Seven new taxa from the butterfly subtribe Euptychiina (Lepidoptera: Nymphalidae: Satyrinae) with revisional notes on Harjesia Forster, 1964 and Pseudeuptychia Forster, 1964

Center for SyStematiC entomology, inC., Gainesville, FL Insecta MundIA Journal of World Insect Systematics 0639 Seven new taxa from the butterfly subtribe Euptychiina (Lepidoptera: Nymphalidae: Satyrinae) with revisional notes on Harjesia Forster, 1964 and Pseudeuptychia Forster, 1964 Shinichi Nakahara McGuire Center for Lepidoptera and Biodiversity Florida Museum of Natural History University of Florida, Gainesville, FL 32611, USA Keith R. Willmott McGuire Center for Lepidoptera and Biodiversity Florida Museum of Natural History University of Florida, Gainesville, FL 32611, USA Olaf H. H. Mielke Laboratório de Estudos de Lepidoptera Neotropical, Departamento de Zoologia Universidade Federal do Paraná, Caixa postal 19020, 81.530-980 Curitiba, Paraná, Brazil Johanna Schwartz Department of Entomology and Nematology University of Florida, P. O. Box 110620 Gainesville, Florida 32611, USA Thamara Zacca Laboratório de Ecologia e Sistemática de Borboletas Universidade Estadual de Campinas Campinas, São Paulo, Brazil Marianne Espeland Arthropoda Department, Zoological Research Museum Alexander Koenig Adenauer Allee 160, 53113 Bonn, Germany Gerardo Lamas Departamento de Entomología, Museo de Historia Natural Universidad Nacional Mayor de San Marcos, Apartado 14-0434 Lima-14, Peru Date of issue: July 27, 2018 0639 0639 1–38 urn:lsid:zoobank.org:pub:D0C05AD6-4F63-48C6-8A26957AE9A920D2 2018 27 July Shinichi Nakahara, Keith R. Willmott, Olaf H. H. Mielke, Johanna Schwartz, Thamara Zacca, Marianne Espeland and Gerardo Lamas Seven new taxa from the butterfly subtribe Euptychiina (Lepidoptera: Nymphalidae: Satyrinae) with revisional notes on Harjesia Forster, 1964 and Pseudeuptychia Forster, 1964 Insecta Mundi 0639: 1–38 ZooBank Registered: urn:lsid:zoobank.org:pub:D0C05AD6-4F63-48C6-8A26-957AE9A920D2 Published in 2018 by Center for Systematic Entomology, Inc. P.O. Box 141874 Gainesville, FL 32614-1874 USA http://centerforsystematicentomology.org/ Insecta Mundi is a journal primarily devoted to insect systematics, but articles can be published on any non-marine arthropod. Topics considered for publication include systematics, taxonomy, nomenclature, checklists, faunal works, and natural history. Insecta Mundi will not consider works in the applied sciences (i.e. medical entomology, pest control research, etc.), and no longer publishes book reviews or editorials. 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Forsyth 0639: 1–38 2018 Seven new taxa from the butterfly subtribe Euptychiina (Lepidoptera: Nymphalidae: Satyrinae) with revisional notes on Harjesia Forster, 1964 and Pseudeuptychia Forster, 1964 Shinichi Nakahara McGuire Center for Lepidoptera and Biodiversity Florida Museum of Natural History University of Florida Gainesville, FL 32611, USA [email protected] Keith R. Willmott McGuire Center for Lepidoptera and Biodiversity Florida Museum of Natural History University of Florida Gainesville, FL 32611, USA Olaf H. H. Mielke Laboratório de Estudos de Lepidoptera Neotropical, Departamento de Zoologia Universidade Federal do Paraná Caixa postal 19020, 81.530-980 Curitiba, Paraná, Brazil Johanna Schwartz Department of Entomology and Nematology University of Florida P. O. Box 110620 Gainesville, Florida 32611, USA Thamara Zacca Laboratório de Ecologia e Sistemática de Borboletas Universidade Estadual de Campinas Campinas, São Paulo, Brazil Marianne Espeland Arthropoda Department, Zoological Research Museum Alexander Koenig Adenauer Allee 160, 53113 Bonn, Germany Gerardo Lamas Departamento de Entomología, Museo de Historia Natural Universidad Nacional Mayor de San Marcos, Apartado 14-0434 Lima-14, Peru Abstract. Seven new euptychiine (Lepidoptera: Nymphalidae: Satyrinae) taxa are described and named herein, namely Harjesia argentata Nakahara, Zacca and Lamas, n. sp., Orotaygetis Nakahara and Zacca, n. gen., O. surui Nakahara, Zacca and Lamas, n. sp., Euptychoides sanmarcos Nakahara and Lamas, n. sp., Pseudeuptychia cuzquenya Nakahara and Lamas, n. sp., P. languida austrina Nakahara and Lamas, n. ssp., and Godartiana astronesthes Lamas and Nakahara, n. sp. A revisional note is provided for Harjesia Forster, 1964 and Pseudeuptychia Forster, 1964, and as a result, Taygetis vrazi Kheil, 1896 is removed from Harjesia and a new taxonomic arrangement, Pseudodebis vrazi n. comb., is proposed based on both morphology and molecular data. Key words. New combination, new genus, new species, new subspecies, Satyrini, taxonomy. 2 • Insecta MundI 0639, July 2018 Nakahara et al. Introduction Despite the fact that they are a common component of butterfly communities in the Neotropics, the nymphalid subtribe Euptychiina has remained one of the poorly understood groups of Neotropical butterflies (Papilionoidea) until recently. The poor state of our knowledge regarding Euptychiina was partly due to widespread wing pattern homogeneity and the lack of comprehensive taxonomic study, contributing to the chaotic generic classification of this group. For example, only eight out of 23 euptychiine genera were recovered as monophyletic in the most comprehensive phylogenetic analysis of Euptychiina to date (Peña et al. 2010). Several of those genera have subsequently been proven, with denser taxon sampling, to require generic reorganization (e.g. Freitas et al. 2012, 2013, 2016, 2018; Nakahara et al. 2015a). Our state of knowledge regarding Euptychiina is thus improving, although further investigation is still required to better understand their taxonomy and systematics. Currently, over 400 described species and 48 genera are recognized within Euptychiina (Lamas 2004; Brévignon 2005; Peña and Lamas 2005; Peña et al. 2010, 2011; Freitas et al. 2011, 2012, 2013, 2015, 2016, 2018; Brévignon and Benmesbah, 2012; Matos-Maraví et al. 2013; Zacca et al. 2013, 2017; Nakahara et al. 2015a, 2015b, 2016; Costa et al. 2016). The Lamas (2004) checklist listed 80 undescribed species for Euptychiina, a remarkable proportion (20%) for butterflies. Moreover, several recent works (e.g. Cong and Grishin 2014; Barbosa et al. 2015; Nakahara et al. 2015b) have shown remarkable discrepancies between perceived and actual species richness of Euptychiina, which turns out to be significantly more speciose than estimated by Lamas (2004). In fact, we estimate the subtribe to contain over 500 species in 70 genera (unpubl. data). Partially in order to provide a baseline for a firmer understanding of spatial variation in species diversity of Euptychiina, many species-rich genera (e.g. Caeruleuptychia Forster, 1964, Euptychia Hübner, 1818, Taygetis Hübner, [1819], Magneuptychia Forster, 1964, Cissia Doubleday, 1848, Paryphthimoides Forster, 1964, Pareuptychia Forster, 1964, Euptychoides Forster, 1964, Hermeuptychia Forster, 1964, Zischkaia Forster, 1964, Chloreuptychia Forster, 1964, Forsterinaria Gray, 1973, Splendeuptychia Forster, 1964, Yphthimoides Forster, 1964) are the subject of ongoing taxonomic revisionary work by the authors and collaborators (see http://www. flmnh.ufl.edu/museum-voices/euptychiina/). The purpose of this paper is to describe and name undescribed euptychiine taxa that will otherwise not fall into those generic revisions. A new genus, five new species and one new subspecies are described herein, and adults, genitalia illustrations and distribution maps are provided for all species. A brief review and species accounts are provided for Harjesia Forster, 1964 and Pseudeuptychia Forster, 1964, as an updated classification for these two genera from Lamas (2004). In addition, we remove Taygetis vrazi Kheil, 1896 from Harjesia and place this taxon in Pseudodebis Forster, 1964, based on both morphology and molecular data. Materials and Methods Morphological study. External