taxonID	type	description	language	source
D33D878EFFB67462FF33FA815450F8A5.taxon	type_taxon	Type species. Euptychia similis Butler, 1867 by original designation.	en	Zacca, Thamara, Casagrande, Mirna M., Mielke, Olaf H. H., Huertas, Blanca, Espeland, Marianne, Freitas, André V. L., Willmott, Keith R., Nakahara, Shinichi, Lamas, Gerardo (2020): Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman & Salvin, 1889 (Lepidoptera: Nymphalidae Satyrinae). Zootaxa 4858 (1): 1-34, DOI: 10.11646/zootaxa.4858.1.1
D33D878EFFB67462FF33FA815450F8A5.taxon	diagnosis	Diagnosis. Among the species of Euptychiina, the dorsal wing patterns of Vareuptychia stat. rest. species most resemble those of Yphthimoides Forster, 1964 (e. g. Y. manasses (C. Felder & R. Felder, 1867), Y. renata (Stoll, 1780) and Y. affinis (Butler, 1867 )) by the developed ocellus on CuA 1 - CuA 2 on the DHW. However, the VHW of Vareuptychia species are distinct from Yphthimoides, being more similar to some species of Moneuptychia Forster, 1964 (e. g. M. montana Freitas, 2015 and M. pervagata Freitas, Siewert & Mielke, 2015) in the shape and location of the ocelli in M 2 - CuA 1. Vareuptychia species differ from the above-mentioned Moneuptychia species by having four small ocelli in R 4 + 5 - M 1 (ocellus 1), M 1 - M 2 (ocellus 2), M 2 - M 3 (ocellus 3) and M 3 - CuA 1 (ocellus 4) on the VFW (sometimes the latter is not evident in some individuals), with ocellus 3 displaced towards the median line (not displaced in Moneuptychia), vein dci concave and projected into the discal cell on the FW (Fig. 12), and the gnathos developed (absent in Moneuptychia).	en	Zacca, Thamara, Casagrande, Mirna M., Mielke, Olaf H. H., Huertas, Blanca, Espeland, Marianne, Freitas, André V. L., Willmott, Keith R., Nakahara, Shinichi, Lamas, Gerardo (2020): Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman & Salvin, 1889 (Lepidoptera: Nymphalidae Satyrinae). Zootaxa 4858 (1): 1-34, DOI: 10.11646/zootaxa.4858.1.1
D33D878EFFB67462FF33FA815450F8A5.taxon	description	Redescription. Antennae half-length of costal margin of FW, with white scales around base of each flagellomere; chaetosema brown; postgenal area with small cream scales; densely hairy eyes; labial palp about two times length of head with small dark brown scales on dorsal and ventral sides of distal segment, small cream scales in lateral view of entire palp, and mixed elongated dark brown and cream scales at mid and basal segments. Legs covered with mixed small cream and brown scales with a paired tibial spur. Wings. FW sub-triangular, costa slightly convex, apex rounded, external margin slightly convex without indentations, and inner margin straight. Ground color brown with three narrow dark brown lines (median, submarginal and marginal lines) on both DW, and a large ocellus in CuA 1 - CuA 2 with a black ocellar spot surrounded by a yellowish ocellar ring and two tiny silver-bluish pupils (some individuals can have only one pupil) on DHW. VFW with four dark brown or rufous narrow lines (median, submedian, marginal and submarginal), space between median and submarginal lines lighter with a dark umbra extending from costa to 2 A, three to four small ocelli between R-M 1, M 1 – M 2, M 2 – M 3, M 3 – CuA 1, submarginal and marginal lines well-crenulated. HW sub-oval, somewhat longer than wide, costa convex, apex rounded, external margin convex with indentations, inner margin slightly sinuous. VHW with five ocelli from Rs-CuA 2, ocelli in Rs-M 1, M 1 - M 2 and CuA 1 - CuA 2 with black ocellar spot surrounded by yellowish ocellar ring (presence of pupils variable), ocelli in M 2 - CuA 2 with lighter ocellar spot and large silver pupils inside, ocellus in M 2 - CuA 1 elliptical and displaced towards median line. Some individuals can have an additional tiny ocellus in 2 A and inner margin. Venation. FW with veins Sc and CuA strongly swollen at base, 2 A slightly swollen and sinuous at base; discal cell half-length of wing with a reduced recurrent vein and m 2 - m 3 sinuous. HW with discal cell half-length of wing, humeral vein developed and m 1 - m 2 convex. Male genitalia. Tegumen flattened in dorsal view, laterally subtriangular. Uncus robust, almost same length as tegumen, dorsally with mid-apical region ovoid and apex truncated. Gnathos sinuous laterally, half-length of uncus, with apical region hooked. Combination of ventral arms of tegumen and dorsal arms of saccus sinuous. Appendices angulares short. Anterior projection of saccus developed and cylindrical. Fultura superior absent. Fultura inferior a thin sclerotized stripe. Valva rhomboid covered by long hair-like setae latero-ventrally, and short setae at inner side, costa developed and sub-squared, dorsal margin straight, apical projection pointed and longer than apex of uncus, ventral margin slightly projected at median region. Aedeagus slightly curved upwards, short than valva, anterior region bottle-shaped, posterior region about 1.5 times longer than anterior region, distal margin dorsally truncated, distal opening ventral smaller than proximal opening. Vesica with two parallel cornutal patches (Fig. 17). Female genitalia. Eighth tergite rectangular (Fig. 24). Papilla anales somewhat oblong, covered by long hairlike setae at distal region; posterior apophysis absent or reduced. Lateral plate (probably derived from 8 th sternite) can reach or not 8 th tergite and lamella antevaginalis. Lamella antevaginalis rectangular or half-orbicular in ventral view. Lamella postvaginalis absent. Ductus bursae membranous; corpus bursae smaller than ductus bursae, with paired signa of variable location. Systematic position and discussion. Vareuptychia was erected by Forster (1964) to include Euptychia similis Butler, 1867 (the type species) and Euptychia usitata Butler, 1867. In that work, Forster provided a description comparing Vareuptychia with Argyreuptychia Forster, 1964 (currently a subjective synonym of Cissia — see Zacca et al. 2018 b) based on the wing color and shape of the male genitalia, although he did not specify which structures should be used to recognize the species of Vareuptychia. The male genitalia illustrations of V. similis and V. usitata in Forster (1964: 125, figures 145 and 146, respectively) are quite different, and the genitalia of V. usitata agrees with Cissia pompilia (C. Felder & R. Felder, 1867) supporting its synonymy with the latter name (Lamas 2004; Zacca et al. 2018 b). Lamas (2004) treated Vareuptychia as a subjective synonym of Cissia, and several authors accepted this proposal after him. However, based on morphological (Marín et al. 2017) and molecular (Murray & Prowell 2005) phylogenies, and the phylogenetic analysis reported here (Fig. 87), Vareuptychia is not closely related to the type species of Cissia, and morphological features from the wing pattern, venation, and male and female genitalia support recognition of Vareuptychia as a distinct genus. Compared to Cissia species (see Zacca et al. 2018 b), Vareuptychia species can be distinguished by hairy eyes (glabrous in Cissia), FW discal cell half length of wing (2 / 3 length of the wing in Cissia), recurrent vein present in males (absent in Cissia) and m 2 - m 3 sinuous (straight in Cissia) (Fig. 12), robust uncus (slender in Cissia), truncated apex of aedeagus (bifid apex in Cissia) (Fig. 17), vesica with two cornutal patches (cornuti absent in Cissia) (Fig. 17) and lamella antevaginalis with a wrinkled appearance and half-orbicular in ventral view (orbicular or obovate in Cissia) (Fig. 25). Miller (1968) cited