Reassessment of the type locality of Euptychiastigmatica Godman, 1905, with the description of two new sibling species from Amazonia (Lepidoptera, Nymphalidae, Satyrinae, Satyrini).

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

Immature Stages and New Host Plant Records for Three Species in the "Taygetis Clade" of Euptychiina in Southeastern Peru (Lepidoptera: Nymphalidae: Satyrinae).

We report here the immature stages and natural host plants for three species in the so-called "Taygetis clade" of the nymphalid butterfly subtribe Euptychiina, Taygetis echo (Cramer, 1775), Taygetis sosis Hopffer, 1874, and Pseudodebis valentina (Cramer, 1779). The study was carried out at Finca Las Piedras in Madre de Dios department, Peru. Taygetis echo was observed utilizing a species of herbaceous bamboo, Olyra latifolia L. (Poaceae: Bambusoideae: Olyreae), T. sosis was found feeding on another species of herbaceous bamboo, Pariana lunata Nees (Poaceae: Bambusoideae: Olyreae), and P. valentina was found utilizing a species of woody bamboo, Guadua weberbaueri Pilg. (Poaceae: Bambusoideae: Bambuseae). We describe and provide photos of the immatures and host plants, as well as illustrations of the head capsules for each of these three species.

Revision of the "celia clade" of Pseudodebis Forster, 1964, with Two New Species and Notes on Papilio phorcys Fabricius, 1793 (Lepidoptera: Nymphalidae: Satyrinae).

The species-level classification of the "celia clade" of the nymphalid butterfly genus Pseudodebis Forster, 1964, is revised as part of ongoing revisionary work on this genus. The "celia clade" contains three species, of which two, Pseudodebis darrenthroopi Nakahara & Willmott, n. sp. and P. tigrillo Nakahara & Willmott, n. sp., are described and named herein based on morphology and molecular data. Consequently, we increase the described species diversity of Pseudodebis to 13, with a remarkable six species occurring in the trans-Andean region. We discuss five specific epithets associated with the clade and designate a neotype for Papilio celia Cramer, 1780, and lectotype for Taygetis keneza Butler, 1869, based on the same specimen, thus retaining the status of the former name as a senior objective synonym. We also provide a brief historical review for Papilio phorcys Fabricius, 1793, an enigmatic name currently synonymized under Papilio celia. Nevertheless, we were unable to locate the syntype(s) for this name and the identity of Papilio phorcys remains uncertain, so we retain the current synonymy as a parsimonious approach.

Immature stages and new host plant records for four satyrine species feeding on herbaceous bamboos in southeastern Peru (Lepidoptera: Nymphalidae: Satyrinae: Satyrini).

We here document the immature stages of three euptychiine butterflies, Nhambikuara mima (Butler, 1867), Splendeuptychia furina (Hewitson, 1862), and Paryphthimoides brixius (Godart, [1824]), all found feeding on a species of herbaceous bamboo, Taquara micrantha (Kunth) I.L.C.Oliveira R.P.Oliveira (Poaceae: Bambusoideae: Olyreae) in Madre de Dios, Peru. This study is the first to report the life history of these three taxa with their natural host plant. We provide illustrations of immatures, head capsules, and the host plant for each of these three species. The immature morphology of these taxa supports recent generic arrangements of these three species in comparison with their close relatives, namely Splendeuptychia furina to Nhambikuara mima and Paryphthimoides brixius to Paryphthimoides terrestris (Butler, 1867), a species documented in our successive study. Thus, the present study includes taxonomic implications based on immature stages by discovering putative synapomorphic characters of larvae and pupae. These pairs of closely related species occur in micro-sympatry at the study site in southeastern Peru, and our observations possibly suggest niche partitioning between sibling species. Additionally, we report two herbaceous bamboo species, Olyra latifolia L. and Taquara micrantha (Kunth) I.L.C.Oliveira R.P.Oliveira as the first known natural host plants for Magneuptychia harpyia (C. Felder R. Felder, 1867).

Resolving two centuries of mistaken identity: Reinterpretation of Papilio marcus Fabricius, 1787 (Lepidoptera: Nymphalidae, Hesperiidae).

