One of the richest butterfly radiations in the Neotropical lowlands is the nymphalid satyrine subtribe Euptychiina, which is also one of the three most poorly studied groups of ‘true butterflies’ (Papilionoidea). In fact, few other animal groups that are so taxonomically challenging are as speciose, large, conspicuous and commonly encountered by researchers, students and naturalists. Hundreds of thousands of euptychiine specimens in museum collections hold untapped data for studies in evolution, biogeography and conservation. However, we estimate that almost half of these specimens cannot be confidently identified.
More than 400 species in 42 genera are currently recognized (our research now suggests 500+ species and 70+ genera), with at least 20% of species still undescribed, a remarkable fraction among butterflies. Molecular phylogenies suggest that Euptychia itself may not be related to remaining “euptychiines”, and the generic classification is chaotic, with 65% of genera currently invalid. Modern monographs treat only five euptychiine genera (15%), with references for remaining genera now a century old. Because of their drab wing patterns, difficulties in identifying the often highly cryptic species and their sedentary life-style, most euptychiine collections are in disarray, and the biology of most species remains unstudied. In the field, however, euptychiines are a common and diverse component of communities from the rainforest to the cerrado. Reliable guides to the taxonomy and identification of euptychiines should therefore unlock the group’s great potential for broader studies in evolution, ecology, biogeography and conservation.
The USA’s National Science Foundation is funding a multi-year project to revise the systematics of Euptychiina, under the project title: “ARTS: Phylogeny and systematic revision of the diverse and cryptic Euptychiina (Lepidoptera, Nymphalidae, Satyrinae)” (DEB-1256742). The project has 5 main goals:
- (1) to combine morphological and molecular data to produce the first ‘total evidence’ phylogenetic hypothesis for the subtribe, and to use this to reliably define genera;
- (2) to complete monographs for the majority of the largest genera not recently revised, tentatively including: Euptychia, Taygetis, Magneuptychia, Cissia, Paryphthimoides, Pareuptychia, Erichthodes, Euptychoides, Hermeuptychia, Caeruleuptychia. A secondary goal is to describe, or provide support and/or data for collaborating researchers to describe, the 21+ known undescribed species in other genera;
- (3) to provide training for butterfly systematists at the University of Florida and at the University of Campinas, Brazil;
- (4) to use project results to test hypotheses about Euptychiina evolution;
- (5) to develop collection and web-based electronic resources to facilitate communication and to build a collaborative research program on the group.
The project involves collaboration among many researchers in North and South America and Europe, and we would be pleased to hear from anyone with an interest in research on euptychiines (contact Keith Willmott at kwillmott@flmnh.ufl.edu). We hope to provide resources such as scannned original descriptions for euptychiine names, specimen databases, images of type and other specimens, images of morphology, DNA sequence data, and other data to facilitate research on the group.
1. Generic revisions or systematic studies are underway in a number of genera. See People page for researchers that we currently know of who have an interest in euptychiines, and their current main projects. Recently published papers and papers in preparation are listed below under ‘Euptychiina Papers’.
2. Curation and databasing continues in several priority collections, including the NHM (London, UK), the FLMNH (Gainesville, Florida, USA), and the MUSM (Lima, Peru). We are focusing mainly on genera under revision.
3. Field work is ongoing in Ecuador, Peru, French Guiana and Brazil, with a focus on collecting material to contribute to resolving taxonomic questions in ongoing revisions.
4. In the molecular lab we continue sequencing the COI ‘barcode’ to help resolve species complexes. Analysis is in progress of a dataset including >400 Euptychiina species, to help clarify the generic classification and relationships.
Eduardo Barbosa
Graduate student at the University of Campinas, Campinas, Brazil
Phylogeny and revision of Yphthimoides and relatives based on DNA and morphology.
Pierre Boyer
Le Puy Sainte Réparade, France
General
Christián Brévignon
French Guiana
Guianan euptychiines
Andy Brower
Professor at Middle Tennessee State University, Murfreesboro, TN
Euptychiina molecular phylogenetics
Tyler Cook
Undergraduate student at the University of Florida, Gainesville, FL
Morphology of Taygetis laches group
Cassandra Doll
Undergraduate student at the University of Florida, Gainesville, FL
Morphology and DNA barcodes of Taygetis sosis group
Nathan Duerr
Undergraduate student at the University of Florida, Gainesville, FL
Curating Magneuptychia and Taygetis-clade
Marianne Espeland
Postdoctoral research associate at the Florida Museum of Natural History, University of Florida, Gainesville, FL
Coordinating total-evidence higher level phylogeny, Euptychiina phylogenomics and diversification. Mentoring of undergraduate students.
Steve Fratello
USA
Guianan euptychiines
Andre Freitas
Professor the University of Campinas, Campinas, Brazil
Generic revisions, phylogenetics
Nick Grishin
Professor at the University of Texas, Dallas
Revision of Hermeuptychia, genomics
Blanca Huertas
Curator, Natural History Museum, London
Revision of Splendeuptychia, Colombian euptychiines
Akito Kawahara
Assistant curator at the Florida Museum of Natural History, University of Florida, Gainesville, FL
Euptychiina molecular phylogenetics
Gerardo Lamas
Museo de Historia Natural, Universidad Mayor de San Marcos, Lima, Peru
Peruvian euptychiines, generic revisions
Jean-François Le Crom
Colombia
Colombian euptychiines
Mario Marín
Graduate student at the University of Campinas, Campinas, Brazil
Euptychiina morphological phylogenetics, revision of Pareuptychia, Moneuptychia
Olaf Mielke
Professor at the Federal University of Paraná, Curitiba, Brazil
Brazilian euptychiines
Jackie Miller
Curator at the Florida Museum of Natural History, University of Florida, Gainesville, FL
Revision of Taygetis
Brittany Milo
Undergraduate student at the University of Florida, Gainesville, FL
Morphology and DNA barcodes of Taygetis virgilia group
Shinichi Nakahara
Graduate student at the Florida Museum of Natural History, University of Florida, Gainesville, FL
Revisions of Euptychia, Magneuptychia
Vanessa Navas
Undergraduate student at the University of Florida
Barcoding of Taygetis
Andrew Neild
United Kingdom
Venezuelan euptychiines
Nicole Okuthe
Undergraduate student at the University of Florida, Gainesville, FL
Molecular and morphological species delimitation in Forsterinaria
Alexandra Pagac
Undergraduate student at the University of Florida, Gainesville, FL
Databasing Euptychiina in the FLMNH
Carlos Peña
Postdoctoral researcher at the University of Turku, Finland
Euptychiina phylogenetics
Tomasz Pyrcz
Professor at the Zoological Museum of the Jagiellonian University, Krakow, Poland
Andean and southern euptychiines, Forsterinaria
Lindsay Richards
Undergraduate student at the University of Florida
Barcoding of Taygetis laches group
Noemy Seraphim
Graduate student at the University of Campinas, Campinas, Brazil
Revision of Hermeuptychia
Denise Tan
Graduate student at the Florida Museum of Natural History, University of Florida, Gainesville, FL
Euptychiina behavior and speciation, systematics of Hermeuptychia
Stephanie Tyler
Undergraduate student
Revision of Chloreuptychia
Angel Viloria
Venezuela
Venezuelan euptychiines
Niklas Wahlberg
Professor at Lund University, Lund, Sweden
Euptychiina molecular phylogenetics
Haydon Warren-Gash
France
General
Lei Xiao
Molecular lab manager at McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural History, University of Florida, Gainesville, FL
Molecular phylogenetics of Euptychiina.