the tibial spurs on the midlegs as a probable synapomorphy of Cissia. However, such tibial spurs are also found in Vareuptychia species, and also frequently found in other euptychines (e. g. Magneuptychia, Graphita Nakahara, Marín & Barbosa, 2016), although their presence is variable in some genera, such as Euptychia Hübner, 1818 (S. Nakahara, pers. obs.). The male genitalia morphology is somewhat homogeneous in Vareuptychia stat. rest. species while the female genitalia have more informative characters to distinguish both species. In the present study, Vareuptychia stat. rest. was recovered as monophyletic with high support (aLRT = 100 / BS = 100), but its sister relationship with Forsterinaria Gray, 1973 was moderately supported (aLRT = 91.3 / BS = 76). In a more comprehensive phylogenetic study including> 300 species of Euptychiina (Espeland et al., in prep.) Vareuptychia stat. rest. was recovered as sister to a clade containing some species of “ Magneuptychia ” (i. e. “ M ”. analis (Godman, 1905), “ M ”. modesta (Butler, 1867 )) that will be allocated to a new genus (Nakahara et al., in prep.). The ML analysis did not recover V. similis comb. rest. and V. themis comb. nov., as identified based on morphological characters, as reciprocally monophyletic (Fig. 87), which might suggest that the morphological characters actually represent intraspecific variation. Alternatively, this result might be an artifact of the low number of specimens used for DNA analysis, or due to incomplete lineage sorting as a result of a recent speciation event (Freeland 2006). We chose a more conservative approach and considered V. similis comb. rest. and V. themis comb. nov. as distinct species rather than as variations of a single species based on consistent differences in the wing phenotype that are not related to seasonal variation or sexual dichromatism (i. e. presence / absence of pupils in the ocellus at CuA 1 - CuA 2 in VHW), and the morphology of the female genitalia (i. e. lamella antevaginalis), in addition to the fact that these morphologically-defined species are sympatric and syntopic in some localities in Mexico and Guatemala with no intermediate phenotypes (see Examined material in Suppl. Material — S 2; E. Pfeiler pers. comm.). Furthermore, numerous authors with first-hand experience of these butterflies in the field have continued to regard them as representing two species (see references listed below under each species). Nevertheless, future studies should focus on obtaining more DNA samples of Vareuptychia species throughout their entire distributional range (Mexico to Colombia — Trinidad and Tobago), and population genetic approaches, to try to better elucidate the real diversity within the genus.	en	Zacca, Thamara, Casagrande, Mirna M., Mielke, Olaf H. H., Huertas, Blanca, Espeland, Marianne, Freitas, André V. L., Willmott, Keith R., Nakahara, Shinichi, Lamas, Gerardo (2020): Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman & Salvin, 1889 (Lepidoptera: Nymphalidae Satyrinae). Zootaxa 4858 (1): 1-34, DOI: 10.11646/zootaxa.4858.1.1
D33D878EFFBC746CFF33FF485083FE00.taxon	description	(Figs. 1 – 4, 14 – 18, 24 – 25, 28)	en	Zacca, Thamara, Casagrande, Mirna M., Mielke, Olaf H. H., Huertas, Blanca, Espeland, Marianne, Freitas, André V. L., Willmott, Keith R., Nakahara, Shinichi, Lamas, Gerardo (2020): Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman & Salvin, 1889 (Lepidoptera: Nymphalidae Satyrinae). Zootaxa 4858 (1): 1-34, DOI: 10.11646/zootaxa.4858.1.1
D33D878EFFBC746CFF33FF485083FE00.taxon	diagnosis	Diagnosis. Vareuptychia similis comb. rest. can be most easily distinguished from its congener by the absence of pupils in ocellus CuA 1 - CuA 2 on the VHW (two pupils in V. themis comb. nov.), by the shape and size of the lamella antevaginalis (smaller and rectangular in V. similis, comb. rest.) (Fig. 25, black arrow), and by the position of the paired signa (positioned dorso-laterally in V. similis comb. rest.). Male genitalia (Figs. 14 – 18). See ‘ Redescription’ section. Female genitalia (Figs. 24 – 25). In addition to the characters mentioned in the generic description, the lateral plate reaches the 8 th tergite. Lamella antevaginalis is small, wrinkled and rectangular in ventral view. The paired signa are located dorso-laterally.	en	Zacca, Thamara, Casagrande, Mirna M., Mielke, Olaf H. H., Huertas, Blanca, Espeland, Marianne, Freitas, André V. L., Willmott, Keith R., Nakahara, Shinichi, Lamas, Gerardo (2020): Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman & Salvin, 1889 (Lepidoptera: Nymphalidae Satyrinae). Zootaxa 4858 (1): 1-34, DOI: 10.11646/zootaxa.4858.1.1
D33D878EFFBC746CFF33FF485083FE00.taxon	description	Variation. Most of the examined specimens exhibited a grayish brown VW ground color, lighter between the median and marginal lines and with orangey yellow patches on the umbra. However, those yellow patches on the VFW were faded or absent in a few individuals. Ocelli size was also variable on the VHW. In general, individuals of V. similis are consistently larger (FW length: 21 – 23 mm (n = 10) than those of V. themis comb. nov.: 19 – 21 mm (n = 8).	en	Zacca, Thamara, Casagrande, Mirna M., Mielke, Olaf H. H., Huertas, Blanca, Espeland, Marianne, Freitas, André V. L., Willmott, Keith R., Nakahara, Shinichi, Lamas, Gerardo (2020): Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman & Salvin, 1889 (Lepidoptera: Nymphalidae Satyrinae). Zootaxa 4858 (1): 1-34, DOI: 10.11646/zootaxa.4858.1.1
D33D878EFFBC746CFF33FF485083FE00.taxon	biology_ecology	Ecology and distribution. This species is distributed widely from Mexico to Venezuela and Trinidad and Tobago, at altitudes up to 1700 m (Fig. 28). Two historical specimens were found at the NHMUK, labeled as having been collected at Nouveau Chantier, an agricultural camp in Saint Laurent du Maroni, French Guiana. This could be interpreted as a historical occurrence of V. similis in this country or a result of mislabeling of specimens, since intensive surveys over the last 30 years have been made in this country (M. Benmesbah, pers. comm.) with no modern record of this species. A further historical specimen in the NHMUK supposedly from Manaus, Amazonas, Brazil, is also presumed mislabeled. Several recent expeditions to that region conducted by members of ‘ Laboratório de Estudos de Lepidoptera Neotropical, Universidade Federal do Paraná, Brazil’ and ‘ Laboratório de Ecologia e Sistemática de Borboletas, Universidade Estadual de Campinas, Brazil) ’ have not recorded this species, which is otherwise unknown from anywhere in the Amazon basin. Vareuptychia similis comb. nov. is multivoltine and apparently flies all year (Figueroa-Fernández et al. 2014; Miller et al. 2012; plus examined material). It is found in dry forests in association with xerophytic shrubland, secondary vegetation, pastures and riparian areas (Harvey et al. 2005; Montero-Abril et al. 2009; Figueroa-Fernández et al. 2014). It has been recorded feeding on decaying fruits, such as graviola (Annonaceae), sapote (Sapotaceae) and mango (Anacardiaceae) (Hernández-Mejía et al. 2008). Harvey et al. (2005) reported this species flying among Hyparrhenia rufa (Ness) Stapf, Brachiaria brizantha Hochst Stapf and B. decumbens Stapf grasses (Poaceae) in Nicaragua and Costa Rica, and suggested that V. similis might use them as host plants. Apart from that report, there are no other records of the host plants or immature stages.	en	Zacca, Thamara, Casagrande, Mirna M., Mielke, Olaf H. H., Huertas, Blanca, Espeland, Marianne, Freitas, André V. L., Willmott, Keith R., Nakahara, Shinichi, Lamas, Gerardo (2020): Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman & Salvin, 1889 (Lepidoptera: Nymphalidae Satyrinae). Zootaxa 4858 (1): 1-34, DOI: 10.11646/zootaxa.4858.1.1