Here, we untangle an oversight surrounding the application of the name Papilio marcus Fabricius, 1787 (Hesperiidae: Hesperiinae), currently in Troyus Warren & Turland, 2012, which has eluded taxonomists for nearly two centuries. First, we note that P. marcus is a junior primary homonym of Papilio marcus Schaller, 1785, a species currently in Morpho Fabricius, 1807 (Nymphalidae: Satyrinae). Second, we designate a lectotype for P. marcus Fabricius, 1787, and recognize it as conspecific with Papilio phyllus Cramer, 1777, a species currently in Vettius Godman, 1901. Therefore, P. marcus is also a junior subjective synonym of V. phyllus (new synonym). Third, aided by genomic sequencing of the lectotype of Vettius phyllides Röber, 1925, we find that this species is not conspecific with V. phyllus and represents instead a valid species of Troyus Warren & Turland, 2012, so that the relative epithet, currently considered as a junior subjective synonym of P. marcus, has to be reappreciated as Troyus phyllides (reinstated status, new combination). Moreover, T. phyllides is apparently the species that has been misidentified as P. marcus since 1832. As a result of this study, the name P. marcus Fabricius nec Cramer falls in synonymy with P. phyllus (currently in Vettius), and the species currently known as T. marcus becomes Troyus phyllides.

Description of a new genus and species for a common and widespread Amazonian satyrine butterfly (Lepidoptera: Nymphalidae: Satyrinae: Satyrini).

We here propose a new monotypic butterfly genus Scriptor Nakahara & Espeland, n. gen. to accommodate a new species, S. sphenophorus Lamas & Nakahara, n. sp., described and named herein. Scriptor sphenophorus n. gen. and n. sp.is a relatively common and widespread butterfly species which is recovered as a member of the so-called "Splendeuptychia clade" in the nymphalid subtribe Euptychiina, based on our molecular phylogenetic analysis using a maximum likelihood approach. Nevertheless, its sister group is not confidently resolved in any analysis, supporting a relatively distant relationship to any described genus as well as our decision to establish a new monotypic genus. We further discuss the proposed taxonomy in the light of frequent criticism of the description of monotypic taxa, as well as emphasize the importance of incorporating multiple evidence when describing new genera, illustrated by reference to several recent generic descriptions in this subtribe.

Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman amp; Salvin, 1889 (Lepidoptera: Nymphalidae: Satyrinae).

Vareuptychia Forster, 1964 stat. rest. is revalidated and comprises two species, V. similis (Butler, 1867) comb. rest. and V. themis (Butler, 1867) comb. nov. Vanima Zacca, Casagrande Mielke gen. nov. is described to contain Euptychia labe Butler, 1870 (the type species), E. palladia Butler, 1867 and E. lesbia Staudinger, [1886]. The taxonomy of these two genera was initially revised based on morphological and distributional data, and subsequently tested and supported with a Maximum Likelihood analysis using four genes (COI, GAPDH, RpS5 and EF1-a). Lectotypes are designated for Euptychia similis Butler, 1867, E. themis Butler, 1867, E. undina Butler, 1870 and E. lesbia Staudinger, [1886]. No DNA sequences were obtained for Euptychia cleophes Godman Salvin, 1889 but its transfer to Megisto Hübner, [1819] is supported by morphological evidence. For all taxa treated in this study, a taxonomic catalog, diagnosis, (re)description and illustrations of adults, venation and genitalia are provided, as well as comments on intraspecific variation, sexual dichromatism, ecology and distribution maps.

A revision of the new genus Amiga Nakahara, Willmott & Espeland, gen. n., described for Papilioarnaca Fabricius, 1776 (Lepidoptera, Nymphalidae, Satyrinae).