Keith Willmott
Associate curator at the at the Florida Museum of Natural History, University of Florida, Gainesville, FL
Ecuadorian euptychiines, revisions of Hermeuptychia, Chloreuptychia.
Thamara Zacca
Graduate student at Universidade Federal do Parana, Brazil
Revisions of Cissia, Magneuptychia, Paryphthimoides
Previous members
Sarah Nguyentran
Undergraduate student at the University of Florida
Barcoding of various euptychiines
Tatiana Pomerantz
Undergraduate student at the University of Florida, Gainesville, FL
Revision of Euptychoides based on DNA and morphology
Gebreyes Kassu
Undergraduate student at the University of Florida, Gainesville, FL
Extraction and DNA barcoding of Euptychiina
Malaz Idris
Undergraduate student at the University of Florida, Gainesville, FL
DNA barcoding and phylogenetics of Taygetis and relatives
Neha Jagtap
Undergraduate student at the University of Florida, Gainesville, FL
DNA barcoding of Brazilian Euptychiina
The Satyrinae is one of the most diverse nymphalid subfamilies, with 2600 species occurring on all continents except Antarctica (Ackery et al., 1999), and the Euptychiina is an almost entirely Neotropical subtribe of the most diverse tribe, the global Satyrini (Peña & Wahlberg, 2008). Most euptychiines are small with dark brown wings marked by a few simple ocelli, although several genera have white or brilliant blue wings. The majority of euptychiines are lowland species, except for the largely Andean Forsterinaria (Peña & Lamas, 2005). Community diversity peaks in the W. Amazon, where 100 species can coexist (Lamas et al., 1991; Brown, 1996), while the Atlantic region of Brazil also has a diverse, endemic fauna. A single Asian genus, Palaeonympha, is also apparently a euptychiine (Miller, 1968; Peña et al., 2006, 2010).
Moderately detailed descriptions of early stages have been published for only 20 species (5% of the subtribe) (Müller, 1886; Singer et al., 1983; Young, 1984; Murray, 2001, 2003; Freitas, 2003, 2004a,b, 2007; Freitas & Peña, 2006; Kaminski & Freitas, 2008), and hostplant records are similarly scarce. Like most satyrines, larvae of most species feed on Gramineae, grasses and bamboo (Beccaloni et al., 2008), with some records on Cyperaceae and Marantaceae, except for Euptychia which feed on mosses and lycopsids (Singer et al., 1971; Singer & Mallet, 1986; DeVries, 1986; Brévignon, 2008). Development times are relatively long among tropical butterflies, up to three months or more (Freitas, unpub. data from > 50 species and 27 genera). Eggs are generally isolated and lack chorionic sculpturing. Larvae have bifid “tails” and may bear head horns that vary in size. Pupae are generally short, squat and smooth. Adults are rarely encountered at nectar, but often feed on decaying fruits or tree sap, and on carrion and animal feces. Some species are crepuscular (e.g. Taygetis, Murray, 2003), and males of forest species patrol territories in the understory, often on ridgetops, in the late afternoon (Peixoto & Benson, 2009; Willmott, pers. obs.). Males often bear tufts of hair-like scales or other androconia on the wings that presumably function in courtship.
The current classification includes more than 400 species (Lamas, 2004; Freitas, Willmott, unpub.; Huertas et al., 2009; Peña et al., 2010; Huertas, 2011; Pulido et al., 2011; Brévignon & Benmesbah, 2011; Matos et al., 2012; Freitas et al., 2011, 2012; Zacca et al., 2013) in 42 genera, with from 1 (10 genera) to 47 species. Remarkably, 80 (20%) of the known species are undescribed, a statistic exceeded among the true butterflies only by the Lycaenidae (34%) and Pronophilina (27%) (Lamas, 2004). Most known undescribed species are distinctive but poorly represented in historical collections, and many cryptic species (e.g. Willmott & Hall, 1995) surely remain to be discovered. For example, Miller (1974) described 11 of the 28 species of Cyllopsis, and Peña and Lamas (2005) described 12 of the 23 species in Forsterinaria. Most inventories contain undetermined species (e.g. Brévignon, 2005, 2007, 2008; Brévignon & Benmesbah, 2011), with recent fieldwork in Brazil revealing 8 new species in Yphthimoides, Splendeuptychia and Moneuptychia (Freitas 2004b, 2007). Many euptychiines are externally very similar, and some are apparently highly variable; with only five euptychiine genera (15%) the subject of modern revisions (Miller, 1972, 1974, 1976, 1978; Peña & Lamas, 2005), references for generic and species identification are urgently needed.
Miller (1968) defined the subtribe by external morphology, and molecular (Murray & Prowell, 2005; Peña et al., 2006) and morphological (Marin, 2010) studies have largely confirmed the current classification (Lamas, 2004), except for excluding Oressinoma and including Amphidecta and the Asian Palaeonympha. The position of Euptychia itself, however, is still unresolved (Murray & Prowell, 2005); the most comprehensive molecular phylogeny of Euptychiina to date (Peña et al., 2010), based on 4447 bp from 5 genes, for 108 euptychiine species and 18 outgroup taxa, did not recover a monophyletic Euptychiina. However, Marin (2010) did recover the subtribe as monophyletic and found a number of clades in common with Peña et al. (2010). A major goal of our research, therefore, will be to integrate and expand these data matrices to generate a robust higher-level phylogeny for the group.
If the relationships of Euptychiina within the Satyrini are still unclear, the generic classification is arguably the most chaotic for any butterfly group of similar diversity. Weymer (1910-11) classified the majority of Neotropical species (270 spp.) as “Euptychia“, presumably because of their wing pattern homogeneity, and D’Abrera (1988) continued the trend, ignoring a large number of generic names that were introduced by Forster (1964). Forster’s diagnoses were based partly on wing pattern and male genitalia, but also on overall appearance, and his characters are often unreliable. Lamas (2004) moved almost all “Euptychia” (sensu D’Abrera) into Forster’s genera, which constitute 26 of the currently accepted 39 Neotropical genera. Although the Lamas classification is a critical framework for future work, recent cladistic analyses reveal much remaining paraphyly and polyphyly among genera (Murray & Prowell, 2005; Peña et al., 2006). Peña et al. (2010) tested the monophyly of 23 Euptychiina genera and found that only 8 (35%) are monophyletic. Often, supposed congeners occurred within different species groups, such as Splendeuptychia, whose species appear in the Splendeuptychia, Pareuptychia and Hermeuptychiaclades. Marin (2010) found similar results using morphological characters; only 10 of 22 Euptychiina genera tested (45%) proved monophyletic. Recent taxonomic revisions based on a molecular phylogeny of the nine Taygetis-clade genera resulted in the synonymy of three genera, rearrangement of three genera and revelation of two undescribed genera (Matos et al., 2012). Based on an expanded higher-level phylogeny, we hope to revise generic limits and describe useful morphological characters for generic identification.