D33D878EFFBC746CFF33FF485083FE00.taxon	materials_examined	Type material, lectotype designation and taxonomic history. Euptychia similis Butler, 1867 was described based on at least two specimens, one from Central Valleys, Guatemala and the other from an undetermined locality in Nicaragua; Butler did not provide any illustration for this species. Three years later, Butler (1870) stated that the syntype of E. similis from Nicaragua was, in fact, a distinct species from E. similis which he named E. undina (illustrated as E. similis in Butler (1867 b, pl. 12, fig. 10 )) (see further discussion below under Vareuptychia themis comb. nov.). Two syntypes of E. similis were found in the Godman & Salvin collection at the NHMUK. The male specimen from Guatemala is herein designated as the lectotype of Euptychia similis to fix the identity of the name. This lectotype has the following labels (separated by oblique bars): / Type H. T. / Central Guatemala S & G / Centr [al]. Valleys, Guatemala, F [rederick]. D [ucane]. G [odman]. & O [sbert]. S [alvin]. / ♂ / Godman-Salvin Coll. 1904. – 1. B [iologia]. C [entrali]. A [mericana]. Lep [idoptera]. Rhop [alocera]. Euptychia similis Butl [er]. / B. M. TYPE No. Rh. 3202 Euptychia similis ♂ Butl [er]. / Type. Sp. figured / BMNH (E) 982888 /; and two other labels will be added later: / Lectotype / Lectotype Euptychia similis Butler, 1867. T. Zacca det. 2020 /. NHMUK. The other specimen will be labelled as paralectotype. Vareuptychia similis has been misidentified as Cissia themis in several collections (e. g. NHMUK, DZUP, ZMHU) and literature (Lewis 1973; DeVries 1987; Marín & Uribe 2009), as also noticed by Nakahara et al. (2012). Examined material. 308 males and 74 females (14 specimens dissected) — see Supporting Information (S 2).	en	Zacca, Thamara, Casagrande, Mirna M., Mielke, Olaf H. H., Huertas, Blanca, Espeland, Marianne, Freitas, André V. L., Willmott, Keith R., Nakahara, Shinichi, Lamas, Gerardo (2020): Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman & Salvin, 1889 (Lepidoptera: Nymphalidae Satyrinae). Zootaxa 4858 (1): 1-34, DOI: 10.11646/zootaxa.4858.1.1
D33D878EFFBE7469FF33FF485091FD21.taxon	description	(Figs 5 – 13, 19 – 23, 26 – 27, 29)	en	Zacca, Thamara, Casagrande, Mirna M., Mielke, Olaf H. H., Huertas, Blanca, Espeland, Marianne, Freitas, André V. L., Willmott, Keith R., Nakahara, Shinichi, Lamas, Gerardo (2020): Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman & Salvin, 1889 (Lepidoptera: Nymphalidae Satyrinae). Zootaxa 4858 (1): 1-34, DOI: 10.11646/zootaxa.4858.1.1
D33D878EFFBE7469FF33FF485091FD21.taxon	diagnosis	Diagnosis. Vareuptychia themis can be distinguished from its congener by the bipupilled ocellus in CuA 1 - CuA 2 and the shape of the lamella antevaginalis (Fig. 27). Male genitalia (Figs. 19 – 23). See above under ‘ Redescription’ section. Female genitalia (Figs. 26 – 27). In addition to the characters mentioned in the generic description, the lateral plate does not reach the 8 th tergite. Lamella antevaginalis is large, wrinkled and half-orbicular in ventral view. The paired signa are dorsally located.	en	Zacca, Thamara, Casagrande, Mirna M., Mielke, Olaf H. H., Huertas, Blanca, Espeland, Marianne, Freitas, André V. L., Willmott, Keith R., Nakahara, Shinichi, Lamas, Gerardo (2020): Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman & Salvin, 1889 (Lepidoptera: Nymphalidae Satyrinae). Zootaxa 4858 (1): 1-34, DOI: 10.11646/zootaxa.4858.1.1
D33D878EFFBE7469FF33FF485091FD21.taxon	description	Variation. The VW ground color varies significantly from pale to dark brown, as also observed by Nakahara et al. (2012), and it is not associated with seasonal polyphenism or geographical distribution. Some individuals have orangey yellow patches on the umbra similar to those found in V. similis. Some individuals have an additional ocellus without a pupil in 2 A and the inner margin of the VHW (Fig. 8). Examination of the male genitalia of ten individuals from different localities (i. e. Mexico, El Salvador, and Venezuela) showed some variation in the size, shape and width of the tegumen, uncus, valva and aedeagus, in contrast to the observations of Nakahara et al. (2012). Conversely, female genitalia structures were found to be more conservative (mainly the shape of the lamella antevaginalis) but the presence / absence of the posterior apophysis and the size and width of the corpus bursae were variable among some individuals.	en	Zacca, Thamara, Casagrande, Mirna M., Mielke, Olaf H. H., Huertas, Blanca, Espeland, Marianne, Freitas, André V. L., Willmott, Keith R., Nakahara, Shinichi, Lamas, Gerardo (2020): Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman & Salvin, 1889 (Lepidoptera: Nymphalidae Satyrinae). Zootaxa 4858 (1): 1-34, DOI: 10.11646/zootaxa.4858.1.1
D33D878EFFBE7469FF33FF485091FD21.taxon	biology_ecology	Ecology and distribution. This species is distributed widely from Mexico to Colombia and Venezuela, and in Trinidad and Tobago (Barcant 1970; Cock 2014; DeVries 1987; Llorente-Bousquets 1996; Luis-Martínez et al. 2004; Hernández-Mejía et al. 2008; Luis-Martinez et al. 2011; Nakahara et al. 2012; Singer & Ehrlich 1991; plus examined material), at altitudes up to 2000 m (Fig. 29). Vareuptychia themis comb. nov. is sympatric and syntopic with V. similis in several localities in Mexico, such as Jalisco, Malinalco, Oaxaca, Sonora, Veracruz and Yucatán (Llorente-Bousquets 1996; Luis-Martinez et al. 2004; Hernández-Mejía et al. 2008; Luis-Martínez et al. 2011; E. Pfeiler pers. comm.; plus examined material), Guatemala (Baja Verapaz), Honduras (San Pedro Sula) and Venezuela (Aragua). Vareuptychia themis comb. nov. is multivoltine, flying all year. It has been recorded in deciduous and semideciduous forests along forest edges, and occasionally in open areas. In Costa Rica, this species is most abundant during the rainy season (Jun – Aug), and becomes less abundant as the dry season progresses (DeVries 1987). Similar to V. similis comb. rest., V. themis comb. nov. has been recorded feeding on decaying fruits, such as graviola (Annonaceae), sapote (Sapotaceae) and mango (Anacardiaceae) (Hernández-Mejía et al. 2008). Recorded larval host plants include undetermined species of Poaceae (DeVries 1987; Santin 2004).	en	Zacca, Thamara, Casagrande, Mirna M., Mielke, Olaf H. H., Huertas, Blanca, Espeland, Marianne, Freitas, André V. L., Willmott, Keith R., Nakahara, Shinichi, Lamas, Gerardo (2020): Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman & Salvin, 1889 (Lepidoptera: Nymphalidae Satyrinae). Zootaxa 4858 (1): 1-34, DOI: 10.11646/zootaxa.4858.1.1