We here propose a new, monotypic genus, Amiga Nakahara, Willmott & Espeland, gen. n., to harbor a common Neotropical butterfly, described as Papilioarnaca Fabricius, 1776, and hitherto placed in the genus Chloreuptychia Forster, 1964. Recent and ongoing molecular phylogenetic research has shown Chloreuptychia to be polyphyletic, with C.arnaca proving to be unrelated to remaining species and not readily placed in any other described genus. Amigaarnaca gen. n. et comb. n. as treated here is a widely distributed and very common species ranging from southern Mexico to southern Brazil. A neotype is designated for the names Papilioarnaca and its junior synonym, Papilioebusa Cramer, 1780, resulting in the treatment of the latter name as a junior objective synonym of the former. A lectotype is designated for Euptychiasericeella Bates, 1865, which is treated as a subspecies, Amigaarnacasericeella (Bates, 1865), comb. n. et stat. n., based on molecular and morphological evidence. We also describe two new taxa, Amigaarnacaadela Nakahara & Espeland, ssp. n. and Amigaarnacaindianacristoi Nakahara & Marín, ssp. n., new subspecies from the western Andes and eastern Central America, and northern Venezuela, respectively.

Four hundred shades of brown: Higher level phylogeny of the problematic Euptychiina (Lepidoptera, Nymphalidae, Satyrinae) based on hybrid enrichment data.

Relationships within satyrine butterflies have been notoriously difficult to resolve using both morphology and Sanger sequencing methods, and this is particularly true for the mainly Neotropical subtribe Euptychiina, which contains about 400 described species. Known larvae of Euptychiina feed on grasses and sedges, with the exception of the genus Euptychia, which feed on mosses and lycopsids, and the butterflies occur widely in rainforest, cloudforest and grassland habitats, where they are often abundant. Several previous molecular and morphological studies have made significant progress in tackling the systematics of the group, but many relationships remain unresolved, with long-branch-attraction artifacts being a major problem. Additionally, the monophyly of the clade remains uncertain, with Euptychia possibly not being closely related to the remainder of the clade. Here we present a backbone phylogeny of the subtribe based on 106 taxa, 368 nuclear loci, and over 180,000 bps obtained through hybrid enrichment. Using both concatenation and species tree approaches (IQ-TREE, EXABAYES, ASTRAL), we can for the first time strongly confirm the monophyly of Euptychiina with Euptychia being the sister group to the remainder of the clade. The Euptychiina is divided into nine well supported clades, but the placement of a few genera such as Hermeuptychia, Pindis and the Chloreuptychia catharina group still remain uncertain. As partially indicated in previous studies, the genera Cissia, Chloreuptychia, Magneuptychia, Megisto, Splendeuptychia and Euptychoides, among others, were found to be highly polyphyletic and revisions are in preparation. The phylogeny will provide a strong backbone for the analysis of datasets in development that are much more taxonomically comprehensive but have orders of magnitude fewer loci. This study therefore represents a critical step towards resolving the higher classification and studying the evolution of this highly diverse lineage.

A new species of Cyllopsis R. Felder, 1869 from the highlands of Chiapas, Mexico (Lepidoptera: Nymphalidae: Satyrinae).

A new species of Cyllopsis R. Felder, 1869, is described and illustrated from the highlands of Chiapas, Mexico. Cyllopsis tomemmeli A. Warren Nakahara, sp. nov., is currently known from 13 specimens (9 males and 4 females) collected on March 26-28, 1959, southeast of San Cristóbal de Las Casas. Despite extensive studies on the butterfly fauna of this region, this species has not since been encountered. We discuss possible relationships between this new species and other species of Cyllopsis.

Association of MEOX2 polymorphism with nonsyndromic cleft palate only in a Vietnamese population.

To evaluate the association between the single nucleotide polymorphism (SNP) rs227493 in the MEOX2 gene and nonsyndromic cleft palate only, this research was conducted as a case-control study by comparing a nonsyndromic cleft palate only group with an independent, healthy, and unaffected control group who were both examined by specialists. Based on clinical examination and medical records, we analyzed a total of 570 DNA samples, including 277 cases and 293 controls, which were extracted from dry blood spot samples collected from both the Odonto and Maxillofacial Hospital in Ho Chi Minh City and Nguyen Dinh Chieu Hospital in Ben Tre province, respectively. The standard procedures of genotyping the specific SNP (rs2237493) for MEOX2 were performed on a StepOne Realtime PCR system with TaqMan SNP Genotyping Assays. Significant statistical differences were observed in allelic frequencies (allele T and allele G) between the non-syndromic cleft palate only and control groups in female subjects, with an allelic odds ratio of 1.455 (95% confidence interval: 1.026-2.064) and P < 0.05. These study findings suggest that nonsyndromic isolated cleft palate might be influenced by variation of MEOX2, especially SNP rs2237493 in Vietnamese females.