This list includes a non-exhaustive list of papers on Euptychiina systematics and biology, with links to PDF files where possible. In addition, below we list papers in progress on Euptychiina with their lead authors – please email Keith Willmott (kwillmott@flmnh.ufl.edu) to add papers to this list. If you would like to contribute to a paper in progress please contact the lead author.
Papers in progress
Manuscript Submitted or in Press:
Nakahara, S., Huertas, B., Nandi Das, G., Willmott, K. R. Revision of the “celia clade” of Pseudodebis Forster, 1964, with: two new species and notes on Papilio phorcys Fabricius, 1793 (Lepidoptera: Nymphalidae: Satyrinae).
Manuscript Essentially Complete:
Revision of Splendeuptychia (Lead author(s): Huertas (b.huertas@nhm.ac.uk))
Combining target enrichment and Sanger sequencing data to clarify the systematics of the large Neotropical butterfly subtribe Euptychiina (Satyrinae, Nymphalidae). Espeland, M., et al. …
Revision of Euptychia (Lead author(s): Nakahara (snakahara@ufl.edu))
Manuscript Writing:
Inventory of Imataca (Lead author(s): Nakahara/Orellana (snakahara@ufl.edu))
Taxonomic revision of the Euptychiina genus Amphidecta A. Butler, 1867 (Lepidoptera: Nymphalidae, Satyrinae) (Lead author(s): Eduardo Barbosa)
Morphological and molecular marker to disentangling Pareuptychia species complex (Nymphalidae: Satyrinae: Euptychiina). Mario A. Marín, Sandra I. Uribe, Anete P. Souza, André V. L. Freitas
Description of a new genus for Pharneuptychia innocentia André V. L. Freitas, Eduardo Barbosa, Thamara Zacca & Ricardo Siewert
Taxonomic revision of Yphthimoides (Lead author(s): Eduardo Barbosa)
Taxonomic revision of Pharneuptychia (Lead author(s): Eduardo Barbosa)
Yphthimoides renata species complex (Lead author(s): Eduardo Barbosa)
Using genome-wide, single-nucleotide polymorphisms to rigorously investigate phylogenetic relationships and species limits in Ecuadorian Hermeuptychia (Lead author(s): Tan (denisetsh@gmail.com))
At the limit of DNA barcoding: morphology helps resolve taxonomic complexities in Taygetis laches and relatives (Lepidoptera: Nymphalidae: Euptychiina) (Lead author(s): Cook/Willmott (kwillmott@flmnh.ufl.edu))
A taxonomic review of Taygetis virgilia (Cramer, 1776) and similar species (Lepidoptera, Nymphalidae, Satyrinae, Euptychiina) (Lead author(s): Milo/Willmott (kwillmott@flmnh.ufl.edu))
A taxonomic review of the genus Forsterinaria focused on Ecuador (Lead author(s): Okuthe/Willmott (kwillmott@flmnh.ufl.edu))
Data Gathering:
A revision of the Caeruleuptychia caerulea species group (Lead author(s): Nakahara (snakahara@ufl.edu))
A revision of the Caeruleuptychia umbrosa species group (Lead author(s): Nakahara (snakahara@ufl.edu))
Names for Hermeuptychia species: assessment of existing names and descriptions of new taxa (Lead author(s): Tan/Grishin/Willmott (kwillmott@flmnh.ufl.edu))
Taxonomy of Chloreuptychia (Lead author(s): Willmott/Nakahara (kwillmott@flmnh.ufl.edu))
Euptychoides revisions (Lead author(s): Marin/Nakahara/Willmott (kwillmott@flmnh.ufl.edu))
Taxonomic notes on Taygetis sosis and relatives (Lead author(s): Willmott/Nakahara/Miller (kwillmott@flmnh.ufl.edu))
Revision of Taygetis (Lead author(s): Willmott/Nakahara/Miller (kwillmott@flmnh.ufl.edu))
New species of Hermeuptychia from Brazil and Colombia. Noemy Seraphim, Mario A. Marín, Sandra I. Uribe, André V. L. Freitas
Evolutionary history of butterflies in the “Pareuptychia clade” (Nymphalidae: Satyrinae: Euptychiina). Mario A. Marín, Sandra I. Uribe, Anete P. Souza, André V. L. Freitas
Biogeography and diversification of Euptychiina. Marianne Espeland, Keith Willmott
Intended:
Revision of Magneuptychia (Lead author(s): Nakahara (snakahara@ufl.edu)/Zacca)
Project publications on Euptychiina
Corahua-Espinoza, T., Nakahara, S., Kabir, J., Shellman, B., Tejeira, R., Ccahuana, R., Gallice, G. 2022. Immature stages and new host plant records for four satyrine species feeding on herbaceous bamboos in southeastern Peru (Lepidoptera: Nymphalidae: Satyrinae: Satyrini). Zootaxa 5125(1): 37-62. 10.11646/zootaxa.5125.1.3.
Barbosa, E. P., Seraphim, N., Valencia, G., Azeredo-Espin, A. M. L., Freitas, A. V. L. 2022. Phylogenetic systematics of Yphthimoides Forster, 1964 and related taxa, with notes on the biogeographical history of Yphthimoides species. Molecular Phylogenetics and Evolution 168, 107390. 10.1016/j.ympev.2022.107390.
Nakahara, S., Rodríguez-Melgarejo, M., Kleckner, K., Espeland, M., Casagrande, M. M., See, J., Gallice, G., Corahua, T., Barbosa, E. P., Lamas, G., Willmott, K. R. 2022. Systematic revision of a new butterfly genus, Cisandina Nakahara & Espeland, n. gen., with descriptions of three new taxa (Lepidoptera: Nymphalidae: Satyrinae). Insect Systematics and Diversity, 6(1): 2; 1-30. 10.1093/isd/ixab028.
Hurtado, T., Nakahara, S., Rodríguez-Melgarejo, M., Tejeira, R., See, J., Ccahuana, R., Gallice, G. 2021. Complete life cycle of the euptychiine butterfly Taygetis cleopatra (C. Felder & R. Felder, 1867) (Lepidoptera: Nymphalidae: Satyrinae) in southeastern Peru. Tropical Lepidoptera Research 31(3): 179-185. 10.5281/zenodo.5777169.
Nakahara, S., Barbosa, E. P., Nakamura, I., Lamas, G., Freitas, A. V. L. 2021. Description of a new species of Yphthimoides Forster, 1964 from Peru (Lepidoptera: Nymphalidae: Satyrinae). Tropical Lepidoptera Research 31(3): 138-144. 10.5281/zenodo.5776879.