D33D878EFFBE7469FF33FF485091FD21.taxon	materials_examined	Type material, lectotype designation and taxonomic history. Butler (1867 a) first introduced the name Euptychia themis as a nomen nudum, since no description or illustration was provided by the author, which fails to conform to Article 13 of the International Code of Zoological Nomeclature (ICZN 1999). The name became available when Butler (1867 b) published an illustration of the species (pl. 12, fig. 13) in the supplementary material to his “ Monograph of the genus Euptychia ”, although the locality of the specimen illustrated was not provided. One male specimen that agrees with the illustration of E. themis (Butler, 1867 b, pl. 12, fig. 13) was found at NHMUK. This specimen has no locality label, although ‘ Mexico’ is handwritten (but has been erased) on one label (Fig. 10), which also states that the specimen came from Hewitson’s collection, agreeing with the information provided by Butler (1867 a). To fix the identity of the name, this male specimen is herein designated as the lectotype of E. themis; it has the following labels: / Type ♂ / Hewitson Coll. 79 – 69. Euptychia themis. Butl [er]. 1. / Euptychia themis Butler / B. M. Type No. Rh. 3201 Euptychia themis, ♂ Butl [er]. / BMNH (E) 982889 /; and two others will be added later: / Lectotype / Lectotype Euptychia themis Butler, 1867. T. Zacca det. 2020 /. NHMUK. Butler (1870) named Euptychia undina based on a single specimen (holotype) from Nicaragua (Fig. 9), being part of the type series of E. similis (see discussion under V. similis) and bearing Butler’s distinctive blue label for types. Examination of the male holotype of E. undina at the NHMUK confirmed that this specimen is, in fact, E. themis since it has two silvery pupils in ocellus CuA 1 - CuA 2 on the VHW. Examined material. 339 males and 125 females (21 specimens dissected) — see Supporting Information (S 2).	en	Zacca, Thamara, Casagrande, Mirna M., Mielke, Olaf H. H., Huertas, Blanca, Espeland, Marianne, Freitas, André V. L., Willmott, Keith R., Nakahara, Shinichi, Lamas, Gerardo (2020): Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman & Salvin, 1889 (Lepidoptera: Nymphalidae Satyrinae). Zootaxa 4858 (1): 1-34, DOI: 10.11646/zootaxa.4858.1.1
D33D878EFFB8746BFF33FC9A5067FCC4.taxon	type_taxon	Type species. Euptychia labe Butler, 1870, by present designation.	en	Zacca, Thamara, Casagrande, Mirna M., Mielke, Olaf H. H., Huertas, Blanca, Espeland, Marianne, Freitas, André V. L., Willmott, Keith R., Nakahara, Shinichi, Lamas, Gerardo (2020): Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman & Salvin, 1889 (Lepidoptera: Nymphalidae Satyrinae). Zootaxa 4858 (1): 1-34, DOI: 10.11646/zootaxa.4858.1.1
D33D878EFFB8746BFF33FC9A5067FCC4.taxon	etymology	Etymology. The generic name ‘ vanima’ means beautiful in the fictional elvish language Quenya from J. R. R. Tolkien’s Middle-Earth novels. In this case, we explicitly note that we use the name Vanima as a feminine noun in the nominative singular.	en	Zacca, Thamara, Casagrande, Mirna M., Mielke, Olaf H. H., Huertas, Blanca, Espeland, Marianne, Freitas, André V. L., Willmott, Keith R., Nakahara, Shinichi, Lamas, Gerardo (2020): Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman & Salvin, 1889 (Lepidoptera: Nymphalidae Satyrinae). Zootaxa 4858 (1): 1-34, DOI: 10.11646/zootaxa.4858.1.1
D33D878EFFB8746BFF33FC9A5067FCC4.taxon	diagnosis	Diagnosis. Among the Euptychiina genera, Vanima gen. nov. can be distinguished by the following set of characters: DHW with a large bipupilled ocellus occupying the entire width of cell CuA 1 - CuA 2 (also present in some other euptychines, e. g., ‘ Cissia’ confusa (Staudinger, 1887 )), VHW having a reddish-brown quadrate patch at the tornus formed by the expansion of the marginal line (Fig. 31) (also present in other euptychines, such as ‘ Magneuptychia’ andrei Zacca, Casagrande & Mielke, 2017 and related species), and a tiny ocellus in 2 A and inner margin in VHW (Fig. 31) (absent in a few individuals of V. labe comb. nov., and present rarely in other euptychines, e. g. Euptychia sophiae Zacca, Nakahara, Dolibaina & Dias, 2015, and E. attenboroughi Neild, Nakahara, Fratello & LeCrom, 2015). This combination of characters is unique within the larger ‘ Megisto ’ - clade of which Vanima gen. nov. is a member (Espeland et al. 2019). Among Euptychiina, the genitalia of the species of Vanima gen. nov. resembles the otherwise distantly related Omacha pax (Huertas, Lamas, Fagua & Willmott, 2016) (Huertas et al. 2016; Andrade-C. et al. 2019) but can be distinguished by the ovoid uncus in dorsal view (elliptical in O. pax), apex of valva pointed or rounded (double prongs in O. pax), aedeagus bipartite at the distal region (truncated in O. pax), papillae anales without posterior apophysis (present in O. pax) and lateral plate of the 8 th segment with a spiracle dorsally located (spiracle absent in O. pax), lamella antevaginalis absent (present in O. pax). Compared to the phylogenetically closest genera, Vanima gen. nov. is easily distinguished from Carminda Ebert & Dias, 1998 by the ocelli on the DW (absent in Carminda), elongated valva (rhomboid in Carminda) and absence of lamella antevaginalis (present in Carminda), and from Yphthimoides by the reddish-brown quadrate patch at the VHW tornus (absent in Yphthimoides) and the absence of spine-like projections in the apex of the valva (present in Yphthimoides).	en	Zacca, Thamara, Casagrande, Mirna M., Mielke, Olaf H. H., Huertas, Blanca, Espeland, Marianne, Freitas, André V. L., Willmott, Keith R., Nakahara, Shinichi, Lamas, Gerardo (2020): Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman & Salvin, 1889 (Lepidoptera: Nymphalidae Satyrinae). Zootaxa 4858 (1): 1-34, DOI: 10.11646/zootaxa.4858.1.1
D33D878EFFB8746BFF33FC9A5067FCC4.taxon	description	Description. Antennae half length of costal margin of FW, with white scales around base of each flagellomere; chaetosema brown; postgena area with small white or cream scales; labial palp about two times length of head with mixed small white and black scales at distal segment and elongated scales at mid and basal segments. Legs covered with mixed small white, brown and black scales. Wings. FW sub-triangular, costa slightly convex, apex rounded, external margin slightly convex, without indentations, and inner margin straight. DHW ground color brown with elements of VFW visible through translucence, large bipupilled ocellus between Radius and M 1 with black ocellar spot surrounded by a narrow yellow ocellar ring (except in V. lesbia comb. nov. in which this ocellus is only present on VFW). VFW with three broad straight lines (basal, submedian and median) of variable color (dark brown or rufous), two thin crenulated dark brown lines (submarginal and marginal), space between median and submarginal lines with a dark brown umbra extending from costa to 2 A or costa to M 3 - CuA 1 (in latter, with an additional yellow spot between M 3 - CuA 2), two to four ocelli between R-M 1, M 1 - M 2, M 2 - M 3, M 3 - CuA 1; one at M 1 - M 2 is always present and larger with a developed black ocellar spot and two pupils while others are smaller with faded ocellar spot and 1 – 2 large pupils. HW sub-rectangular, costa convex, apex rounded, external margin convex with slight indentations, inner margin slightly convex and curved at tornus. DHW ground color brown with elements of VHW visible through slightly translucent wings, large ocellus in CuA 1 - CuA 2 with black ocellar spot surrounded by a narrow yellow ocellar ring and 1 – 2 tiny pupils. VHW similar to VFW except for five ocelli between Rs-CuA 2 and an additional tiny ocellus in 2 A and inner margin; bipupilled ocelli in M 1 - M 2 and CuA 1 - CuA 2 are larger than others with developed black ocellar spot surrounded by yellow ocellar ring, ocelli between M 2 - CuA 1 with faded ocellar spot and two large pupils; marginal line enlarged at tornus forming a rufous sub-square. Venation. FW with vein Sc strongly swollen at base, CuA swollen and 2 A slightly swollen at base, dcs reduced, dcm concave, dci straight or slightly sinuous (Fig. 45). HW with developed humeral vein (Figs. 45 – 46). Male genitalia. Tegumen