Discovery of a rare and striking new pierid butterfly from Panama (Lepidoptera: Pieridae).

We here describe and name a distinctive new pierid species in the subfamily Pierinae, Catasticta sibyllae Nakahara, Padrón MacDonald, n. sp. from western Panama. Catasticta. sibyllae n. sp. is known from only two male specimens collected at two sites which are approximately 130 km apart in western Panama. This new species is the only species in the genus without markings in the median area of both surfaces of forewing and hindwing, and our molecular data suggest that the Peruvian species C. lisa Baumann Reissinger, 1969 is its sister species.

Description of a new species of Euptychia Hübner, 1818 (Lepidoptera: Nymphalidae: Satyrinae) from the western Andes.

We describe a new nymphalid butterfly, Euptychia favonius sp. nov., from the western Andes. Adults and genitalia for this new taxon are illustrated and a distribution map is provided. The taxonomy of its sister species, E. meta Weymer, 1911, is discussed, and a lectotype is designated for E. meta.

Description of a new genus for Euptychia hilara (C. Felder & R. Felder, 1867) (Lepidoptera: Nymphalidae: Satyrinae).

Based on external morphology, food plant records for caterpillars, and molecular analysis, Euptychia hilara (C. Felder & R. Felder, 1867) is removed from Euptychia Hübner 1818. A new genus, Inbio Nakahara & Espeland gen. nov., is proposed for this taxon. Inbio hilara comb. nov. is a member of a monophyletic clade containing Cyllopsis Felder, 1869, Paramacera Butler, 1868, and Atlanteuptychia Freitas, Barbosa & Mielke, 2013, although it can be morphologically distinguished from these genera. Lectotypes for Neonympha hilara C. Felder & R. Felder, 1867 and Euptychia anacleta Butler, 1877 (a synonym of E. hilara) are designated herein.

Nomenclatural status of Euptychia mollina Hübner, 1818 (Lepidoptera: Nymphalidae: Satyrinae).

The purpose of this note is to clarify the nomenclatural status of Euptychia mollina Hübner, 1818, the type species of Euptychia Hübner, 1818, as there seems to be confusion regarding its year of publication. Due to an unfortunate oversight, Lamas (2004) listed the name as Euptychia mollina (Hübner, [1813]), and this mistake has been repeated in the subsequent literature (e.g. Brévignon 2005; Warren et al. 2014; Neild et al. 2014).

Taxonomic status and redescription of Magneuptychianebulosa (Butler, 1867) (Lepidoptera, Nymphalidae, Satyrinae) with a lectotype designation.

A redescription of Magneuptychianebulosa (Butler, 1867), a poorly known euptychiine butterfly, is given here, and accurate distributional data are provided for the first time. Taxonomic status of this taxon has been discussed by comparing its morphology against its possible congeners. In addition, lectotype designation for Magneuptychianebulosa is provided in order to objectively establish the identity of this taxon and consequently stabilize the nomenclature.

ResumenSe presenta una redescripción de Magneuptychianebulosa (Butler, 1867), una mariposa euptychina poco conocida y se brinda por primera vez datos de su distribución. Se discute el estatus taxonómico de este taxón por comparación de su morfología con la de posibles congéneres. En adición, se provee un lectotipo para Magneuptychianebulosa, en búsqueda de establecer objetivamente la identidad de este taxa y consecuentemente estabilizar su nomenclatura.

Two new species of Euptychia Hübner, 1818 from the upper Amazon basin (Lepidoptera, Nymphalidae, Satyrinae).