Tejeira, R., Ccahuana, R., Hurtado, T., Nakahara, S., See, J., Rodríguez-Melgarejo, M., Corahua-Espinoza, T., Gallice, G. 2021. Immature stages of Chloreuptychia marica (Weymer, 1911) (Lepidoptera: Nymphalidae: Satyrinae: Satyrini). Tropical Lepidoptera Research 31(1): 96-100. 10.5281/zenodo.5600408.
Freitas, A. V. L., Rosa, A. H. B., Kaminski, L. 2021. Immature stages of Stegosatyrus ocelloides (Nymphalidae: Euptychiina), a grassland specialist butterfly. Tropical Lepidoptera Research 31(2): 127-133. 10.5281/zenodo.5600441.
Freitas, A. V. L., Brabosa, E. P., Carreira, J. Y. O. 2021. Immature stages and natural history of Yphthimoides borasta (Nymphalidae: Euptychiina). Tropical Lepidoptera Research 31(1): 42-47. 10.5281/zenodo.4721637.
Nakahara, S., Willmott, K. R., MacDonald, J. R., Thurman, A. 2021. Description of an enigmatic new species of Chloreuptychia Forster, 1964 from Panama (Lepidoptera: Nymphalidae: Satyrinae). Tropical Lepidoptera Research, 31(3): 145-150. DOI: 10.5281/zenodo.577691.
Rosa, A. H. B., Huertas, B., Willmott, K. R., Barbosa, E. P., Machado, P. A., Mielke, O. H. H., Canaan, C. H. P., Freitas, A. V. L. 2021. Fifty years without a name: a new species of Splendeuptychia Forster (Lepidoptera: Nymphalidae: Satyrinae). Zootaxa 5061(1): 95-114. 10.11646/zootaxa.5061.1.
Zacca, T., Casagrande, M. M., Mielke, O. H. H., Huertas, B., Freitas, A. V. L., Marín, M. A., Espeland, M., Willmott, K. R. 2021. A new euptychiine butterfly species from south Brazil and taxonomic rearrangements for Taydebis Freitas, 2013 and Hermeuptychia Forster, 1964 (Lepidoptera: Nymphalidae: Satyrinae). Zootaxa 5023(4): 555-570. 10.11646/zootaxa.5023.4.5
Nakahara, S., Matos-Maraví, P., Willmott, K. R., Nakamura, I., MacDonald, J. R. 2021. Description of a new species of Pseudodebis Forster, 1964 from Central America (Lepidoptera: Nymphalidae: Satyrinae). Tropical Lepidoptera Research, 31(1): 35-41. 10.5281/zenodo.4721627
Tan, D., Parus, A., Dunbar, M., Espeland, M., Willmott, K. R. 2021. COI barcode species delineation methods imply critically underestimated diversity in “common” Hermeuptychia butterflies (Lepidoptera: Nymphalidae: Satyrinae). Zoological Journal of the Linnean Society, XXX: 1-15. 10.1093/zoolinnean/zlab007.
Nakahara, S., Matos-Maraví, P., Schwartz, J., Willmott, K. R. 2021. Assessing generic synapomorphy of Pseudodebis Forster, 1964 (Lepidoptera: Nymphalidae: Satyrinae) and a recent speciation between two new species with a shift in elevation in the western Andes. Invertebrate Systematics 35: 158-180 10.1071/IS20024
Nakahara, S., Lamas, G., Willmott, K. R., Espeland, M. 2020. Description of a new genus and species for a common and widespread Amazonian satyrine butterfly (Lepidoptera: Nymphalidae: Satyrinae: Satyrini). PeerJ 8:e10324, 1-25 10.7717/peerj.10324.
Nakahara, S., Kleckner, K., Lamas, G., Huertas, B., Willmott, K. R. 2020. A contribution towards the systematics of Magneuptychia Forster, 1964: Caeruleuptychia francisca (Butler, 1870), n. comb. (Lepidoptera: Nymphalidae: Satyrinae). Tijdschrift voor Entomologie 163: 51-61.
Zacca, T., Casagrande, M. M., Mielke, O H. H., Huertas, B., Espeland, M., Freitas, A. V. L., Willmott, K. R., Nakahara, S., Lamas, G. 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): 001-034.
Zacca, T., Casagrande, M., Mielke, O. H. H., Huertas, B., Barbosa, E., Freitas, A. V. L., Willmott, K. R. 2020. No more in limbo! Description of Emeryus gen. nov. to accommodate three species formerly placed in Paryphthimoides Forster, 1964 (Lepidoptera: Nymphalidae, Satyrinae). Austral Entomology 59: 505-523.
Zacca, T., Casagrande, M. M., Mielke O. H. H., Huertas, B., Barbosa E. P., Freitas, A. V. L., Lamas, G., Espeland, M., Brévignon, C., Nakahara, S., Checa, M. F., Willmott, K. R. 2020. Systematics of the Neotropical butterfly genus Paryphthimoides Forster, 1964 (Lepidoptera: Nymphalidae: Satyrinae), with descriptions of seven new taxa. Insect Systematics and Evolution 52(1): 42-96.
Nakahara, S., Barbosa, E. P., Willmott, K. R., Lamas, G., Freitas, A. V. L. 2019. Two new species of Taygetina Forster, 1964 from Ecuador, Peru and Brazil with a possible case of “juxta loss” in butterflies (Lepidoptera: Nymphalidae: Satyrinae). Insect Systematics and Diversity 3(6): 9; 1-13.
Nakahara, S., Gallardo, R. 2019. Scientific Note: Reinstatement of Euptychia sericeella Bates, 1865: Amiga sericeella stat. rev., with corrigenda to Nakahara et al. (2019) (Lepidoptera: Nymphalidae: Satyrinae). Tropical Lepidoptera Research 29(2): 111-114.
Melo, D. H. A., Filgueiras, B. K. C., Iserhard, C. A., Iannuzzi, I., Freitas, A. V. L., Leal, I. R. 2019. Effect of habitat loss and fragmentation on fruit-feeding butterflies in the Brazilian Atlantic Forest. Canadian Journal of Zoology 97: 588-596.
Lourenço, G. M., Soares, G. R., Santos, T. P., Dáttilo, W., Freitas, A. V. L., Ribeiro, S. P. 2019. Equal but different: Natural ecotones are dissimilar to anthropic edges. PLoS ONE 14(3): e0213008.
Freitas, A. V. L., Mota, L. L., Barbosa, E. P., Carreira, J. Y. O. 2019. Immature stages of the Selaginella-feeding Euptychia mollina (Nymphalidae: Satyrinae) from the Brazilian Amazon. Zoologia 36: e32053.
Rosa, A. H. B., Perillo, L. N., Neves, F. S., Ribeiro, D. B., Freitas, A. V. L. 2019. Butterflies collected using malaise traps as useful bycatches for ecology and conservation. Journal of Threatened Taxa 11(9): 14235-14237.
Freitas, A. V. L., Mota, L. L., Zacca, T., Barbosa, E. P. 2019. Description of a new and highly distinctive genus and species of Euptychiina (Lepidoptera: Nymphalidae: Satyrinae) from the Brazilian southern Amazon. Revista Brasileira de Entomologia 63: 254-261.
