, 4819, 128 - 142.","Freitas, A. V. L., Barbosa, E. P., Siewert, R. R., Mielke, O. H. H., Zacca, T. & Azeredo-Espin, A. M. L. (2015) Four new species of Moneuptychia (Lepidoptera: Satyrinae: Euptychiina) from Brazil. Zootaxa, 3981, 521 - 541.","Henao-Banol, E. R. & Meneses, L. H. (2018) A new species of Pharneuptychia Forster, 1964 from the Estoraques natural area, north of Santan- der, Colombia (Leepidoptera: Nymphalidae: Satyridae). Boletin Cientifico. Centro de Museos. Museo de Historia Natural Universidad de Caldas, 21, 173 - 183.","Zhang, J., Cong, Q., Shen, J., Opler, P. A. & Grishin, N. V. 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R. (1983) The Cissia confusa species-group in Costa Rica and Trinidad (Lepidoptera: Satyrinae). Zoological Journal of the Linnean Society, 79, 101 - 119.","Barbosa, E. P., Siewert, R. R., Mielke, O. H. H., Lamas, G., Willmott, K. R. & Freitas, A. V. L. (2018) Redescription of Yphthimoides patricia (Hayward, 1957), with taxonomic notes on the names Euptychia saltuensis Hayward, 1962 and Yphthimoides manasses (C. Felder & R. Felder, 1867) (Nymphalidae: Satyrinae). Zootaxa, 4422, 537 - 557.","Freitas, A. V. L. (2007) A new species of Moneuptychia Forster (Lepidoptera: Satyrinae, Euptychiina) from the highlands of southeastern Brazil. Neotropical Entomology, 36, 919 - 925."]}