sub-triangular in lateral view, in dorsal view with a median concavity at posterior region. Uncus about twice length of tegumen, robust at posterior region turning slender from median region to apex in lateral view and dorsally elliptical. Gnathos narrow and curved upwards, almost same length as uncus. Appendices angulares of variable size. Anterior projection of saccus developed and cylindrical. Fultura superior absent. Fultura inferior a thin stripe. Valva sub-triangular covered by long hair-like setae latero-ventrally, and short setae at inner side, costae developed, and apex pointed. Aedeagus straight, shorter than valva, cylindrical, anterior region bottleshaped, posterior region about two times longer than anterior region, with bifid apex in dorsal view, distal opening ventral and almost same length as proximal opening. Vesica without cornuti. Female genitalia. Eighth tergite rectangular. Papilla anales somewhat triangular, sclerotized at posterior region and covered by long hair-like setae at distal region; posterior apophysis absent. Lateral plate (probably derived from 8 th sternite) can reach or not 8 th tergite, spiracle present. Lamellae antevaginalis and postvaginalis absent. Ductus bursae membranous and of variable size; corpus bursae membranous with paired signa dorsally or dorso-laterally. Systematic position and discussion. Vanima gen. nov. comprises three species previously treated in Cissia by Lamas (2004): V. labe comb. nov., V. lesbia comb. nov. and V. palladia comb. nov. Previous molecular (Murray & Prowell 2005; Peña et al. 2006, 2010; Wahlberg et al. 2009) and morphological (Marín et al. 2017) phylogenies did not sample any species of Vanima gen. nov. Recently, Espeland et al. (2019) included V. labe comb. nov. and V. palladia comb. nov. in a backbone phylogeny of Euptychiina based on hybrid enrichment data. According to that study, V. labe comb. nov. and V. palladia comb. nov. are distantly related to Cissia but sister to a clade that comprises some species of Yphthimoides (e. g. Y. pacta (Weymer, 1911), Y. borasta (Schaus, 1902 )) and Carminda. The results of our present study also corroborate that Vanima gen. nov. is monophyletic (aLRT = 89.8 / BS = 94) (Fig. 87). With the inclusion of V. lesbia comb. nov. in the analysis we found that this species is sister to V. labe comb. nov. + V. palladia comb. nov. Some studies have recovered Yphthimoides as polyphyletic (Marín et al. 2017; Espeland et al. 2019; Barbosa et al., in prep.) and an ongoing study will describe a new genus to contain these species (Barbosa et al., in prep.). Conversely, Carminda is monophyletic based on molecular data and the morphology of the wing pattern, venation, male and female genitalia (Dias 2011; Aguiar et al., in prep.) are quite distinct from Vanima gen. nov. Although the male genitalia morphology of the species of Vanima gen. nov. shares some similarities with the monotypic Omacha, analysis of sequence data from> 300 species of Euptychiina shows that both genera are not closely related and Omacha appears as sister of the clade that includes Vareuptychia (Espeland et al., in prep.).	en	Zacca, Thamara, Casagrande, Mirna M., Mielke, Olaf H. H., Huertas, Blanca, Espeland, Marianne, Freitas, André V. L., Willmott, Keith R., Nakahara, Shinichi, Lamas, Gerardo (2020): Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman & Salvin, 1889 (Lepidoptera: Nymphalidae Satyrinae). Zootaxa 4858 (1): 1-34, DOI: 10.11646/zootaxa.4858.1.1
D33D878EFFBB7475FF33FA425083FA4C.taxon	description	(Figs. 30 – 33, 42, 47 – 51, 62 – 63, 68, 71 – 72)	en	Zacca, Thamara, Casagrande, Mirna M., Mielke, Olaf H. H., Huertas, Blanca, Espeland, Marianne, Freitas, André V. L., Willmott, Keith R., Nakahara, Shinichi, Lamas, Gerardo (2020): Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman & Salvin, 1889 (Lepidoptera: Nymphalidae Satyrinae). Zootaxa 4858 (1): 1-34, DOI: 10.11646/zootaxa.4858.1.1
D33D878EFFBB7475FF33FA425083FA4C.taxon	diagnosis	Diagnosis. Vanima labe comb. nov. resembles V. palladia comb. nov. but can be easily distinguished by its larger size, VFW and VHW with doubled submarginal lines (Fig. 31) (simple line in V. palladia comb. nov.), VHW with tiny ocellus in M 2 - CuA 1 (Fig. 35) (larger ocellus in V. palladia comb. nov.) and tornus with the rufous quadrate spot well developed (Fig. 31). Male genitalia (Figs. 47 – 51). In addition to the characters mentioned in the generic description, the apex of the valva is serrated. Female genitalia (Figs. 62 – 63). In addition to the characters mentioned in the generic description, the lateral plate does not reach the 8 th tergite and the signa are located dorsally (Fig. 62).	en	Zacca, Thamara, Casagrande, Mirna M., Mielke, Olaf H. H., Huertas, Blanca, Espeland, Marianne, Freitas, André V. L., Willmott, Keith R., Nakahara, Shinichi, Lamas, Gerardo (2020): Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman & Salvin, 1889 (Lepidoptera: Nymphalidae Satyrinae). Zootaxa 4858 (1): 1-34, DOI: 10.11646/zootaxa.4858.1.1
D33D878EFFBB7475FF33FA425083FA4C.taxon	description	Variation. Two specimens from Guatemala, one from Zapote (voucher number BMNH (E) 1420892) and another from Sinanja (BMNH (E) 1420768), stand out by having a notably faded ocellus on the DHW and sinuous median line between M 1 - CuA 1 on the VHW. Considering that there are no differences in major wing pattern elements and genitalia, they are here treated as intraspecific variation of Vanima labe comb. nov. rather than as a distinct species. Females of V. labe comb. nov. are easily distinguished from males by the rounded FW apex and well-developed subapical ocellus in M 1 - M 2 on the DFW.	en	Zacca, Thamara, Casagrande, Mirna M., Mielke, Olaf H. H., Huertas, Blanca, Espeland, Marianne, Freitas, André V. L., Willmott, Keith R., Nakahara, Shinichi, Lamas, Gerardo (2020): Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman & Salvin, 1889 (Lepidoptera: Nymphalidae Satyrinae). Zootaxa 4858 (1): 1-34, DOI: 10.11646/zootaxa.4858.1.1
D33D878EFFBB7475FF33FA425083FA4C.taxon	biology_ecology	Ecology and distribution. This species is distributed widely from Mexico to Colombia and western Ecuador (Singer et al. 1983; DeVries 1987; plus examined material) at altitudes up to 1800 m (Fig. 68). Vanima labe comb. nov. is multivoltine, flying all year. In Costa Rica, this species has been reported as being most abundant during the dry season (DeVries 1987). It is found in primary and secondary forest, usually along riparian edges or in large light gaps in rainforest (Singer et al. 1983; DeVries 1987; Ehrlich et al. 1994; Garwood & Jaramillo 2016; Garwood et al. 2016), but also in association with disturbed areas (Singer et al. 1983). Adults (Fig. 71) feed on rotting fruits and decomposing fungi (DeVries 1987). In western Ecuador (Fig. 72), males were observed perching in small groups in small light gaps and sunflecks in the forest understory on ridge tops, from 1 – 4 m above the ground, from 09: 30 – 10: 30 hr (Willmott & Hall, unpubl. data). Recorded larval host plants include undetermined species of Paspalum L. and Ichananthus Beauv. (Poaceae) (DeVries 1987; Ackery 1988; Beccaloni et al. 2008). Descriptions of the egg, first instar and pupa are available in Singer et al. (1983).	en	Zacca, Thamara, Casagrande, Mirna M., Mielke, Olaf H. H., Huertas, Blanca, Espeland, Marianne, Freitas, André V. L., Willmott, Keith R., Nakahara, Shinichi, Lamas, Gerardo (2020): Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman & Salvin, 1889 (Lepidoptera: Nymphalidae Satyrinae). Zootaxa 4858 (1): 1-34, DOI: 10.11646/zootaxa.4858.1.1