Uehara-Prado, M., Freitas, A. V. L. 2019. Population structure of Taygetis ypthima (Nymphalidae, Euptychiina) in southeast Brazil. Journal of the Lepidopterists’ Society 73(1): 63-64.
Nakahara, S., Zacca, T., Dias, F. M. S., Dolibaina, D. R., Xiao, L., Espeland, M., Casagrande, M. M., Mielke, O. H. H., Lamas, G., Huertas, B., Kleckner, K., Willmott, K. R. 2019. Revision of the poorly known Neotropical butterfly genus Zischkaia Forster, 1964 (Lepidoptera, Nymphalidae, Satyrinae), with descriptions of nine new species. European Journal of Taxonomy 551: 1-67. DOI: 10.5852/ejt.2019.551.
Brévignon, C., Rosant, T., Lamas, G., Tyler, S., Willmott, K. R. 2019. Description d’une nouvelle espèce d’Euptychiina du genre Chloreuptychia (Lepidoptera, Nymphalidae, Satyrinae). Bulletin de la Société Entomologique de France 124(2): 127-138. DOI: 10.32475/bsef_2044.
Willmott, K. R., Marín, M. A., Nakahara, S., Pomerantz, T., Lamas, G., Espeland, M., Xiao, L., Hall, J. P. W., Robinson Willmott, J. I., Freitas, A. V. L. 2019. A revision of the new Andean butterfly genus Optimandes Marín, Nakahara & Willmott, n. gen., with the description of a new species (Nymphalidae: Satyrinae: Euptychiina). Tropical Lepidoptera Research 29(1): 29-44. DOI: 10.5281/zenodo.2650482.
Filgueiras, B. K. C., Melo, D. H. A., Uehara-Prado, M., Freitas, A. V. L., Leal, I. R., Tabarelli, M. 2019. Compensatory dynamics on the community structure of fruit-feeding butterflies across hyper-fragmented Atlantic forest habitats. Ecological Indicators 98: 276-284.
Iserhard, C. A., Duarte, L., Seraphim, N., Freitas, A. V. L. 2019. How urbanization affects multiple dimensions of biodiversity in tropical butterfly assemblages. Biodiversity and Conservation 26: 621-638.
Melo, D. H. A., Duarte, M., Mielke, O. H. H., Robbins, R. K., Freitas, A. V. L. 2019. Butterflies (Lepidoptera: Papilionoidea) of an urban park in northeastern Brazil. Biota Neotropica 19(1): e20180614.
Santos, J. P. dos, Freitas, A. V. L., Brown, K. S., Carreira, J. Y. O., Gueratto, P. E., Rosa, A. H. B., Lourenço, G. M., Accacio, G. M., Uehara-Prado, M., Iserhard, C. A., Richter, A., Gawlinski, K., Romanowski, H. P., Mega, N. O., Teixeira, M. O., Moser, A., Ribeiro, D. B., Araujo, P. F., Filgueiras, B. K. C., Melo, D. H. A., Leal, I. R., Beirão, M. do V., Ribeiro, S. P., Cambuí, E. C. B., Vasconcelos, R. N., Cardoso, M. Z., Paluch, M., Greve, R. S., Voltolini, J. C., Galetti, M., Regolin, A. L., Sobral-Souza, T., Ribeiro, M. C. 2019. Fruit-feeding butterflies from the Atlantic forests. Bulletin of the Ecological Society of America 100(1): e01484.
Nakahara, S., Lamas, G., Tyler, S., Marín, M., Huertas, B., Willmott, K. R., Mielke, O. H. H., Espeland, M. 2019. A revision of the new genus Amiga Nakahara, Willmott & Espeland, gen. n., described for Papilio arnaca Fabricius, 1776 (Lepidoptera, Nymphalidae, Satyrinae). Zookeys 821: 85-152.
Freitas, A. V. L., Batista Rosa, A. H., Oliveira Carreira, J. Y., Eyng Gueratto, P., Pereira Santos, J., Tacioli, A. 2018. Immature stages of two Moneuptychia from southeastern Brazil (Nymphalidae: Euptychiina). Tropical Lepidoptera Research 28(2): 100-105.
Espeland, M., Breinholt, J., Barbosa, E., Casagrande, M. M., Huertas, B., Lamas, L., Marín, M. A., Mielke, O. H. H., Miller, J. Y., Nakahara, S., Tan, D., Warren, A. D., Zacca, T., Kawahara, A., Freitas, A. V. L., Willmott, K. R. 2018. Four hundred shades of brown: Higher level phylogeny of the problematic Euptychiina (Lepidoptera, Nymphalidae, Satyrinae) based on hybrid enrichment data. Molecular Phylogenetics and Evolution 131: 116-124. DOI: 10.1016/j.ympev.2018.10.03.
See, J., Nakahara, S., Gallice, G. 2018. Immature stages of Splendeuptychia quadrina (Butler, 1869) (Lepidoptera: Nymphalidae: Satyrinae). Tropical Lepidoptera Research 28(2): 49-53.
Nakahara, S., Willmott, K. R., Mielke, O. H. H., Schwartz, J., Zacca, T., Espeland, M., Lamas, G. 2018. 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.
Willmott, K. R., Lamas, G., Radford, J., Marín, M. A., Nakahara, S., Espeland, M., Xiao, L., Hall, J. P. W. 2018. A distinctive new species of cloud forest Euptychiina (Lepidoptera: Nymphalidae: Satyrinae) from Ecuador and Peru. Tropical Lepidoptera Research 28(1): 39-45.
Zacca, T., Casagrande, M. M., Mielke O. H. H., Huertas, B., Barbosa E. P., Freitas, A. V. L., Magaldi, L. M., Espeland, M., Nakahara, S., Willmott, K. R. 2018. Systematics of the butterfly genus Cissia Doubleday, 1848 (Lepidoptera: Nymphalidae, Satyrinae) using an integrative approach. Arthropod Systematics and Phylogeny, 76(2): 349-376.
Benmesbah, M., Zacca, T., Casagrande, M. M., Mielke, O. H. H., Lamas, G., Willmott, K. R. 2018. Taxonomic notes on Papilio ocypete Fabricius, 1776 and Papilio helle Cramer, 1779 with description of two new similar species from South America (Lepidoptera: Nymphalidae: Satyrinae). Zootaxa 4425(1): 115-145.
Barbosa, E. P., Siewert, R. R., Mielke, O. H. H., Lamas, G., Willmott, K. R., Freitas, A. V. L. 2018. Redescription of Yphthimoides patricia (Hayward, 1957), with taxonomic notes on the names Euptychia saltuensis Hayward, 1962 and Yphthimoides manasses (C. Felder & R. Felder, 1867) (Nymphalidae: Satyrinae). Zootaxa, 4422(4): 537-557. DOI: 10.11646/zootaxa.4422.4.5
Freitas, A. V. L., Barbosa, E., Zacca, T., Marín, M., Beirão, M., Silva, A., Casagrande, M. M., Espeland, M., Willmott, K. R. 2018. Before it is too late: description of a new genus and species of butterfly from a highly threatened Brazilian biome. Revista Brasileira de Entomologia 62: 148-158, DOI: 10.1016/j.rbe.2018.02.00.