morphological characters were studied by soaking legs, labial palpi, and abdomens in hot 10% KOH solution for 5–10 minutes, dissecting, and storing in glycerine after examination. Genitalia were stained with chlorazol black in order to visualize the membranous structures. Drawings were done using a camera lucida attached to either a Leica MZ 16 stereomicroscope or a Nikon SMZ2800. Terminology for wing venation and wing pattern elements follows Nakahara et al. (2018); nomenclature of genitalia also follows Nakahara et al. (2018). However, we apply the terms “ductus ejaculatorius”, “manica” and “vesica” sensu Pierce (1914); “phallus”, “phallobase” and “aedeagus” sensu Klots (1956: 102–103) (see Fig. 3). Snodgrass (1935) apparently introduced the term “phallobase”, but since the definition of this term was vaguely defined in terms of Lepidoptera, we follow Klots (1956). The following collection codens are used throughout the text: DZUP Entomological Collection Padre Jesus Santiago Moure, Universidade Federal do Paraná, Curitiba, Brazil MGCL McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural History, Gainesville, USA Insecta MundI 0639, July 2018 • 3New taxa from the subtribe euptychiiNa MNHU Museum für Naturkunde, Leibniz-Institut für Evolutionsund Biodiversitätsforschung an der Humboldt Universität, Berlin, Germany MUSM Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Lima, Peru NHMUK Natural History Museum, London, UK NMPC National Museum of Prague, Prague, Czech Republic USNM National Museum of Natural History, Smithsonian Institution, Washington, D.C., USA ZSM Zoologische Staatssammlung München, München, Germany ZUEC Museu de Zoologia da Universidade Estadual de Campinas, Unicamp, Campinas, Brazil The following abbreviations are also used: DFW – Dorsal forewing DHW – Dorsal hindwing VFW – Ventral forewing VHW – Ventral hindwing Molecular work. DNA was extracted from leg or thoracic tissue samples that were either dried or stored in 96% ethanol. Voucher specimens are deposited at MGCL, DZUP and ZUEC. DNA extraction methods, PCR conditions and primers used follow Nakahara et al. (2015a) for cytochrome c oxidase I (COI) and Elongation factor 1-alpha (EF1-α), and Nakahara et al. (2018) for GAPDH. Additionally, a 591bp fragment of RPS5 (ribosomal protein S5) was amplified using the primers RPS5degF (5ʹ ATGGCNGARGARAAYTGGAAYGA 3ʹ) and RPS5degR (5ʹ CGGTTRGAYTTRGCAACACG 3ʹ) (Wahlberg and Wheat 2008). Sequences for additional specimens were taken from Peña et al. (2011) and Matos-Maraví et al. (2013) (Table 1). Sequences were assembled using Geneious 10 (Biomatters) and aligned using MAFFT v.7 (Katoh and Standley 2013). The genes were partitioned by codon position and partitions and models were selected using ModelFinder (Kalyaanamoorthy et al. 2017) in IQ-TREE 1.6.0 (Nguyen et al. 2015). Thereafter, 200 tree searches were performed in IQ-TREE to find the most likely tree. Support was calculated based on 2000 non-parametric bootstrap replicates. The tree was rooted with Neonympha mitchellii French, 1889 based on prior information (Matos-Maraví et al. 2013). The genetic distance was calculated using DNADIST version 3.5c. Databasing and georeferencing. Specimens and their label data were databased using Microsoft Access. The locality data were subsequently georeferenced with as much detail as possible, unless such information was already available on the label. Locality coordinates are provided in the “Examined specimens” section for each taxon. For holotype and lectotype specimens, information on the labels were written verbatim with labels separated by double forward slashes. Taxonomy Taxa are listed in phylogenetic order, following Espeland et al. (in review). Pseudodebis Forster, 1964 Pseudodebis: Forster 1964: 75. Type species. Papilio valentina Cramer 1779: 82, pl. 242, fig. A. ― by original designation. A revision of Pseudodebis is underway by SN, JS and collaborators, but due to the relevance of this particular species to Harjesia Forster, 1964, a genus discussed in this article, we decided to treat this species herein in order to contribute to the definition of Harjesia and to the forthcoming revision of Pseudodebis. Pseudodebis vrazi (Kheil, 1896), new combination (Fig. 2, 3, 16) Taygetis vrazi: Kheil 1896a: 151–152, fig. TL: Río Atabapo [Venezuela] and Rio Negro [Brazil]. D’Abrera 1988: 754–755, fig. [misidentified, represents Pseudodebis dubiosa Forster, 1964]. 4 • Insecta MundI 0639, July 2018 Nakahara et al. Table 1. GenBank accession numbers for sequences used in this study. Voucher code Genus Species Genes COI EF1a GAPDH RPS5 MGCL-LOAN-050 Forsterinaria pronophila MH592919 MH592940 MH592957 MH592974 NW127-20 Forsterinaria pronophila DQ338797 DQ338941 GQ864975 GQ865444 CP02-57 Forsterinaria pseudinornata JQ392594 JQ392723 JQ392827 JQ392932 CP14-07 Forsterinaria quantius GQ864772 GQ864866 GQ864972 GQ865442 BC-DZ-Willmott-293 Harjesia argentata MH592922 MH592927 MH592944 MH592961 CP01-13 Harjesia blanda DQ338800 DQ338945 GQ357436 GQ357565 MGCL-LOAN-267 Harjesia blanda MH592913 MH592941 MH592958 MH592975 CP23-22 Harjesia obscura JQ392610 JQ392737 JQ392839 JQ392946 KW-140718-01 Megeuptychia antonoe MH592909 MH592930 MH592947 MH592964 LEP-19590 Neonympha mitchelli MH592912 MH592937 MH592954 MH592971 KW-15-001 Orotaygetis surui MH592920 MH592932 MH592949 MH592966 LEP-14787 Parataygetis albinotata MH592915 MH592935 MH592952 MH592969 CP04-09 Parataygetis albinotata JQ392615 JQ392741 JQ392843 JQ392950 CP01-06 Posttaygetis penelea JQ392620 JQ392745 —JQ392955 KW-140708-01 Posttaygetis penelea MH592910 MH592928 MH592945 MH592962 CP22-02 Pseudodebis celia GU205874 GU205930 GU205988 GU206049 KW-140716-03 Pseudodebis celia 01 MH592911 MH592929 MH592946 MH592963 PM01-23 Pseudodebis marpessa JQ392624 JQ392747 JQ392849 JQ392957 CP22-05 Pseudodebis puritana JQ392712 ——— LEP-16938 Pseudodebis valentina MH592908 MH592936 MH592953 MH592970 CP01-94 Pseudodebis valentina JQ392630 ——— KW-15-003 Pseudodebis vrazi MH592918 ——— CP23-21 Sepona punctata JQ392607 JQ392734 JQ392838 JQ392943 MGCL-LOAN-490 Sepona punctata MH592917 MH592942 MH592959 MH592976 LEP-10084 Taygetina banghaasi MH592925 MH592933 MH592950 MH592967 KW-140719-01 Taygetina gulnare MH592914 MH592931 MH592948 MH592965 CP02-13 Taygetina gulnare JQ392613 JQ392740 JQ392842 JQ392949 MGCL-LOAN-022 Taygetina kerea MH592923 MH592939 MH592956 MH592973 PM02-04 Taygetina kerea JQ392645 JQ392763 JQ392866 JQ392976 CP22-01 Taygetina peribaea JQ392583 JQ392714 JQ392817 JQ392922 PM03-03 Taygetina weymeri JQ392708 JQ392814 JQ392918 JQ393027 MGCL-LOAN-003 Taygetina ypthima MH592924 MH592938 MH592955 MH592972 NW149-8 Taygetina ypthima GU205873 GU205929 GU205987 GU206048 BC-DZ-Willmott-015 Taygetis chiquitana MH592921 MH592926 MH592943 MH592960 PM04-12 Taygetis laches JQ392658 ——— PM01-07 Taygetis rufomarginata JQ392679 ——— LEP-10421 Taygetis virgilia 01 MH592916 MH592934 MH592951 MH592968 PM01-03 Taygetis larua JQ392667 ——— Insecta MundI 0639, July 2018 • 5New taxa from the subtribe euptychiiNa Taygetis Vrázi [sic]: Kheil 1896b: 230. Taygetis valentina vrazi: Weymer 1910: 191. Taygetis valentina var. vrazi: Gaede 1931: 434. Taygetis rectifascia: Forster 1964: 65, nota [as possible synonym of this species]. Harjesia vrazi: Lamas 1999: 142, figs. 1–2; lectotype designation; Lamas 2004: 220. Harjesia [n. sp.] Lamas MS (Nymphalidae: Satyrinae 1369); Lamas 2004: 220. Lectotype male. //LECTOTYPE ♂ Taygetis vrazi Kheil, 1896 By G. LAMAS, ‘99// atabapo [underlined] Tayg. vrázi // Mus. Nat. Pragae Inv. P5p/493/24// Genitalic vial SN-17-21 S. Nakahara// (NMPC) [examined]. Redescription. Male. Forewing length: 28mm (n = 2). Head: Eyes with hair-like setae, with whitish scales at base; frons brownish, covered with whitish scales and hair-like scales; post-genal area with greyish hair-like scales and greyish scales; first segment of labial palpi adorned with white