D33D878EFFBB7475FF33FA425083FA4C.taxon	materials_examined	Type material and taxonomic history. The description of Euptychia labe Butler, 1870 was based on three specimens from the Godman & Salvin collection, two collected by Arcé in Calobre and Santa Fe (Veraguas, Panama) and the other by Hague in the Polochic Valley (Guatemala). Singer et al. (1983) designated the syntype from Santa Fe as lectotype of E. labe. Although lacking its abdomen (Fig. 42), there is no doubt that it is a female based on the well-developed subapical ocellus on the DFW, and not a male as stated by Singer et al. (1983). Forster (1964) transferred E. labe to Argyreuptychia (currently a synonym of Cissia, see Zacca et al. 2018 b). Subsequently, this species was transferred to Cissia by Singer et al. (1983), mainly based on the immature stages (coloration and number of instars). In that study, Singer et al. (1983) treated Cissia labe together with C. palladia and C. penelope (Fabricius, 1775) in the ‘ labe subgroup’ defined by some features of the larval head capsule and absence of projections or knobs in the pupa. This classification was followed by Lamas (2004), but recently Zacca et al. (2018 b) noted that the species should be removed from Cissia based on morphological (wing pattern, venation, male and female genitalia) and molecular data (nuclear and mitochondrial markers). Examined material. 81 males and 142 females (14 specimens dissected) — see Supporting Information (S 2).	en	Zacca, Thamara, Casagrande, Mirna M., Mielke, Olaf H. H., Huertas, Blanca, Espeland, Marianne, Freitas, André V. L., Willmott, Keith R., Nakahara, Shinichi, Lamas, Gerardo (2020): Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman & Salvin, 1889 (Lepidoptera: Nymphalidae Satyrinae). Zootaxa 4858 (1): 1-34, DOI: 10.11646/zootaxa.4858.1.1
D33D878EFFA47477FF33FA30505BF876.taxon	description	(Figs 34 – 37, 43, 57 – 61, 64 – 65, 69)	en	Zacca, Thamara, Casagrande, Mirna M., Mielke, Olaf H. H., Huertas, Blanca, Espeland, Marianne, Freitas, André V. L., Willmott, Keith R., Nakahara, Shinichi, Lamas, Gerardo (2020): Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman & Salvin, 1889 (Lepidoptera: Nymphalidae Satyrinae). Zootaxa 4858 (1): 1-34, DOI: 10.11646/zootaxa.4858.1.1
D33D878EFFA47477FF33FA30505BF876.taxon	diagnosis	Diagnosis. Vanima lesbia comb. nov. is easily distinguished from its congeners by the VHW having a wide yellowish-ochre spot between the median and submarginal lines from M 3 to the inner margin (Figs. 35, 37). Male genitalia (Figs. 57 – 61). In addition to the characters mentioned in the generic description, the apex of the valva is well developed and curved inwards in dorsal view, slightly serrated, and the anterior projection of the saccus is well-developed. Female genitalia (Figs. 64 – 65). In addition to the characters mentioned in the generic description, the lateral plate reaches the 8 th tergite, the ductus bursae is about two times longer than the corpus bursae and the signa are located dorso-laterally.	en	Zacca, Thamara, Casagrande, Mirna M., Mielke, Olaf H. H., Huertas, Blanca, Espeland, Marianne, Freitas, André V. L., Willmott, Keith R., Nakahara, Shinichi, Lamas, Gerardo (2020): Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman & Salvin, 1889 (Lepidoptera: Nymphalidae Satyrinae). Zootaxa 4858 (1): 1-34, DOI: 10.11646/zootaxa.4858.1.1
D33D878EFFA47477FF33FA30505BF876.taxon	description	Variation. The ocellus in 2 A and the inner margin on the VHW can vary in size, but it is always present. Females and males are very similar in size and wing patterns.	en	Zacca, Thamara, Casagrande, Mirna M., Mielke, Olaf H. H., Huertas, Blanca, Espeland, Marianne, Freitas, André V. L., Willmott, Keith R., Nakahara, Shinichi, Lamas, Gerardo (2020): Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman & Salvin, 1889 (Lepidoptera: Nymphalidae Satyrinae). Zootaxa 4858 (1): 1-34, DOI: 10.11646/zootaxa.4858.1.1
D33D878EFFA47477FF33FA30505BF876.taxon	biology_ecology	Ecology and distribution. This species is known from the tropical ombrophilous lowland forest in French Guiana (Saint-Laurent-du-Maroni and Cayenne) and the Brazilian Amazon rainforest (Amazonas and Pará) (Fig. 69). Vanima lesbia comb. nov. flies from May to October, during late morning to mid-afternoon (Brévignon 2008; plus examined material). The host plants and immature stages are unknown.	en	Zacca, Thamara, Casagrande, Mirna M., Mielke, Olaf H. H., Huertas, Blanca, Espeland, Marianne, Freitas, André V. L., Willmott, Keith R., Nakahara, Shinichi, Lamas, Gerardo (2020): Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman & Salvin, 1889 (Lepidoptera: Nymphalidae Satyrinae). Zootaxa 4858 (1): 1-34, DOI: 10.11646/zootaxa.4858.1.1
D33D878EFFA47477FF33FA30505BF876.taxon	materials_examined	Type material, lectotype designation and taxonomic history. Euptychia lesbia Staudinger, [1886] was described based on an unstated number of females collected by Dr. Paul Hahnel in Maçauari, Fonte Boa and Tefé, Amazonas, Brazil. Three female syntypes of E. lesbia were found at ZMHU from the same localities mentioned above. Although L. D. Miller added labels indicating them as lectotype and paralectotypes of E. lesbia in 1988, the lectotype designation was never published. The female syntype from Maçauari is herein designated as the lectotype of Euptychia lesbia to fix the identity of the name (Fig. 43); this specimen has the following labels: / Origin / abgebildet / Massauary Hhl. [Hahnel] / ex collect [io] Staudinger / Eu [ptychia]. lesbia Stgr. [Staudinger] / Eigentum Mus [eum]. Berlin / genitalia vial M- 9060 ♀ Lee D. Miller /; and two other labels to be added: / Lectotypus / Lectotypus Euptychia lesbia Staudinger, [1886]. T. Zacca, 2020 / ZMHU. The other two syntypes become paralectotypes and will be accordingly labelled. Weymer (1911) treated E. lesbia in the “ Batesi group ” together with E. batesii Butler, 1867, E. analis, E. thalessa Möschler, 1877, E. juani Staudinger, 1887, E. tricolor Hewitson, 1850, E. fulgora Butler, 1869, E. nortia Hewitson, 1862 and E. segesta Weymer, 1911, species currently placed in Magneuptychia and Nubila Viloria, Andrade & Henao, 2019, but actually probably representing four different genera (Espeland et al., in prep.). Forster (1964) transferred E. lesbia to Argyreuptychia. Lamas (2004) placed it in Cissia, and then Zacca et al. (2018 b) noted that it would need to be removed from the latter genus. Examined material. 8 males and 7 females (2 specimens dissected) — see Supporting Information (S 2).	en	Zacca, Thamara, Casagrande, Mirna M., Mielke, Olaf H. H., Huertas, Blanca, Espeland, Marianne, Freitas, André V. L., Willmott, Keith R., Nakahara, Shinichi, Lamas, Gerardo (2020): Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman & Salvin, 1889 (Lepidoptera: Nymphalidae Satyrinae). Zootaxa 4858 (1): 1-34, DOI: 10.11646/zootaxa.4858.1.1
D33D878EFFA77470FF33FAD25067FDEC.taxon	description	(Figs. 38 – 41, 44 – 46, 52 – 56, 66 – 67, 70, 73)	en	Zacca, Thamara, Casagrande, Mirna M., Mielke, Olaf H. H., Huertas, Blanca, Espeland, Marianne, Freitas, André V. L., Willmott, Keith R., Nakahara, Shinichi, Lamas, Gerardo (2020): Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman & Salvin, 1889 (Lepidoptera: Nymphalidae Satyrinae). Zootaxa 4858 (1): 1-34, DOI: 10.11646/zootaxa.4858.1.1