Warren, A. D., Nakahara, S., Llorente-Bousquets, J., Luis-Martínez, A., Miller, J. Y. 2018. A new species of Cyllopsis R. Felder, 1869 from the highlands of Chiapas, Mexico (Lepidoptera: Nymphalidae: Satyrinae). Zootaxa 4403(3): 570-577.
Santos, J. P. dos, Freitas, A. V. L., Brown, K. S., Carreira, J. Y. O., Gueratto, P. E., Rosa, A. H. B., Lourenço, G. M., Accacio, G. M., Uehara-Prado, M., Iserhard, C. A., Richter, A., Gawlinski, K., Romanowski, H. P., Mega, N. O., Teixeira, M. O., Moser, A., Ribeiro, D. B., Araujo, P. F., Filgueiras, B. K. C., Melo, D. H. A., Leal, I. R., Beirão, M. do V., Ribeiro, S. P., Cambuí, E. C. B., Vasconcelos, R. N., Cardoso, M. Z., Paluch, M., Greve, R. S., Voltolini, J. C., Galetti, M., Regolin, A. L., Sobral-Souza, T., Ribeiro, M. C. 2018. Atlantic butterflies: a data set of fruit-feeding butterfly communities from the Atlantic forests. Ecology 99(12): 2875 pp., http://onlinelibrary.wiley.com/doi/10.1002/ecy.2507/suppinfo.
Braga, L., Ramos, R. R., Aguiar, T. M. C., Freitas, A. V. L. 2018. Distribution extension of Godartiana byses (Nymphalidae: Satyrinae) and first record for the state of São Paulo, Brazil. Journal of the Lepidopterists’ Society 72(2): 176-179.
Espeland, M., Breinholt, J., Willmott, K. R., Warren, A. D., Vila, R., Toussaint, E. F. A., Maunsell, S. C., Aduse-Poku, K., Talavera, G., Eastwood, R., Jarzyna, M. A., Ries, L., Guralnick, R., Lohman, D. J., Pierce, N. E., Kawahara, A. 2018. Comprehensive higher-level phylogeny of butterflies (Papilionoidea) inferred from genomic data. Current Biology, 28: 770-778, DOI: 10.1016/j.cub.2018.01.061
Freitas, A. V. L. 2017. Immature stages of the Neotropical satyrine Taygetis acuta (Nymphalidae: Euptychiina). Tropical Lepidoptera Research, 27: 1-5.
Nakahara, S., Zacca, T., Huertas, B., Neild, A. F. E., Hall, J. P. W., Lamas, G., Holian, L. A., Espeland, M., Willmott, K. R. 2017. Remarkable sexual dimorphism, rarity and cryptic species: a revision of the ‘aegrota species group’ of the Neotropical butterfly genus Caeruleuptychia Forster, 1964 with the description of three new species (Lepidoptera, Nymphalidae, Satyrinae). Insect Systematics and Evolution, DOI 10.1163/1876312X-00002167.
Marin, M. A., Peña, C., Uribe, S. I., Freitas, A. V. L. 2017. Morphology agrees with molecular data: phylogenetic affinities of Euptychiina butterflies (Nymphalidae: Satyrinae). Systematic Entomology, 42: 768-785.
Zacca, T., Paluch, M., Siewert, R., Freitas, A. V. L., Barbosa, E., Mielke, O. H. H., Casagrande, M. M. 2017. Revision of Godartiana Forster (Lepidoptera: Nymphalidae), with the description of a new species from northeastern Brazil. Austral Entomology, 56: 169-190.
Zacca, T., Casagrande, M. M., Mielke, O. H. H., Huertas, B., Neild, A. F. E., Benmesbah, M. 2017. Description of a new species of Euptychiina (Lepidoptera: Nymphalidae: Satyrinae) from South America. Zootaxa, 4231(3): 442-450.
de Andrade, R. B., Balch, J. K., Carreira, J. Y. O., Brando, P. M., Freitas, A. V. L. 2017. The impacts of recurrent fires on diversity of fruit-feeding butterflies in a south-eastern Amazon forest. Journal of Tropical Ecology, 33 (1): 22-32.
Nakahara, S., Barbosa, E. P., Freitas, A. V. L. 2017. A potentially endangered new species of Euptychia Hübner, 1818 (Lepidoptera: Nymphalidae: Satyrinae) from the Atlantic coastal forest of Brazil. Neotropical Entomology, DOI: 10.1007/s13744-016-0466-y
Nakahara, S., Vega, G., Willmott, K. R. 2016. Description of a new species of Euptychia Hübner, 1818 (Lepidoptera: Nymphalidae: Satyrinae) from the western Andes. Zootaxa, 4184: 358-366. DOI: 10.11646/zootaxa.4184.2.7
Huertas, B., Lamas, G., Fagua, G., Mallet, J., Nakahara, S., Willmott, K. R. 2016. A remarkable new butterfly species from western Amazonia (Lepidoptera: Nymphalidae: Satyrinae). Conservación Colombiana, 24: 1-5.
Barbosa E. P., Marín M.A., Giraldo C. E., Uribe S. I., Freitas A. V. L. 2016. Description of two new species of the Neotropical genus Yphthimoides Forster, 1964 (Lepidoptera: Nymphalidae: Satyrinae) from the “renata clade”. Neotropical Biodiversity, 2(1): 87-98.
Freitas, A. V. L., Barbosa, E. P., Marin, M. A. 2016. Immature stages and natural history of the neotropical satyrine Pareuptychia ocirrhoe interjecta (Nymphalidae: Satyrinae). Journal of the Lepidopterists’ Society, 70(4): 271-276.
Nakahara, S., Tan, D., Lamas, G., Parus, A., Willmott, K. R. 2016. A distinctive new species of Hermeuptychia Forster, 1964 from the eastern tropical Andes (Lepidoptera: Nymphalidae: Satyrinae). Tropical Lepidoptera Research, 26: 77-84.
Nakahara, S., Marín, M. A., Neild, A. F. E. 2016. A new species of Pareuptychia Forster, 1964 (Lepidoptera, Nymphalidae, Satyrinae) from the northwestern Amazon basin with characterisation of two potential synapomorphies for the genus. Tropical Lepidoptera Research, 26(1): 6-12.
Freitas, A. V. L., Carreira, J. Y. O., Santos, J. P., Barbosa, E. P. 2016. Immature stages and natural history of two species of Forsterinaria from southeastern Brazil (Lepidoptera: Nymphalidae). Tropical Lepidoptera Research, 26(1): 13-18.
Nakahara, S., Barbosa, E. P., Marín, M. A., Freitas, A. V. L., Pomerantz, T., Willmott, K. R. 2016. Graphita gen. nov., a new genus for Neonympha griphe C. Felder & R. Felder, 1867 (Lepidoptera, Nymphalidae, Satyrinae). Neotropical Entomology, 45(6): 675-691.