Two new species of Euptychia Hübner, 1818 are described from the upper Amazon basin: Euptychia attenboroughi Neild, Nakahara, Fratello & Le Crom, sp. n. (type locality: Amazonas, Venezuela), and Euptychia sophiae Zacca, Nakahara, Dolibaina & Dias, sp. n. (type locality: Acre, Brazil). Their unusual facies prompted molecular and phylogenetic analyses of one of the species resulting in support for their classification in monophyletic Euptychia. Diagnostic characters for the two species are presented based on wing morphology, wing pattern, presence of androconial patches on the hindwing, and genitalia. Our results indicate that the projection of the tegumen above the uncus, previously considered a synapomorphy for Euptychia, is not shared by all species in the genus. The adults and their genitalia are documented, and distribution data and a map are provided.

ResumoDuas novas espécies de Euptychia Hübner, 1818 são descritas da bacia do alto Rio Amazonas: Euptychia attenboroughi Neild, Nakahara, Fratello & Le Crom, sp. n. (localidade tipo: Amazonas, Venezuela), Euptychia sophiae Zacca, Nakahara, Dolibaina & Dias, sp. n. (localidade tipo: Acre, Brasil). Suas aparências não usuais levaram a análises molecular e filogenética de uma das espécies que sustentam sua classificação como Euptychia. São apresentados caracteres diagnósticos para as duas espécies baseados na morfologia e no padrão de coloração de suas asas, presença de manchas androconiais nas asas posteriores e suas genitálias. Os resultados indicam que a projeção do tegume sobre o unco, previamente considerada uma sinapomorfia para Euptychia, não é compartilhada por todas as espécies do gênero. São ilustrados os adultos e suas genitálias, bem como dados e mapa de distribuição.

ResumenSe describen dos nuevas especies de Euptychia Hübner, 1818 para la cuenca alta del Río Amazonas: Euptychia attenboroughi Neild, Nakahara, Fratello & Le Crom, sp. n. (localidad típica: Amazonas, Venezuela) y Euptychia sophiae Zacca, Nakahara, Dolibaina & Dias, sp. n. (localidad típica: Acre, Brasil). Sus apariencias inusuales motivaron el análisis molecular y filogenético de una de las especies, lo cual sustenta su clasificación como Euptychia. Se presentan caracteres diagnósticos para las dos especies basados en la morfología y patrón de coloración alar, presencia de parches androconiales en las alas posteriores y sus genitales. Los resultados indican que la proyección del tegumen sobre el uncus, previamente considerada una sinapomorfía para Euptychia, no es compartida por todas las especies del género. Se ilustran los adultos y sus genitales, y se proporcionan datos y mapa de distribución.

Role of Caribbean Islands in the diversification and biogeography of Neotropical Heraclides swallowtails.

Numerous hypotheses on the evolution of Neotropical biodiversity have stimulated research to provide a better understanding of diversity dynamics and distribution patterns of the region. However, few studies integrate molecular and morphological data with complete sampling of a Neotropical group, and so there has been little synthesis of the multiple processes governing biodiversity through space and time. Here, a total-evidence phylogenetic approach is used to reconstruct the evolutionary history of the butterfly subgenus Heraclides. We used DNA sequences for two mitochondrial genes and one nuclear gene and coded 133 morphological characters of larvae and adults. A robust and well-resolved phylogeny was obtained using several analytical approaches, while molecular dating and biogeographical analyses indicated an early Miocene origin (22 Mya) in the Caribbean Islands. We inferred six independent dispersal events from the Caribbean to the mainland, and three from the mainland to the Caribbean, and we suggest that cooling climates with decreasing sea levels may have contributed to these events. The time-calibrated tree is best explained by a museum model of diversity in which both speciation and extinction rates remained constant through time. By assessing both continental and fine-scale biodiversity patterns, this study provides new findings, for instance that islands may act as source of diversity rather than as a sink, to explain spatio-temporal macroevolutionary processes within the Neotropical region.

A new species of Euptychia Hübner, 1818 (Lepidoptera: Nymphalidae: Satyrinae: Satyrini) from Mount Roraima, Guyana.

A new nymphalid species in the subtribe Euptychiina, Euptychia roraima Nakahara, Fratello & Harvey n. sp., is described from Mount Roraima, Guyana. Both internal and external morphology of E. roraima are compared against several Euptychia species and the relationship between E. roraima and congeners is briefly discussed. A strong case is put forth for further and extensive exploration of the Pantepui region concerning its poorly known butterfly fauna.