long hair-like scales and brown long hair-like scales ventrally, whitish scales laterally and dorsally, second segment in length almost twice as great as eye depth and covered with whitish scales and hair-like scales laterally, dorsally adorned with whitish hair-like scales, ventrally adorned with brown long hair-like scales and long white hair-like scales 3–4× as long as segment width, third segment about one-third of second segment in length and covered with brownish scales, with band of white scales laterally; antennae approximately two-fifths of forewing length, with approximately 36 antennomeres (n = 1), distal 9–10 antennomeres composing club, distal 5 antennomeres appear darker, club not prominent. Thorax: Dorsally and laterally covered with long light greyish hair-like scales and light greyish scales; ventrally adorned with long whitish hair-like scales and whitish scales. Legs: Foreleg whitish, with tarsus, tibia, and femur similar in length; midleg and hindleg with femur whitish ventrally, tibia and tarsus greyish dorsally, ocher ventrally, tarsus and tibia adorned with spines ventrally, pair of tibial spurs present at distal end of tibia. Abdomen: Eighth tergite appears as thin sclerotized band at base of eighth abdominal segment; eighth sternite appears as a single plate. Wing venation: Basal half of forewing subcostal vein swollen; base of cubitus inflated; forewing recurrent vein absent; hindwing humeral vein developed; origin of M2 nearer M1 than M3. Male genitalia (Fig. 3a, b): Tegumen subtriangular in lateral view; uncus longer than tegumen in lateral view, narrow, sparsely adorned with hair-like setae at base, slightly curved in lateral view, tapered towards terminal point; brachia reduced, appearing as somewhat “ear-like” projections at posterior margin of tegumen; combination of ventral arms of tegumen and dorsal arms of saccus almost straight, broadens in lateral view near saccus; appendices angulares developed; saccus straight, similar to uncus in length; juxta shallow “V” shaped; in lateral view, narrow apical process of valva approximately one-third of valva length, valva broadens towards terminal point, almost twice in width, distal margin appears straight in lateral view, basal two-thirds of valva appear somewhat as parallelogram in lateral view, ventral margin slightly concave, costa rectangular, distal half of valva with hair-like setae; phallus roughly straight, phallobase about one-third of phallus in length, ductus ejaculatorius visible, manica covering about half of aedeagus, vesica visible. Female. Forewing length: 29mm (n = 1). Similar to male except as follows: Foreleg whitish, foretarsus divided into five distinct tarsomeres; ground colour of both wing surface slightly lighter. Female genitalia (Fig. 3c–e): Lamella antevaginalis sclerotized, appearing as “finger-like” sclerotized plate projecting posteriorly; lateral side of 8th abdominal segment sclerotized, apparently fused to lamella antevaginalis at its anterior margin; posterior one-third of ductus bursae slightly sclerotized, origin of ductus seminalis located at anterior end of this slightly sclerotized region, anterior two-thirds of ductus bursae membranous; corpus bursae roughly oval in dorsal view, together with ductus bursae extending length of entire abdomen, with two signa located in middle, virtually extending length of corpus bursae, parallel to each other. Specimens examined. (3 ♂, 1 ♀) Brazil: Rondônia: [Jaru], Santa Cruz da Serra (Vitt, L.), 7 Aug 1985, 1 ♂ [FLMNH-MGCL-281490], (MGCL). Peru: Loreto: Río Samiria, Estación Biológica Pithecia, 6 • Insecta MundI 0639, July 2018 Nakahara et al. [5°11′S,74°42′W], 180 m, (Pacheco, V.), 7 Nov 1979, 1 ♀ (Genitalic vial SN-17-97 S. Nakahara) [MUSMLEP-102415], (MUSM); Madre de Dios: Tambopata Research Center, La Colpa, 13°09ʹS, 69°37ʹW, 250 m, (Aibar, P.), 15 Nov 1999, 1 ♂ (Genitalic vial SN-16-63 S. Nakahara) [MUSM-LEP-102426], (MUSM). Venezuela: Amazonas: Atabapo, [4°2ʹ31ʺN,67°42ʹ27ʺW], 1 ♂ (Genitalic vial SN-17-21 S. Nakahara) (NMPC). Systematic placement and diagnosis. Our molecular phylogeny (Fig. 1) recovered Pseudodebis as a monophyletic group with high support (bs = 100), including the type species, Taygetis valentina (Cramer, 1779), as sister to the remainder of the sampled Pseudodebis taxa. Two species formerly placed in Taygetomorpha Miller, 2004 (P. celia (Cramer, 1779) and P. puritana (Weeks, 1902)) are found to be part of the monophyletic Pseudodebis. This is in accordance with Matos-Maraví et al. (2013), where Taygetomorpha was synonymized under Pseudodebis. Pseudodebis vrazi n. comb. is placed as sister to P. puritana (Weeks, 1902) with a relatively high support (bs = 91). Its placement in Pseudodebis is also supported based on morphology, namely the reduction of the brachia. All other described species of Pseudodebis appear to have reduced (or almost absent) brachia, although the degree of reduction varies across species (Forster 1964; SN pers. obs.). Pseudodebis vrazi n. comb. is readily distinguished from P. puritana by the following characters (in addition to others): 1) relatively small size (forewing length of P. puritana: 38–40 mm (n = 3)); 2) VHW postdiscal band appears as a pair of very close, parallel lines in P. vrazi n. comb. (appears as a regular, single band in P. puritana); 3) ocellus in VHW cell Cu1 present with a pupil in a black area, ringed with orange in P. vrazi n. comb. (present as a creamy-white smudge in P. puritana); 4) apical process of valva broadens at terminal point (in lateral view) in P. vrazi n. comb. (rather consistent in width in P. puritana); 5) lamella antevaginalis appears as a “finger-like” sclerotized plate projecting posteriorly in P. vrazi n. comb. (appears as a broad, rather rectangular plate in P. puritana). Pseudodebis vrazi n. comb. is also a lowland species, whereas P. puritana occurs in cloud forest habitats mostly above 1300 m. Forster (1964) considered Taygetis rectifascia Weymer, 1907 to be a possible synonym of Taygetis vrazi, although these two species are clearly not conspecific. Distribution. This species is known to date from four localities scattered across the Amazon basin in Venezuela (Amazonas), Brazil (Rondônia) and Peru (Loreto and Madre de Dios). Remarks. Until the present study, P. vrazi n. comb. was a rather enigmatic taxon known only from two specimens, the male lectotype from the Río Atabapo, Venezuela, housed in NMPC, and a female specimen from the Río Samiria, Peru, in the MUSM (Lamas 1999). Kheil (1896a) described Taygetis vrazi based on the previously mentioned Venezuelan male lectotype and a paralectotype from Rio Negro, Brazil, although we were unable to locate this Brazilian specimen supposedly housed in MNHU (Lamas 1999). Despite the fact that subsequent authors (e.g. Weymer 1910; Forster 1964) provided differing taxonomic opinions about this species, it remained in Taygetis for more than 100 years. Lamas (1999) reviewed its history of classification, designated a lectotype for T. vrazi, and reported the aforementioned Peruvian female for the first time. Moreover, Lamas (1999) transferred the species from Taygetis to the genus Harjesia Forster, 1964, stating that the “male syntype held at NMPC shows conclusively that T. vrazi is a member of the genus Harjesia”, although no further justification for this taxonomic change was provided. Subsequently, Lamas (2004) retained this systematic placement, and the species remained in Harjesia until now (e.g. Warren et al. 2017). Recently, during the course of examining the euptychiine collection at MUSM, SN discovered that one worn male specimen from La Colpa, Peru, identified as “Harjesia sp. n. 4” (listed as one of the undescribed Harjesia species in Lamas (2004)) actually represents P. vrazi n. comb. based on comparison of the male genitalia of this specimen and the male lectotype. This male specimen from La