Freitas, A. V. L., Barbosa, E., Willmott, K. R., Wahlberg, N., Lamas, G. 2016. ‘Species’ from two different butterfly genera combined into one: description of a new genus of Euptychiina (Nymphalidae: Satyrinae) with unusually variable wing pattern. Revista Brasileira de Entomologia, doi:10.1016/j.rbe.2016.01.004
Nakahara, S., Hall, J. P. W., Lamas, G., Willmott, K. R. 2015. Seven new species and one new subspecies of Euptychia Hübner, 1818 (Lepidoptera: Nymphalidae: Satyrinae) from the tropical Andes. Tropical Lepidoptera Research 25(2): 63-79.
Neild, A. F. E., Nakahara, S., Zacca, T., Fratello, S., Lamas, G., Le Crom J-F., Dolibaina, D. R., Dias, F. M. S., Casagrande, M. M., Mielke, O. H. H., Espeland, M. 2015. Two new species of Euptychia Hübner, 1818 from the upper Amazon basin (Lepidoptera, Nymphalidae, Satyrinae). ZooKeys 541: 87-108.
Nakahara, S., Marín, M. A., Ríos-M, C. 2015. Taxonomic status and redescription of Magneuptychia nebulosa (Butler, 1867) (Lepidoptera: Nymphalidae: Satyrinae) with a lectotype designation. ZooKeys 503: 135-147.
Nakahara, S., Llorente-Bousquets, J. E., Luis-Martínez, A., Miller, J. Y., Warren, A. D. 2015. Two new species of Euptychia Hübner, 1818 (Lepidoptera: Nymphalidae: Satyrinae) from Mexico and Guatemala. Journal of Research on the Lepidoptera 48: 51-57.
Nakahara, S., Janzen, D. H., Hallwachs, W., Espeland, M. 2015. Description of a new genus for Euptychia hilara (Lepidoptera, Nymphalidae, Satyrinae). Zootaxa 4012(3): 525-541.
Lamas, G., Nakahara, S. 2015. Nomenclatural status of Euptychia mollina Hübner, 1818 (Lepidoptera: Nymphalidae: Satyrinae). Zootaxa 3946(4): 587-590.
Kaminski, L. A., Dell’Erba, R., Barbosa, E. P., Freitas, A. V. L. 2015. New distribution records and notes on the habitat of Magneuptychia flavofascia Zacca & Siewert, 2014 (Lepidoptera: Nymphalidae). Check List 11(4): 1692, pp. 1-3.
Freitas, A. V. L., Barbosa, E. P., Siewert, R. R., Mielke, O. H. H., Zacca, T., Azeredo-Espin, A. N. 2015. Four new species of Moneuptychia (Lepidoptera: Satyrinae: Euptychiina) from Brazil. Zootaxa 3981(4): 521-541.
Fratello, S. A., Nakahara, S., Brévignon, R. C., Harvey, D. J. 2015. Two new species of Euptychia Hübner, 1818 (Lepidoptera: Nymphalidae: Satyrinae) from the Guiana Shield, with notes on E. marceli Brévignon, 2005 and E. rufocincta Weymer, 1911. Journal of the Lepidopterists’ Society 69(4): 293-306.
Barbosa E. P., Silva A. K., Paluch M., Azeredo-Espin A. M. L., Freitas A. V. L. 2015. Uncovering the hidden diversity of the Neotropical butterfly genus Yphthimoides Forster (Nymphalidae: Satyrinae): description of three new species based on morphological and molecular data. Organisms Diversity and Evolution 15: 577-589.
Nakahara, S., Fratello, S. A., Harvey, D. J. 2014. A new species of Euptychia Hübner, 1818 (Lepidoptera: Nymphalidae: Satyrinae: Satyrini) from Mount Roraima, Guyana. Zootaxa, 3881(3): 291-300.
Neild, A. F. E., Nakahara, S., Fratello, S. A., Harvey, D. J. 2014. A new species of Euptychia Hübner, 1818 (Nymphalidae: Satyrinae: Satyrini) from the Amazon basin and the Guianas. Tropical Lepidoptera Research, 24(1): 4-9.
Cosmo, L. G., Barbosa, E. P., Freitas, A. V. L. 2014. Biology and morphology of the immature stages of Hermeuptychia atalanta (Lepidoptera: Nymphalidae: Satyrinae). Annales de la Société Entomologique de France, 50(1): 82-88.
Warren, A. D., Tan, D., Willmott, K. R. Grishin, N. V. 2014. Refining the diagnostic characters and distribution of Hermeuptychia intricata (Nymphalidae: Satyrinae: Satyrini). Tropical Lepidoptera Research 24(1): 44-51.
Freitas, A. V. L., Barbosa, E. P., Santos, J. P., Mielke, O. H. H. 2013. A new genus, Atlanteuptychia gen. nov., for Euptychia ernestina (Lepidoptera: Nymphalidae: Satyrinae), Zoologia (Curitiba), 30: 661-668.
Siewert, R. R., Zacca, T., Dias, F. M. S., Freitas, A. V. L., Mielke, O. H. H., Casagrande, M. M. 2013. The “Taygetis ypthima species group” (Lepidoptera, Nymphalidae, Satyrinae): taxonomy, variation and description of a new species. Zookeys, 356: 11-29.
Other published papers on Euptychiina
Ackery, P.R., de Jong, R., Vane-Wright, R.I. 1999. The butterflies: Hedyloidea, Hesperoidea and Papilionoidea. In: Kristensen, N.P. (Ed.), Lepidoptera: Moths and Butterflies. 1. Evolution, Systematics and Biogeography. Handbook of Zoology, vol. IV. Berlin, Walter de Gruyter. Part 35.
Alvaréz-García, H., Salinas-Gutiérrez, J. L., Luis-Martínez, A., Vásquez, A. I. 2014. New records and notes about Splendeuptychia kendalli L. Miller, 1978 (Satyrinae: Satyrini: Euptychiina) in Mexico. Southwestern Entomologist, 39: 337-342.
Anken, R. H. 1994. Zur Kenntnis neuer Euptychiini der Genus Pareuptychia Forster aus Brasilien (Lepidoptera: Satyridae). Entomologische Zeitschrift, 104(12): 221-231.
Anken, R. H. 1994. Neue Taxa des Genus Hermeuptychia Forster aus Brasilien (Lepidoptera: Satyridae). 2. Beitrag zur Kenntnis neuer neotropischer Euptychiini. Entomologische Zeitschrift, 104(14): 283-291.
Anken, R. H. 1994. Beschreibung einer neuen Art des Genus Yphthimoides Forster aus Brasilien (Lepidoptera: Satyridae). 3. Beitrag zur Kenntnis neuer neotropischer Euptychiini. Entomologische Zeitschrift, 104(16): 327-331.
Anken, R. H. 1994. Eine neue Species des Genus Paryphthimoides Forster aus Brasilien (Lepidoptera: Satyridae). 4. Beitrag zur Kenntnis neuer neotropischer Euptychiini. Entomologische Zeitschrift, 104(17): 348-351.