D33D878EFFA77470FF33FAD25067FDEC.taxon	diagnosis	Diagnosis. Vanima palladia comb. nov. differs from its congeners by its smaller size (FW length: 16 – 18 mm), the VFW having two developed but faded ocelli in M 2 - M 3 and M 3 - CuA 1 and by having the rufous spot at the tornus smaller and more elongate rather than quadrate (well developed and quadrate in V. labe comb. nov. and V. lesbia comb. nov.) (Figs. 39, 41). Male genitalia (Figs. 52 – 56). In addition to the characters mentioned in the generic description, the apex of valva is broad and strongly serrated. Female genitalia (Figs. 66 – 67). In addition to the characters mentioned in the generic description, the lateral plate reaches the 8 th tergite, the ductus bursae is short and the signa are located dorso-laterally.	en	Zacca, Thamara, Casagrande, Mirna M., Mielke, Olaf H. H., Huertas, Blanca, Espeland, Marianne, Freitas, André V. L., Willmott, Keith R., Nakahara, Shinichi, Lamas, Gerardo (2020): Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman & Salvin, 1889 (Lepidoptera: Nymphalidae Satyrinae). Zootaxa 4858 (1): 1-34, DOI: 10.11646/zootaxa.4858.1.1
D33D878EFFA77470FF33FAD25067FDEC.taxon	description	Variation. Females have a rounded FW (triangular in males), well developed subapical ocellus in M 1 - M 2 on the DFW (absent in males), and the VFW with the median and submarginal lines farther away from each other than they are in males. In both sexes, the ocelli in M 2 - M 3 and M 3 - CuA 1 on the VFW can be faded in some individuals, but always present.	en	Zacca, Thamara, Casagrande, Mirna M., Mielke, Olaf H. H., Huertas, Blanca, Espeland, Marianne, Freitas, André V. L., Willmott, Keith R., Nakahara, Shinichi, Lamas, Gerardo (2020): Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman & Salvin, 1889 (Lepidoptera: Nymphalidae Satyrinae). Zootaxa 4858 (1): 1-34, DOI: 10.11646/zootaxa.4858.1.1
D33D878EFFA77470FF33FAD25067FDEC.taxon	biology_ecology	Ecology and distribution. This species is distributed widely from Nicaragua to Colombia, Trinidad and Tobago, French Guiana (St-Laurent du Maroni), Brazil (Acre, Roraima, Rondônia, Amazonas, Maranhão, Pernambuco, Alagoas, Goiás and Mato Grosso), Peru (Cuzco, Loreto and Madre de Dios) and Bolivia (El Beni) (Singer et al. 1983; DeVries, 1987; Singer & Ehrlich 1993; Brévignon & Benmesbah, 2012; plus examined material) at altitudes up to 1400 m (Fig. 70). According to DeVries (1987), V. palladia comb. nov. is rare in Costa Rica, being known only from remnant forests near Atenas. The species flies throughout the year. Similar to V. labe comb. nov., the peak of abundance of V. palladia comb. nov. is during the dry season in Trinidad (Singer et al. 1983). The species occurs very locally in association with deciduous and premontane forests, along riparian edges (Fig. 73) (Singer et al. 1983; DeVries, 1987). Recorded larval host plants include undetermined species of Cyperus L. and Seleria (Cyperaceae), Ichnanthus pallens (Sw.), Lasiacis sloanei (Griseb.), Oplismenus hirtellus (L.), Panicum pilosum Sw., P. polygonatum Schrad., Paspalum conjugatum P. J. Bergius, Pasp. convexum Willd. ex Döll, Pasp. decumbens Sw., Setaria paniculifera (Steud.) E. Fourn. (= S. palmifolia (J. Koenig) Stapf and Tripsacum sp. (Poaceae) (Singer et al. 1971; Singer et al. 1983; DeVries 1987; Ackery 1988; Singer & Erhlich 1993; Beccaloni et al. 2008). Nevertheless, most of these records, and perhaps all of them, come from larvae reared in captivity, and it is not clear whether the host plant used in nature is known. Descriptions of the egg and all four instars (coloration and head capsule shape) are given by Singer et al. (1983).	en	Zacca, Thamara, Casagrande, Mirna M., Mielke, Olaf H. H., Huertas, Blanca, Espeland, Marianne, Freitas, André V. L., Willmott, Keith R., Nakahara, Shinichi, Lamas, Gerardo (2020): Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman & Salvin, 1889 (Lepidoptera: Nymphalidae Satyrinae). Zootaxa 4858 (1): 1-34, DOI: 10.11646/zootaxa.4858.1.1
D33D878EFFA77470FF33FAD25067FDEC.taxon	materials_examined	Type material, lectotype designation and taxonomic history. Euptychia palladia Butler, 1867 was described based on an unstated number of specimens collected by H. W. Bates in Tapajós, Pará, Brazil. The female lectotype of E. palladia (Fig. 44) was designated by Singer et al. (1983) and is deposited at the NHMUK. The specimen cited and illustrated as E. palladia in Weymer (1911: 200, pl. 47 b) corresponds to ‘ Cissia’ myncea (Cramer, 1780) (see discussion below), as also pointed out by Singer & Ehrlich (1993). Weymer described his specimen as having five ocelli on the VHW, which disagrees with the original description of E. palladia, which noted six ocelli on the VHW, including a reduced ocellus in 2 A and the inner margin (Butler, 1867 a: 462). Barcant (1970) also misidentified ‘ Cissia’ myncea as E. palladia. Brown et al. (2007) cited Cissia sp. nr. palladia from Quibdó, Chocó, Colombia, but it has not been possible to locate this specimen. Forster (1964) transferred E. palladia to Argyreuptychia. Subsequently, this species was transferred to Cissia (Singer et al. 1983) and placed in the ‘ labe subgroup’ together with C. labe and C. penelope. This classification was followed by Lamas (2004), but recently Zacca et al. (2018 b) suggested that the species would need to be removed from Cissia based on morphological (wing pattern, venation, male and female genitalia) and molecular data (nuclear and mitochondrial markers). Examined material. 41 males and 56 females (7 specimens dissected) — see Supporting Information (S 2).	en	Zacca, Thamara, Casagrande, Mirna M., Mielke, Olaf H. H., Huertas, Blanca, Espeland, Marianne, Freitas, André V. L., Willmott, Keith R., Nakahara, Shinichi, Lamas, Gerardo (2020): Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman & Salvin, 1889 (Lepidoptera: Nymphalidae Satyrinae). Zootaxa 4858 (1): 1-34, DOI: 10.11646/zootaxa.4858.1.1
D33D878EFFA27473FF33FF485132FF22.taxon	type_taxon	Type-species. Papilio eurytus Fabricius, 1775, by subsequent designation by Butler (1868).	en	Zacca, Thamara, Casagrande, Mirna M., Mielke, Olaf H. H., Huertas, Blanca, Espeland, Marianne, Freitas, André V. L., Willmott, Keith R., Nakahara, Shinichi, Lamas, Gerardo (2020): Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman & Salvin, 1889 (Lepidoptera: Nymphalidae Satyrinae). Zootaxa 4858 (1): 1-34, DOI: 10.11646/zootaxa.4858.1.1
D33D878EFFA2747CFF33FE995069FA7D.taxon	description	(Figs 74 – 86)	en	Zacca, Thamara, Casagrande, Mirna M., Mielke, Olaf H. H., Huertas, Blanca, Espeland, Marianne, Freitas, André V. L., Willmott, Keith R., Nakahara, Shinichi, Lamas, Gerardo (2020): Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman & Salvin, 1889 (Lepidoptera: Nymphalidae Satyrinae). Zootaxa 4858 (1): 1-34, DOI: 10.11646/zootaxa.4858.1.1