Colpa, Peru, was sequenced (KW_15_003_Pseudodebis_vrazi_01; see Fig. 1) to help clarify its systematic status, and we consequently regard this species as a member of Pseudodebis Forster, 1964. Morphology also reinforces removal of the species from Harjesia since the type species of that genus, H. blanda (Möschler, 1877), does have a rather developed brachia unlike P. vrazi n. comb. and other Pseudodebis. In addition, a male specimen from Rondônia, Brazil, in MGCL was discovered as the only known specimen of P. vrazi n. comb. from that country, increasing the number of known specimens for this species to four. Insecta MundI 0639, July 2018 • 7New taxa from the subtribe euptychiiNa Harjesia Forster, 1964 Harjesia: Forster 1964: 78. Type species. Taygetis blanda Möschler 1877: 325, pl. 3, fig. 12. ― by original designation. Forster (1964) erected Harjesia to accommodate two species, Taygetis blanda Möschler, 1877 and Euptychia obscura Butler, 1867, and designated T. blanda as the type species of the genus, even though T. blanda was considered a subspecies of Taygetis sylvia Bates, 1866 by Weymer (1910). Concerning Harjesia, Forster (1964) stated “Characteristic for this genus is mainly the male genitalia with the narrow tip of the valva”, providing no comparison with other genera. Subsequently, Lamas (2004) followed this classification and also included Euptychia oreba Butler, 1870 (regarding Euptychia gulnare Butler, 1870 and Euptychia civica Weymer, 1911 (nomen nudum) as junior subjective synonyms of this species), Euptychia griseola Weymer, 1911 and Taygetis vrazi Kheil, 1896 as members of Harjesia, in addition to proposing Euptychia eremita Weymer, 1911 and Taygetis indecisa Ribeiro, 1931 as junior subjective synonyms of E. obscura and E. gulnare, respectively. Lamas (2004) also listed four undescribed species of Harjesia, of which two (H. argentata n. sp. and Orotaygetis surui n. gen. et n. sp.) are described below. Freitas et al. (2016) recently treated E. griseola as a synonym of E. punctata Weymer, 1911 and moved this taxon out of Harjesia by placing it in its own monotypic genus, Sepona Freitas and Barbosa, 2016, based on molecular and morphological data. As discussed above, Pseudodebis vrazi n. comb. is removed from Harjesia and considered conspecific with one of the new species of Harjesia listed by Lamas (2004). Three species, namely Harjesia blanda, H. obscura and H. argentata n. sp., do form a monophyletic group with a high support (bs = 99, see Fig. 1), and we regard Harjesia as a small monophyletic genus to accommodate these three species. Finally, the remaining species in Harjesia (sensu Lamas 2004) appear not to belong in the clade containing the type for the genus (see statement above), and thus will need to be transferred to another genus (Espeland et al. unpubl. data). Systematic placement and diagnosis. Harjesia appears to be a member of a clade consisting of Forsterinaria Gray, 1973, Posttaygetis Forster, 1964, and Parataygetis Forster, 1964 (Fig. 1), but support in this part of the tree is low. Harjesia is distinguished from Forsterinaria by having a ringed ocellus with white pupil in VFW cell M1 and often in adjacent cells, whereas these ocelli appear just as white pupils or a white patch in Forsterinaria; ocelli in VHW cells M2 and M3 are ringed, and in Harjesia these are large, whereas in Forsterinaria they are small or appear just as simple white pupils; the brachia are about half the length of the uncus in Harjesia (except for H. obscura, in which the brachia are reduced), whereas the brachia are absent or reduced in Forsterinaria. Harjesia is distinguished from Posttaygetis by the absence of a horizontal creamy band in the VHW discal cell that also extends along M3, in addition to the absence of a reduced and basally placed ocellus in VHW cell M2, whereas these characters are present in Posttaygetis. As mentioned above, the brachia are about half the length of the uncus in Harjesia (except for H. obscura), whereas the brachia are absent or reduced in Posttaygetis. Among other characters, Harjesia is distinguished from Parataygetis by having a narrow, rounded apical process of the valva, whereas the distal side is broad and serrated in Parataygetis. Species account Harjesia Forster, 1964 argentata Nakahara, Zacca and Lamas, n. sp. blanda (Möschler, 1877) obscura (Butler, 1867) (= eremita Weymer, 1911) Harjesia argentata Nakahara, Zacca and Lamas, new species (Fig. 4–6, 16) Harjesia [n. sp.] Lamas, MS (Nymphalidae: Satyrinae 1367); Lamas 2004: 220. Description. Male. Forewing length: 29–30mm (n = 3). Head: Eyes sparsely hairy, with white scales at base; first segment of labial palpi similar to second segment in width, similar in length to third segment, adorned with white long hair-like scales and 14 • Insecta MundI 0639, July 2018 Nakahara et al. bursae membranous, origin of ductus seminalis located at one-fifth distance from ostium bursae to corpus bursae, ductus bursae posterior to origin of ductus seminalis apparently slightly sclerotized; corpus bursae roughly oval in dorsal view, extending to third abdominal segment, with two signa parallel to each other located posteriorly. Types. Holotype. MALE: PERU: MADRE de DIOS Cerro Pantiacolla, E slope nr. Summit, ca. 4 km. ENE Shintuya; 960–1030 m. 25.vii.1980 J. F. Douglass 1342// Allyn Museum Acc. 1980 – 14// Genitalia vial SN-14-149 S. Nakahara// (MGCL, to be deposited in MUSM). Paratypes. (2 ♂, 7 ♀) Bolivia: La Paz: Río Zongo, [16°3′40′′S,68°1′2′′W], 1200 m, (Garlepp), 1895-1896, 1 ♂, (MNHU). Peru: Cuzco: Cosñipata Valley, Quebrada Quitacalzón, [13°01′13″S,71°29′50″W], 1050 m, (Harris, B.), 12 Aug 2009, 1 ♀ (Genitalic vial SN-14-18, MUSM) ([MUSM-LEP-103661], (MUSM); (Kinyon, S.), 25 Sep 2011, 1♀ [MUSM-LEP-103662], (MUSM); Quebrada Quitacalzón, [13°1′S,71°30′W], 1100 m, (Gibson, L.), 10 May 2012, 1 ♀ [MUSM-LEP-103663], (MUSM); (Kinyon, S.), 22 Sep 2014, 1 ♀ [MUSMLEP-103666], (MUSM); (Lamas, G.), 22 Sep 2014, 1 ♀ (Genitalic vial SN-16-17) [MUSM-LEP-103664], (MUSM); (Lamas, G.), 23 Oct 2010, 1 ♂ [MUSM-LEP-103667], (MUSM); Huánuco: Cordillera del Sira, ca. [9°25′S,74°45′W], 1380 m, (Exp. Universidad Viena), Sep 1987–Aug 1988, 1 ♀ [MUSM-LEP-103660], (MUSM); Puno: Tambopata - Candamo, Río Távara, [13°25–27′S,69°38′W], 1000 m, (Baynes, H.), 1 Aug 1995, Expedition Cambridge Peru ’95, 1 ♂ [MUSM-LEP-103659], (MUSM); (Grados, J.), 8 Aug 1995, 1 ♀ [MUSM-LEP-103665], (MUSM). Etymology. This species is named after the Peruvian institution Universidad Nacional Mayor de San Marcos, the oldest university on the American continent. The majority of the type series are deposited in the museum (MUSM) belonging to this university. The specific epithet is treated as a latinized masculine noun in apposition. Distribution. This species is known from the eastern foothills of the central Peruvian Andes to Bolivia, where it is likely sympatric with E. fida. Systematic placement and diagnosis. This species is placed as sister to Ecuadorian specimens phenotypically similar to Euptychoides fida (Weymer, 1911) based on molecular data (unpublished). Although there are some slight wing pattern differences between Ecuadorian specimens and the syntypes of E. fida from Bolivia, they appear to be conspecific or at least sister taxa, and we thus describe this new species provisionally in Euptychoides. Euptychoides sanmarcos n. sp. is distinguished from syntypes of Euptychoides fida in MNHU (two males and one female, examined) by having a relatively straight VHW postdiscal band and lacking the whitish coloration distal to VHW postdiscal band. The VHW postdiscal band appears somewhat jagged in syntypes of E. fida, and there is a whitish coloration distal of this band, which also extends along the VHW inner margin. In addition, the VFW postdiscal band seems to terminate close to 2A in E. sanmarcos n. sp., whereas this band terminates half way between Cu2 