Anken, R. H. 1998. Eine neue Art der Gattung Splendeuptychia Forster aus dem Minas Gerais Brasiliens (Lepidoptera: Nymphalidae: Satyrinae: Euptychiini). 5. Beitrag zur Kenntnis neuer neotropischer Euptychiini. Entomologische Zeitschrift, 108(5): 184-192.
Anken, R. H. 1999. Beschreibung einer neuen Art der Gattung Paryphthimoides Forster aus dem Minas Gerais Brasiliens (Lepidoptera: Nymphalidae: Satyrinae: Euptychiini). 6. Beitrag zur Kenntnis neuer neotropischer Euptychiini. Entomologische Zeitschrift, 109(3): 108-113.
Anken, R. H. 1999. Eine neue Art der Gattung Pareuptychia Forster aus dem Minas Gerais Brasiliens (Lepidoptera: Nymphalidae: Satyrinae). 7. Beitrag zur Kenntnis neuer neotropischer Euptychiini. Entomologische Zeitschrift, 109(4): 165-172.
Anken, R. H. 1999. Zur Kenntnis einer zweiten Art der Gattung Capronnieria Forster 1964 nebst Beschreibung einer neuen Art des Genus Yphthimoides Forster 1964 aus dem Minas Gerais Brasiliens (Lepidoptera, Satyrinae, Euptychiini). 8. Beitrag zur Kenntnis neuer neotropischer Euptychiini. Entomologische Zeitschrift, 109(7): 269-279.
Aurivillius, P. O. C. 1929. Wissenschaftliche Ergebnisse der schwedischen Entomologischen Reisen des Herrn Dr. A. Roman in Amazonas 1914-1915 und 1923-1924. 13. Rhopalocera. Entomologisk Tidskrift, 50(3/4): 153-168.
Beccaloni, G. W., Viloria, A. L., Hall, S. K., Robinson, G. S. 2008. Catalogue of the Hostplants of the Neotropical Butterflies / Catálogo de las Plantas Huésped de las Mariposas Neotropicales. m3m-Monografías Tercer Milenio, Volume 8. Zaragoza, Spain: Sociedad Entomológica Aragonesa (SEA)/Red Iberoamericana de Biogeografía y Entomología Sistemática (RIBES)/Ciencia y Tecnología para el Desarrollo (CYTED)/Natural History Museum, London, U. K. (NHM)/Instituto Venezolano de Investigaciones Científicas, Venezuela (IVIC). 1-536 pp., 1 fig, 3 tabs.
Benmesbah, M. 2015. Description de deux nouvelles espèces de Satyrinae de Guyane française (Lepidoptera, Nymphalidae). Bulletin de la Société entomologique de France, 120(2): 147-156.
Benmesbah, M., Viloria, A. L., Murienne, J. 2020. Taxonomic notes on Euptychia modesta Butler, 1867, Neonympha alcinoe C. Felder & R. Felder, 1867 and Euptychia pamela Hayward, 1957 (Lepidoptera: Nymphalidae: Satyrinae), with descriptions of three new genera, five new species and two new subspecies from Central and South America. Anartia 31: 7-62.
Beutelspacher, C. R. 1982. Una nueva especie de Cyllopsis Felder (Lepidoptera: Satyridae). Anales del Instituto de Biologia Universidad Nacional Autónoma de México. Serie zoologia, 52: 361-366. PDF
Breckheimer, I., Haddad, N. M., Morris, W. F., Trainor, A. M., Fields, W. R., Jobe, T. R., Hudgens, B. R., Moody, A., Walters, J. R. 2014. Defining and evaluating the umbrella species concept for conserving and restoring landscape connectivity. Conservation Biology, 28(6): 1584-1593. PDF
Brévignon, C. 2005. Description de nouveau Satyrinae provenant de Guyane française (Lepidoptera, Nymphalidae). Lambillionea, 105: 393-404.
Brévignon, C. 2007. Description d’une nouvelle espèce du genre Taygetis Hübner, [1819] provenant de Guyane française (Lepidoptera, Nymphalidae, Satyrinae). Lambillionea, 107: 235-237.
Brévignon, C. 2008. Inventaire des Satyrinae de Guyane Française (Lepidoptera: Nymphalidae), pp. 62-94. In: Lacomme, D., Manil, L. (Eds.),Lépidoptères de Guyane, Tome 3, Rhopalocères 2. Paris, Lepidopteristes de France.
Brévignon, C., Benmesbah, M. 2011. Complément à l’inventaire des Satyrinae de Guyane (Lepidoptera: Nymphalidae), p. 36-52. In: Lacomme, D., Manil, L. (Eds.), Lépidoptères de Guyane, Tome 7, Nymphalidae. Paris, Lepidopteristes de France.
Brown, K. S., Jr. 1996. Conservation of threatened species of Brazilian butterflies. Decline and Conservation of Butterflies in Japan, 3: 45-62.
Butler, A. G. 1867. A monograph of the genus Euptychia, a numerous race of butterflies belonging to the family Satyridae; with descriptions of sixty specis new to science, and notes to their affinities. Proceedings of the Zoological Society of London, 1866(3): 484-504, pls. 39-40.
Butler, A. G. 1867b. Descriptions of some new species of Satyridae belonging to the genus Euptychia. Proceedings of the zoological Society of London 1867(1): 104-110, pls. 11-12.
Butler, A. G. 1874. Descriptions of some new species and a new genus of Diurnal Lepidoptera, in the collection of Herbert Druce, Esq. Transactions of The Royal Entomological Society of London, 22: 423-436. PDF
Butler, A. G. 1877. On new species of the genus Euptychia, with a tabular view of those hitherto recorded. 13(67): 116-128. PDF
Butler A. G., Druce, H. 1874. List of the butterflies of Costa Rica, with descriptions of new species. Proceedings of the Zoological Society of London, 42(1): 330-370. PDF
Chacón, I., Nishida, K. 2002. A new species of Cyllopsis (Lepidoptera: Nymphalidae: Satyrinae) from Costa Rica. Revista de Biología Tropical, 50:679-683. PDF
Chermock, R. L. 1947. A revisional study of the Euptychia pyracmon group. Entomological News 58(8): 193-204.
Cock, M. J. W., 2014. An updated and annotated checklist of the larger butterflies (Papilionoidea) of Trinidad, West Indies: Papilionidae, Pieridae and Nymphalidae. Insecta Mundi, 0353: 1-14. PDF
Cong, Q., Grishin, N. 2014. A new Hermeuptychia (Lepidoptera, Nymphalidae, Satyrinae) is sympatric and synchronic with H. sosybius in southeast US coastal plains, while another new Hermeuptychia species – not hermes – inhabits south Texas and northeast Mexico. Zookeys, 379: 43-91. PDF
Costa, M., Viloria, A. L., Attal, S., Neild, A. F. E., Fratello, S. A., Nakahara, S. 2016. Lepidoptera del Pantepui. Parte III. Nuevos Nymphalidae Cyrestinae y Satyrinae. Bulletin de la Société Entomologique de France, 121(2): 179-206.
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