D33D878EFFA2747CFF33FE995069FA7D.taxon	diagnosis	Diagnosis. Megisto cleophes comb. nov. resembles M. cymela (Cramer, 1777) but can be easily distinguished from that species by the absence of an ocellus in CuA 1 - CuA 2 on both the DFW and VFW, and by having crenulated submarginal and marginal lines on the VW (straight in M. cymela). The wing pattern of M. cleophes comb. nov. also resembles that of Cissia rubricata (Edwards, 1871), but differs by the absence of the orange spot in the median region of the DFW and VFW, and by having the median and submarginal lines running parallel from the costal to the inner margin (the lines converge between 2 A and the inner margin in C. rubricata). Male genitalia (Figs. 79 – 83). Tegumen dorsally convex, laterally subtriangular. Uncus robust at posterior region, tapering at apex, dorsally ovoid with apex truncated. Gnathos curved upwards, about 2 / 3 length of uncus, slightly sinuous, larger at base and tapering at apex. Combination of ventral arm of tegumen and dorsal arm of saccus sinuous. Appendices angulares developed, wider at base with apex curved downwards. Anterior projection of saccus short, laterally lanceolated, and smaller than gnathos. Fultura superior absent. Fultura inferior as a broad sclerotized plate in U-shape. Valva elongated, covered by long hair-like setae latero-ventrally, and short setae at inner side; costae well-developed; dorsal margin sinuous with a dorsal projection at median region, ventral margin concave at median region; apex pointed, dorsally with wide spiny projection. Aedeagus straight, cylindrical, almost same length as valva; anterior region bottle-shape; posterior region about twice longer than anterior region with bifid apex; distal opening ventral and longer than proximal opening. Vesica without cornuti. Female genitalia (Figs. 84 – 85). Eighth tergite rectangular. Papillae anales somewhat oblong covered by long hair-like setae at distal region; posterior apophysis absent. Lamella antevaginalis subtriangular in ventral view. Ductus bursae membranous; corpus bursae membranous, longer than ductus bursae, with paired signa ventrally located.	en	Zacca, Thamara, Casagrande, Mirna M., Mielke, Olaf H. H., Huertas, Blanca, Espeland, Marianne, Freitas, André V. L., Willmott, Keith R., Nakahara, Shinichi, Lamas, Gerardo (2020): Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman & Salvin, 1889 (Lepidoptera: Nymphalidae Satyrinae). Zootaxa 4858 (1): 1-34, DOI: 10.11646/zootaxa.4858.1.1
D33D878EFFA2747CFF33FE995069FA7D.taxon	description	Variation. Females have a rounded FW (triangular in males), and the submedian and median lines are broader than in males. Males have a dark androconial patch in the median region of the DFW and no pupils in the ocelli on both the VFW and VHW (Fig. 75).	en	Zacca, Thamara, Casagrande, Mirna M., Mielke, Olaf H. H., Huertas, Blanca, Espeland, Marianne, Freitas, André V. L., Willmott, Keith R., Nakahara, Shinichi, Lamas, Gerardo (2020): Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman & Salvin, 1889 (Lepidoptera: Nymphalidae Satyrinae). Zootaxa 4858 (1): 1-34, DOI: 10.11646/zootaxa.4858.1.1
D33D878EFFA2747CFF33FE995069FA7D.taxon	biology_ecology	Ecology and distribution. This species is endemic to Mexico in the states of Guerrero, Morelos, Oaxaca, Puebla and Veracruz, at altitudes up to 1400 m (Luis-Martínez et al. 1996, 2003, 2011; Michán et al. 2005; plus examined material) (Fig. 86). Megisto cleophes comb. n. flies from March to October. The immature stages and host plants are unknown, although the related M. cymela has been documented feeding on Xyris L. (Poales, Xyridaceae) (Tietz 1972; Ferris & Brown 1981; Ackery 1988).	en	Zacca, Thamara, Casagrande, Mirna M., Mielke, Olaf H. H., Huertas, Blanca, Espeland, Marianne, Freitas, André V. L., Willmott, Keith R., Nakahara, Shinichi, Lamas, Gerardo (2020): Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman & Salvin, 1889 (Lepidoptera: Nymphalidae Satyrinae). Zootaxa 4858 (1): 1-34, DOI: 10.11646/zootaxa.4858.1.1
D33D878EFFA2747CFF33FE995069FA7D.taxon	materials_examined	Type material, lectotype designation and taxonomic history. Euptychia cleophes Godman & Salvin, 1889 was described based on at least one male and one female from Tierra Colorada and Dos Caminos, Mexico. Two female syntypes from Tierra Colorada and Dos Caminos were found at NHMUK, and one of them is herein designated as the lectotype of E. cleophes to fix the identity of the species (Fig. 74); this specimen has the following labels: / Type / Type of species / E [uptychia]. cleophes type ♀ / B. M. Type No Rh 3144 Euptychia cleophes, ♀ G [odman]. & S [alvin]. / ♀ / Dos Caminos, Guerrero, 2700 ft. [822 m], Sept [ember]., H. H. Smith. / Godman-Salvin Coll. 1904. – 1. B [iologia]. C [entrali]. A [mericana]. Lep [idoptera]. Rhop [alocera]. Euptychia cleophes, G [odman]. & S [alvin]. / Sp. figured / BMNH (E) 1267115 /; and two other labels will be added later: / Lectotypus / Lectotypus Euptychia cleophes Godman & Salvin, 1889. T. Zacca det. 2020 /. NHMUK. The female syntype from Tierra Colorada will be labeled as a paralectotype. Another female specimen found at the NHMUK, from Acahuizotla, also from Smith’s collection, is not part of the type series because it was not mentioned in the original description and is, in fact, Vareuptychia themis comb. nov. As clarified by Godman (1901: 653), what Godman & Salvin (1889) supposed was the male of E. cleophes also turned out to be V. themis comb. nov. Based on the wing pattern, Godman & Salvin (1889) and Godman (1901) suggested that this species was closely allied to ‘ Cissia’ myncea and Vanima labe comb. nov.. Later, Euptychia cleophes was transferred to Cissia by Singer et al. (1983) based on the immature stages, and this classification was followed by subsequent authors. Recently, Zacca et al. (2018 b) suggested that the species needed to be removed from Cissia, mainly based on morphological data (wing patterns, venation, male and female genitalia), since DNA sequences were not available for this species. In the present study, we place this species in Megisto based on morphological similarities with M. cymela (see illustrations in Miller 1976: 6, figs. 7 – 12), including the wing pattern, venation, male genitalia with elongated gnathos, short anterior projection of saccus and absence of cornuti and female genitalia with a developed lamella antevaginalis. Although M. cleophes comb. nov. shows some similarities in wing pattern with Cissia rubricata, these might be the result of convergent evolution; the presence of androconial scales on the DFW is also shared with M. cymela, and absent in all species of Cissia. Superficial similarity in wing pattern is also true of M. cleophes comb. nov. and Llorenteana pellonia (Godman, 1901), but wing venation and morphology of the male genitalia are helpful to distinguish both taxa. Until very recently, L. pellonia was treated as ‘ incertae sedis’ (Lamas 2004), but the species was recently transferred to the new genus Llorenteana Viloria & Luis-Martínez, 2019, in the subtribe Ypthimina (Viloria & Luis-Martínez 2019). Nevertheless, no morphological or molecular phylogenetic analysis has yet confirmed this tentative subtribal placement. Systematic position and discussion. Megisto is sister to the Oriental Palaeonympha opalina (Peña et al. 2006, 2010; Espeland et al. 2019), with the split of both genera from the remaining South American Euptychiina genera (except Euptychia) estimated to have occurred in the early Miocene (Peña et al. 2010). Unfortunately, we were unable to obtain DNA sequences for M. cleophes comb. nov. and our classification of this taxon is based only on comparison of morphological characters. Clearly, DNA sequence data for this species, as well as the enigmatic Llorenteana pellonia, would be valuable to test these taxonomic proposals and better understand the relationships of these restricted Mexican species. Examined material. 3 males and 19 females (2 specimens dissected) — see Supporting Information (S 2).	en	Zacca, Thamara, Casagrande, Mirna M., Mielke, Olaf H. H., Huertas, Blanca, Espeland, Marianne, Freitas, André V. L., Willmott, Keith R., Nakahara, Shinichi, Lamas, Gerardo (2020): Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman & Salvin, 1889 (Lepidoptera: Nymphalidae Satyrinae). Zootaxa 4858 (1): 1-34, DOI: 10.11646/zootaxa.4858.1.1