and 2A in those syntypes of E. fida. Based on the female genitalia of one of the syntypes of E. fida dissected by Lee D. Miller (genitalia vial M-9151), the female genitalia of E. fida and this species appear to be similar. The genitalia vial number (M-9051) associated with this female syntype does not correspond to the genitalia of this E. fida, and the correct vial (M-9151) was determined by the genitalia dissection list prepared by Lee D. Miller, with Jacqueline Y. Miller’s assistance. It should also be noted that this female syntype has been erroneously identified as a male and is indicated as such on labels pinned with the specimen. Pseudeuptychia Forster, 1964 Pseudeuptychia: Forster 1964: 86. Type species: Euptychia languida Butler, 1871: 282 ― by original designation. Forster (1964) described Pseudeuptychia to contain two species, namely Euptychia languida Butler, 1871 and E. hemileuca Staudinger, [1886], and designated the former as the type species of the genus. Pseudeuptychia was regarded as a somewhat ‘isolated and distinctive genus’ by Forster, in regard to its DHW black ocelli and distinctive male genitalia. Presumably based on its overall appearance, Pseudeuptychia was associated with Euptychia Hübner, 1818 and Pareuptychia Forster, 1964 in the Insecta MundI 0639, July 2018 • 15New taxa from the subtribe euptychiiNa original description of the genus, but the genitalia do not show any strong affinities to support a close relationship to these two genera, and our molecular data show that these three genera are not closely related (unpublished data). Lamas (2004) retained Forster’s (1964) classification and listed two additional undescribed species in the genus. We here describe one of them, in addition to describing a new subspecies of Pseudeuptychia languida. In addition to describing these two new taxa, we also figure the female genitalia of P. hemileuca (Fig. 13a, b), a poorly known Pseudeuptychia species known only from the single female syntype in the MNHU (Fig. 12a). The new species described and named below, Pseudeuptychia cuzquenya n. sp., is also known only from a single female specimen, and clearly a discovery of the male, as well as any data regarding these taxa, would be extremely valuable in improving our understanding of these rare Neotropical butterflies. Systematic placement and diagnosis. Our molecular data suggest that Pseudeuptychia is sister to Chloreuptychia herseis (Godart, [1824]) (unpublished data), which affinity is also supported by the form of lamella antevaginalis, but a comprehensive phylogenetic study of Euptychiina, to confirm the most appropriate generic classification, is still underway. Pseudeuptychia is distinguished from Chloreuptychia herseis by having a whitish coloration on the dorsal wing surface(s), which is absent in C. herseis, in addition to having the ocelli in VHW cells M2 and M3 more rounded (the ocelli in VHW cells M2 and M3 are elongated in C. herseis). Species account Pseudeuptychia Forster, 1964 cuzquenya Nakahara and Lamas, n. sp. hemileuca (Staudinger, [1886]) languida languida (Butler, 1871) languida austrina Nakahara and Lamas, n. ssp. languida [undescribed ssp.] Nakahara et al., MS Pseudeuptychia cuzquenya Nakahara and Lamas, new species (Fig. 12b, 13, 16) Pseudeuptychia [n. sp.] Lamas MS (Nymphalidae: Satyrinae 1475): Lamas 2004: 221. Description. Male. unknown or unrecognized. Female. Forewing length: 21 mm (n = 1) Head: Eyes naked, with white scales at base; first segment of labial palpi white, adorned with white long hair-like scales and brownish long hair-like scales, second segment length almost twice as great as eye depth and covered with white hair-like scales and white scales laterally, and with black hair-like scales along edge of distal two-thirds of dorsal surface, ventrally adorned with black hair-like scales and some white hair-like scales about 3-4× as long as segment width, third segment about one-third of second segment in length and covered with black scales dorsally and ventrally, with creamy-white scales laterally; antennae approximately two-fifths of forewing length, with ca. 35 antennomeres (n = 1), distal 13–14 antennomeres composing club. Thorax: Dorsally scattered with white scales and brown scales, greenish long hair-like scales present anteriorly, mesoand metathorax covered with sparse white hair-like scales; ventrally scattered with white scales and brown scales. Legs: Foreleg whitish, first, second and third segments appear as distinct tarsomeres, fourth and fifth tarsomeres fused; midleg with femur white ventrally, tibia and tarsus greyish dorsally, tarsus ventrally ocher and tibia adorned with spines ventrally, pair of tibial spurs present at distal end of tibia. Abdomen: See genitalia description below. Wing venation: Basal half of forewing subcostal vein swollen; base of cubitus swollen; forewing recurrent vein absent; hindwing humeral vein developed; origin of M2 slightly nearer M1 than M3. Wing shape: Forewing triangular, apex rounded, costal margin slightly convex, outer margin almost straight, inner margin almost straight, but rounded towards thorax near base; hindwing slightly 16 • Insecta MundI 0639, July 2018 Nakahara et al. elongate, rounded, costal margin almost straight, angles inwards at base, outer margin convex, inner margin slightly concave near tornus, anal lobe convex, slightly round. DFW: Ground colour light brown, distally darker, white coloration extending from base towards half way through discal cell, cells Cu1, Cu2, and 2A, slightly translucent, thus subtly revealing ventral ocelli in cell M1. DHW: Ground colour white, similar to white coloration on DFW, marginal area light brown, slightly translucent, thus subtly revealing ventral submarginal and marginal bands and ocelli in cells M1 and Cu1, distal side of ocelli marked with rufous tint. VFW: Ground colour pale brown; pale reddish-brown discal band extends from radial vein, crossing discal cell, narrower below cubital vein, bent inwards and fading away before reaching 2A; whitish coloration along inner margin, extending up to origin of Cu; pale reddish-brown postdiscal band extending from radial vein towards inner margin until reaching vein 2A, slightly narrowing towards posterior end, bent outwards below Cu2; broad, faint, indistinct dark shading covering middle of cells R5, M1, M2, M3, Cu1, and Cu2 (around submarginal ocelli); sinuate submarginal band, almost concolorous, extending from apex towards tornus, jagged above Cu1, straight below this vein; concolorous marginal band, traversing along marginal area from apex to tornus; fringe brownish; ocellus in cell M1, spilling out from veins M1 and M2, black with two slate grey pupils in center, ringed with orange, tiny satellite ocellus present in cell M2, appears as slate grey patch surrounded by indistinct orange ring, small ocellus in cell M3, similar to ocellus in cell M1. VHW: Ground colour similar to forewing; general wing pattern similar to forewing except as follows: discal and post discal band broader, postdiscal band inflated distally when traversing along discal cell; submarginal band jagged after Cu1 and broadens towards tornus; five submarginal ocelli, those in cells M1 and Cu1 similar to that in VFW cell M1, those in cells M2 and M3 similar to those in VFW cells M2 and M3, ocellus in cell Rs black with slate grey smudge-like pupil in center, ringed with orange. Female genitalia (Fig. 13c): Lamella antevaginalis appears as a slightly sclerotized region surrounded by a membraneous area, somewhat semi-circular in ventral view; lateral sclerotized plate of 8th abdominal segment not fused with lamella antevaginalis at anterior margin; weakly sclerotized region present in ventral surface of intersegmental membrane of seventh and eighth abdominal segments; bursa copulatrix not examined due to damage in the abdomen. Types. Holotype. FEMALE: PERU, CU[ZCO] VALLE DE MARCAPATA// MUSM-LEP 104263// Photographed By K. Willmott June 2015// Genitalia vial SN-16-65 S. Nakahara// (MUSM). Etymology. The specific epithet is derived from the Spanish word ‘cuzqueña’, meaning ‘a woman from Cuzco’, in reference to the fact that this species is represented by a single female from Cuzco department. This specific epithet is treated as a latinized feminine noun in apposition. Distribution. To date, this species is known only from the type locality, Valle de Marcapata, Cuzco, Peru. Systematic placement and diagnosis. Although we do not have DNA data to justify the placement of this species in Pseudeuptychia, we describe this species in this genus due to its morphological resemblance to the other two other species placed in this genus, namely P. languida (Butler, 1871) (type species) and P. hemileuca Staudinger, [1886]. All three species share a similar lamella antevaginalis which appears as a simple rounded plate surrounded by a membranous area. The female of Pseudeuptychia cuzquenya n. sp. is distinguished from the female of P. languida by lacking the prominent white area between the VHW discal band and postdiscal band, in addition to having the postdiscal band crossing the origin of Cu1, whereas the postdiscal band is located distal to the origin of Cu1 in P. languida. The lamella antevaginalis is similar to that of P. languida, but is distinguished by being semi-circular in ventral view, whereas this structure is somewhat oval in ventral view in P. languida. The male of P. cuzquenya n. sp. is unknown. Pseudeuptychia languida austrina Nakahara and Lamas, new subspecies (Fig. 12c–d, 13d–g, 16) Description and diagnosis. The male description below is based on Bolivian specimens, since no definitive southern Peruvian males of this taxon are known to us. Insecta MundI 0639, July 2018 • 17New taxa from the subtribe euptychiiNa Male. Forewing length 22 mm (n = 1): Differs from the nominotypical subspecies in the following respects: curved portion of VHW postdiscal band in cells M2 and M3 less undulating, not reaching the ocelli in cells M2 and M3. Female. Forewing length 21–22 mm (n = 2): Differs from the nominotypical subspecies in the following respects: white area of DFW does not extend above origin of Cu1, and only slightly extends into discal cell or does not extend beyond cubital vein, this white area also does not extend beyond origin of Cu2 where the discal band is located; white area of VFW same as that of DFW; VFW discal band crossing origin of Cu2; curved portion of VHW postdiscal band in cells M2 and M3 less undulating, crossing origin of M3 (two Bolivian females possess VHW postdiscal band passing distal of origin of M3). Types. Holotype. FEMALE: //PERU: MADRE de DIOS Cerro Pantiacolla, E slope nr. summit, ca. 4 km. ENE Shintuya; 960–1030 m. 25.vii.1980 J. F. Douglass 1346// Allyn Museum Acc. 1980-19// (MGCL, to be deposited in MUSM). Paratypes. (7 ♀) Peru: Cuzco: Cosñipata Valley, Quebrada Quitacalzón, [13°1ʹS,71°30ʹW], 1050 m, (Gibson, L.), 2 Feb 2010, 1 ♀ [MUSM-LEP-104270], (MUSM); Quebrada Quitacalzón, [13°1ʹS,71°30ʹW], 1100 m, (Kinyon, S.), 10 May 2012, 1 ♀ [MUSM-LEP-104271], (MUSM); 31 Jan 2010, 1 ♀ [MUSMLEP-104267], (MUSM); (Lamas, G.), 2 Feb 2010, 1 ♀ [MUSM-LEP-104268], (MUSM); 1 ♀ (Genitalia vial SN-16-67) [MUSM-LEP-104269], (MUSM); 8 Nov 2007, 1 ♀ [MUSM-LEP-104266], (MUSM); 24 Oct 2016, 1 ♀, (MUSM). Other specimens examined. Bolivia: Cochabamba: Ipiri, Bosque Amazónico, [16°03ʹ26ʺS,66°40ʹ45ʺW], (Vidaurre, T.), 18 Sep 2004, 1 ♂ [MUSM-LEP-104265], (MUSM); Yungas, Bosque Amazónico, [16°02ʹ27ʺS,66°38ʹ54ʺW], (Vidaurre, T.), 23 Sep 2004, 1 ♂ (Genitalia vial SN-16-66 S. Nakahara) [MUSMLEP-104264], (MUSM); La Paz: Río Zongo, [16°3ʹ40ʺS,68°1ʹ2ʺW], (Garlepp), 1 ♀, (MNHU); Río Zongo, [16°3ʹ40ʺS,68°1ʹ2ʺW], 1200 m, (Garlepp), 1895–1896, 1 ♀, (MNHU). Etymology. The subspecific epithet is based on the feminine Latin adjective ‘austrina’, meaning southern, in reference to the southern distribution of this taxon compared to related taxa. Distribution. This subspecies is known to date from Peru (Cuzco and Madre de Dios) and Bolivia (La Paz and Cochabamba). Systematic placement and remarks. Based on existing museum specimens, we currently regard P. languida as likely representing three subspecies (but see discussion below): the nominate race of P. languida known from Colombia, with a large white patch on the DFW in the female and a strongly undulating VHW postdiscal band; an undescribed subspecies of P. languida from Ecuador to northcentral Peru (Huánuco), with an almost entirely dark DFW in the female and a strongly undulating VHW postdiscal band; and P. languida austrina n. ssp., from central Peru (Junín) to Bolivia (Cochabamba), with a white patch on the DFW in the female and less undulating VHW postdiscal band. However, given that we are not aware of any Ecuadorian females nor Colombian males of this species, we are unable to assess the range of the nominate race and the neighboring undescribed subspecies. Thus, here we limit the discussion to the southernmost subspecies, which is named and described herein, and wait to draw a conclusion regarding the taxonomic status of the aforementioned P. languida populations in the future. Although P. languida austrina n. ssp. was initially regarded as a distinct species by GL, based on the less undulating VHW postdiscal band, the sequenced specimen (MUSM-LEP-104271; DNA voucher: KW-15-034) grouped with Ecuadorian P. languida (DNA voucher: LEP-10517) with trivial COI sequence difference (0.0018) (unpubl. data). In addition, the female and putative male genitalia (Fig. 13d–g) of this subspecies are identical to those of specimens from further north in Peru (Huánuco), with similar VHW postdiscal bands to the nominate subspecies. These facts influenced our decision regarding the taxonomic status of this taxon. However, one of the known Bolivian females from Río Zongo (in MNHU, with the label “Euptychia languida Btl. det. T.G. Howarth. 1961.”) exhibits slight phenotypic differences in comparison with Peruvian females, especially in terms of the position of VHW postdiscal band, which reaches the ocelli in cells M2 and M3. The other female from the same site is similar to southern Peruvian females in that the VHW postdiscal band passes an area just distal of the origin of M3. The phenotype of 18 • Insecta MundI 0639, July 2018 Nakahara et al. the former specimen corresponds with two Bolivian males in the MUSM from Cochabamba. However, given that males and females of Bolivian specimens are not from the same site, combined with the lack of known southern Peruvian male specimens, we are unable to confidently evaluate differences between the Peruvian population and the Bolivian population, and also the phenotypic stability of the Bolivian population. Therefore, we restricted the type series to southern Peru, but given the small phenotypic difference, which could simply be broadly clinal, we currently consider the Bolivian specimens to represent the same taxon as that found in southern Peru. It should also be noted that two male specimens figured in D’Abrera (1988: 759) from the NHMUK, one from Chanchamayo, Peru and the other from Bolivia, , possess a white patch on the DFW, in contrast to the Bolivian males in the MUSM. Based on the less undulating VHW postdiscal band, we tentatively identify both specimens figured by D’Abrera (1988) as also representing P. l. austrina n. ssp. It remains to be determined whether the presence of white on the DFW is variable within or between populations. Clearly, a number of taxonomic questions remain in this genus, which can only be answered once new material and DNA sequence data become available. Godartiana Forster, 1964 Godartiana Forster 1964: 118, nota. Type species. Satyrus byses Godart, [1824]: 466 ― by original designation. Zacca et al. (2017) recently revised the classification of Godartiana, recognising five species (G. byses (Godart, [1824]), G. muscosa (Butler, 1870), G. amadoi Paluch, Zacca and Freitas, 2017, G. armilla (Butler, 1867) and G. luederwaldti (Spitz, 1931)) in the genus. An additional, undescribed species of Godartiana, in MUSM, is described and named here, increasing the number of species in the genus to six. Godartiana astronesthes Lamas and Nakahara, new species (Fig. 14–16) Praefaunula sp. n.: Lamas 1994: 180. Praefaunula [n. sp.] Lamas MS (Nymphalidae: Satyrinae 1471): Lamas 2004: 221. Description. Male. Forewing length: 14.0–14.5 mm (n = 3) Head: Eyes naked, with golden scales at base; first segment of labial palpi about half of third segment in length, whitish, adorned with golden long hair-like scales, second segment length almost twice as great as eye depth and covered with golden scales and white scales laterally, and with somewhat brownish hair-like scales along dorsal surface, ventrally adorned with long golden hair-like scales and some white long hair-like scales, about 3-4× as long as segment width, third segment about one-third of second segment in length and covered with golden scales; antennae approximately two-fifths of forewing length, with approximately 35 antennomeres (n = 1), distal 13–14 antennomeres composing club. Thorax: Dorsally and laterally covered with golden scales and long golden hair-like scales; ventrally scattered with greyish scales. Legs: Foreleg greyish, tarsus, tibia and femur almost same in length; midleg and hindleg with femur creamy, slightly paler than tarsus and tibia, tibia and tarsus dorsally greyish, ventrally brownish, tarsus and tibia adorned with spines ventrally, tibial spurs present at distal end of tibia. Abdomen: Eighth tergite reduced, apparent only along basal margin of dorsal surface of eighth abdominal segment; eighth sternite appearing as two sclerotized regions. Wing venation: Basal half of forewing subcostal vein swollen; base of cubitus swollen; forewing recurrent vein absent; hindwing humeral vein developed; origin of M2 slightly nearer M1 than M3. Wing shape: Forewing subtriangular, apex rounded, costal margin slightly convex, outer margin convex, inner margin straight, but rounded towards thorax near base; hindwing slightly elongate, rounded, costa slightly convex, angular near base towards thorax, outer margin very slightly undulating, inner margin slightly concave near tornus, anal lobe convex, slightly round. DFW: Ground colour chocolate-brown, slightly translucent, thus subtly revealing ventral ocelli. DHW: Ground colour similar to forewing, slightly translucent, thus subtly revealing ocelli. VFW: Ground colour chestnut brown; area basal to postdiscal band mottled with dark brown patches; narrow sinuate dark-brown postdiscal band extending from radial vein towards inner margin until Insecta MundI 0639, July 2018 • 19New taxa from the subtribe euptychiiNa reaching vein 2A, curved outwards, traversing area distal to discall cell; concolorous sinuate submarginal band extending from apex towards tornus, terminating at 2A-Cu2 (apparently variable), area distal to submarginal band somewhat ocher; concolorous marginal band, extending from apex towards tornus, almost parallel to submarginal band, area distal to marginal band somewhat greyish; fringe light brownish; five submarginal ocelli in cells R5, M1, M2, M3, and Cu1 (ocellus in cell Cu1 absent in three males: MUSM-LEP 104242, 104243, 104252), those in cells M1 and M2 white-pupilled black spot ringed with orangish, remaining ocelli sometimes lack black spot and ring somewhat indistinct. VHW: Ground colour similar to forewing; general wing pattern similar to forewing, except as follows: postdiscal and submarginal bands occasionally appear fused to each other at anterior and posterior ends; six submarginal ocelli in cells Rs, M1, M2, M3, Cu1, and Cu2, those in cells M1 to Cu1 white-pupilled black spot ringed with orangish, ocelli in Rs sometimes lack black spot and ring somewhat indistinct, ocelli in Cu2 sometimes very small. Male genitalia (Fig. 15a–c): Tegumen appears somewhat semi-circular in lateral view, dorsally curved in lateral view, ventrally almost straight in lateral view; uncus about twice as long as tegumen in length, posterior half curved downwards, posterior end appears as downwardly projecting flange, sparse hairs visible on dorsal surface; brachia long, about 1.5× longer than uncus, apical point higher than uncus in lateral view, slightly curved inwards in dorsal view; combination of ventral arms from tegumen and dorsal arms from saccus curved near appendices angulares; appendices angulares present; saccus similar to uncus in terms of length; juxta present; valvae setose; bristle-like hairs present on distal side of valvae; distal margin concave, ventral margin longer than dorsal margin, ventral margin convex, dorsal margin basal to costa concave; phallus roughly straight, similar in length with tegumen plus uncus, distinction between phallobase and aedeagus unclear, cornuti absent; ventral surface of anal tube apparently not sclerotized. Female. Forewing length: 14.5 mm (n = 1). Similar to male except as follows: female foretarsus divided into five distinct tarsomeres; forewing appears somewhat broader; ground colour somewhat paler. Female genitalia (Fig. 15d, e): Lamella antevaginalis sclerotized, appearing as elongated subtriangular plate in ventral view; lateral plate of 8th abdominal segment sclerotized at posterior edge, anteriorly less sclerotized, fused with lamella antevaginalis at anterior margin; ductus bursae membranous, origin of ductus seminalis located at one-third distance from ostium bursae to corpus bursae, ductus bursae slightly sclerotized around origin of ductus seminalis; corpus bursae roughly circular in dorsal view, extending to fourth abdominal segment, with two signa located in middle, parallel to each other. Types. Holotype. MALE: PERU, MD, S.N. Pampas del Heath, Refugio Juliaca 220 m 1257/6853 14.vi.92 G. Lamas// PAMPAS// Photographed By K. Willmott June 2015// MUSM-LEP 104245// (MUSM). Paratypes. (11 ♂, 1 ♀) Peru: Madre de Dios: Río Heath, Pampas del Heath, [12°57′S,68°53′W], 220 m, (Bohórquez, I.), 25 Jun 1987, 1 ♂ [MUSM-LEP-104242], (MUSM); Refugio Juliaca, [12°57′S,68°53′W], 220 m, (Lamas, G.), 14 Jun 1992, 1 ♂ [MUSM-LEP-104244], 1 ♂ [MUSM-LEP-104246], (MUSM); 15 Jun 1992, 1 ♂ [MUSM-LEP-104249], (MUSM); 1 ♂ [MUSM-LEP-104251], (MUSM); 1 ♂ (GENITALIA # MM-13, MUSM) [MUSM-LEP-104253], (MUSM); 18 Jun 1992, 1 ♂ [MUSM-LEP-104243], (MUSM); 19 Jun 1992, 1 ♂ [MUSM-LEP-104247], (MUSM); 1 ♂ [MUSM-LEP-104248], (MUSM); 1 ♂ [MUSMLEP-104250], (MUSM); 1 ♂ (Genitalia vial SN-16-70 S. Nakahara) [MUSM-LEP-104252]; 1 ♀ (Genitalia vial SN-16-68 S. Nakahara) [MUSM-LEP-104254], (MUSM). Etymology. The specific epithet is a combination of the Greek neuter noun astron, meaning ‘star’, and the Greek feminine noun esthes, meaning ‘attire’, alluding to the ventral ocelli that are reminiscent of a starry sky, or ‘clothed with stars’. This specific epithet is treated as a latinized feminine noun in apposition. Distribution. This species is so far only known from Pampas del Heath, Madre de Dios, Peru. Systematic placement and diagnosis. Godartiana appears to be a distinctive lineage within Euptychiina, being sister to a large clade containing many species-rich genera such as Caeruleuptychia, Cissia, Taygetis, etc. (Espeland et al. in review). Godartiana astronesthes n. sp. can be confidently placed in Godartiana based on its rippled/mottled ventral wing pattern and well-developed brachia (brachia longer than uncus). Godartiana astronesthes n. sp. is similar to G. armilla but is distinguished by the 20 • Insecta MundI 0639, July 2018 Nakahara et al. combination of following characters: 1) relatively small adult size; 2) DFW androconial patch absent or indistinct; 3) presence of rather prominent ocellus in VFW cell M2, in addition to variable presence of ocellus in cell Cu1; 4) presence of prominent pupil in submarginal ocelli of VFW and VHW; 5) presence of ocher area distal to VFW and VHW submarginal band. Remarks. Zacca et al. (2017) illustrated high intraspecific variation in several Godartiana species, especially in apparent association with seasonally varying precipitation. Godartiana astronesthes n. sp. has only been sampled in June (all known specimens were collected from 14–19 June 1992, except for a single specimen from 25 June 1987), which is the driest month of the year at the type locality (Cañas and Waylen 2012). Given the absence of sampled specimens from wetter months of the year, it is impossible to assess whether seasonally induced phenotypic variation exits in this species, and if such variation is found it may well influence the diagnostic characters provided above. Acknowledgments We are extremely grateful to the following persons for their generous help and support in diverse phases during the course of preparing the manuscript: Mirna Casagrande (Curitiba, Brazil), Fernando Dias (Curitiba, Brazil), André Freitas (Campinas, Brazil), Maryzender Rodríguez (Lima, Peru), Karla Mantilla (Lima, Peru), Juan Grados (Lima, Peru), Diana Silva (Lima, Peru), Gerardo Gutiérrez (Lima, Peru), César Aguilar (Lima, Peru), Xiaodi Wang (Gainesville, USA), Stacey Huber (Gainesville, USA), Evelyn Gamboa (Lima, Peru), Jason Hall (Washington DC, USA), Lei Xiao (Gainesvile, USA), Tatiana Pomerantz (Gainesville, USA), Stephanie Tyler (Gainesville, USA), Gebreyes Kassu (Gainesville, USA), Karunakar Kallam (Gainesville, USA), Alexandra Pagac (Gainesville, USA), Wolfram Mey (Berlin, Germany), Jan Sumpich (Prague, Czech Republic), Kelly Dexter (Gainesville, USA), Brian Harris (Washington, D.C., USA), Blanca Huertas (London, UK) and Axel Hausmann (Munich, Germany). SN and KRW thank Santiago Villamarín (Quito, Ecuador), the Instituto Nacional de Biodiversidad (INABIO, Ecuador) and the Ecuadorian Ministerio del Ambiente for supporting field research in Ecuador. This research was supported in part by a National Science Foundation grant (DEB–1256742), in addition to the Florida Museum of Natural History. FLMNH Museum Associates and the University of Florida. TZ thanks FAPESP for a Post-Doc fellowship (2017/02264-6). Finally, we thank Eduardo Barbosa (Campinas, Brazil) and Jacqueline Miller (Gainesville, USA) for reviewing this manuscript prior to submission. Literature Cited Barbosa, E. P., A. K. Silva, M. Paluch, A. M. L. Azeredo-Espin, and A. V. L. Freitas. 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(3): 577–589. Brévignon, C. 2005. 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Freitas, and P. Boyer. 2013. Stegosatyrus, a new genus of Euptychiina from the grasslands of neotropical realm (Lepidoptera: Nymphalidae: Satyrinae). Zootaxa 3682(2): 331–350. Insecta MundI 0639, July 2018 • 23New taxa from the subtribe euptychiiNa Zacca, T., M. Paluch, R. R. Siewert, A. V. L. Freitas, E. P. Barbosa, O. H. H. Mielke, and M. M. Casagrande. 2017. Revision of Godartiana Forster, 1964 (Lepidoptera: Nymphalidae: Satyrinae) with the description of a new species from northeastern Brazil. Austral Entomology 56(2): 169–190. Received June 4, 2018; accepted July 2, 2018. Review editor David Plotkin. Figure 1. Phylogeny of selected taxa of the “Taygetis clade” sensu Peña et al. (2010) based on maximum likelihood with support shown as non-parametric bootstrap. Taxa discussed or described in this paper are highlighted in bold. 0.02 BC-DZ-Willmott-293 Harjesia argentata n. sp. CP23-22 Harjesia obscura CP01-13 Harjesia blanda MGCL-LOAN-267 Harjesia blanda CP01-06 Posttaygetis penelea KW-140708-01 Posttaygetis penelea 100 100 100 100 100 100 100 100 100 100 100 100 99 61 54 53 85 87 99 46 62 37 89 98 82 91 99 98 94 90 77 83 79 59 CP04-09 Parataygetis albinotata LEP-14787 Parataygetis albinotata MGCL-LOAN-050 Forsterinaria pronophila NW127-20 Forsterinaria pronophila CP02-57 Forsterinaria pseudinornata CP14-07 Forsterinaria quantius CP23-21 Sepona punctata MGCL-LOAN-490 Sepona punctata KW-140718-01 Megeuptychia antonoe LEP-19590 Neonympha mitchelli KW-15-001 Orotaygetis surui n. gen et n. sp. NW149-8 Taygetis ypthima MGCL-LOAN-003 Taygetis ypthima CP01-94 Pseudodebis valentina LEP-16938 Pseudodebis valentina PM01-23 Pseudodebis marpessa CP22-05 Pseudodebis puritana KW-15-003 Pseudodebis vrazi n. comb. KW-140716-03 Pseudodebis celia 01 CP22-02 Pseudodebis celia BC-DZ-Willmott-015 Taygetis chiquitana LEP-10422 Taygetis virgilia 01 PM01-03 Taygetis larua PM01-07 Taygetis rufomarginata PM04-12 Taygetis laches LEP-10084 Taygetina banghaasi CP02-13 Taygetina gulnare KW-140719-01 Taygetina gulnare PM03-03 Taygetina weymeri CP22-01 Taygetina peribaea PM02-04 Taygetina kerea MGCL-LOAN-022 Taygetina kerea 30 • Insecta MundI 0639, July 2018 Nakahara et al. Figure 8. Orotaygetis surui n. sp., genitalia. a) Male genitalia in lateral view. b) Phallus in lateral view. c) Female genitalia with posterior portion enlarged. d) Lamella antevaginalis in ventral view. e) Corpus bursae with focus on signa (illustrated genitalia: MUSM-LEP 102416 for male; SN-16-60 for female). Insecta MundI 0639, July 2018 • 31New taxa from the subtribe euptychiiNa Figure 9. Orotaygetis surui n. sp., wing venation. a) Male forewing and hindwing. b) Female forewing and hindwing. 32 • Insecta MundI 0639, July 2018 Nakahara et al. Figure 10. Euptychoides sanmarcos n. sp., adults. Top row, holotype male, dorsal on left, ventral on right; bottom row, paratype female (MUSM-LEP 103661), dorsal on left, ventral on right. Insecta MundI 0639, July 2018 • 33New taxa from the subtribe euptychiiNa Figure 11. Euptychoides sanmarcos n. sp., genitalia. a) Male genitalia in lateral view with juxta in posterior view on the left. b) Phallus in lateral view. c) Female genitalia with location of signa indicated on the right. d) Lamella antevaginalis in ventral view with intersegmental membrane (illustrated genitalia: SN-14-149 for male; SN-16-17 for female). 34 • Insecta MundI 0639, July 2018 Nakahara et al. Figure 12. Pseudeuptychia specimens (dorsal on left, ventral on right). a) Pseudeuptychia hemileuca syntype female. b) Pseudeuptychia cuzquenya n. sp. holotype female. c) Pseudeuptychia languida austrina n. ssp. (MUSM-LEP 104264). d) Pseudeuptychia languida austrina n. ssp. holotype female. Insecta MundI 0639, July 2018 • 35New taxa from the subtribe euptychiiNa Figure 13. Pseudeuptychia genitalia. a) Female genitalia of Pseudeuptychia hemileuca syntype. b) Lamella antevaginalis in ventral view of P. hemileuca syntype. c) Lamella antevaginalis in ventral view of P. cuzquenya n. sp. d) Male genitalia in lateral view of P. languida austrina n. ssp., with tip of uncus shown above. e) Phallus in lateral view of P. languida austrina n. ssp. f) Female genitalia of P. languida austrina n. ssp. g) Lamella antevaginalis in ventral view of P. languida austrina n. ssp. (illustrated genitalia: Lee D. Miller M-9056 for P. hemileuca; SN-16-65 for P. cuzquenya n. sp.; SN-16-66 and SN-16-67 for P. languida austrina n. ssp.). 36 • Insecta MundI 0639, July 2018 Nakahara et al. Figure 14. Godartiana astronesthes n. sp., adults. Top row, holotype male (MUSM-LEP 104245), dorsal on left, ventral on right; bottom row, paratype female (MUSM-LEP 104254), dorsal on left, ventral on right. Insecta MundI 0639, July 2018 • 37New taxa from the subtribe euptychiiNa Figure 15. Godartiana astronesthes n. sp., genitalia. a) Male genitalia in lateral view. b) Valva in lateral view. c) Phallus in lateral view. d) Female genitalia with location of signa indicated on the right. e) Lamella antevaginalis in ventral view with intersegmental membrane (illustrated genitalia: SN-16-70 for male; SN-16-68 for female). 38 • Insecta MundI 0639, July 2018 Nakahara et al. Figure 16. Distribution maps of the euptychiine taxa discussed herein.