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Published: 2020-09-29
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Revalidation of Vareuptychia Forster, 1964, description of Vanima gen. nov., and notes on Euptychia cleophes Godman & Salvin, 1889 (Lepidoptera: Nymphalidae: Satyrinae)

Departamento de Biologia Animal and Museu da Biodiversidade, Instituto de Biologia, Laboratório de Ecologia e Sistemática de Borboletas, Universidade Estadual de Campinas, Campinas, São Paulo, Brazil.
Department of Zoology, Laboratório de Estudos de Lepidoptera Neotropical, Universidade Federal do Paraná, Curitiba, Paraná, Brazil.
Department of Zoology, Laboratório de Estudos de Lepidoptera Neotropical, Universidade Federal do Paraná, Curitiba, Paraná, Brazil.
Department of Life Sciences, Natural History Museum, London, UK.
Research Museum Alexander Koenig, Bonn, Germany.
Departamento de Biologia Animal and Museu da Biodiversidade, Instituto de Biologia, Laboratório de Ecologia e Sistemática de Borboletas, Universidade Estadual de Campinas, Campinas, São Paulo, Brazil.
McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural History, University of Florida, Gainesville, Florida, USA.
McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural History, University of Florida, Gainesville, Florida, USA.
Departament of Entomology, Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Lima, Peru.
Lepidoptera Amazon rainforest Central America Cissia dry forests Euptychiina integrative taxonomy Megisto phylogeny

Abstract

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.

 

References

  1. Ackery, P.R. (1988) Host plants and classification: A review of nymphalid butterflies. Biological Journal of the Linnean Society, 33 (2), 95–203.

    https://doi.org/10.1111/j.1095-8312.1988.tb00446.x

    Andrade-C., M.G., Viloria, A.L., Henao-Bañol, E.R. & LeCrom, J.-F. (2019) Generic reassignments of satyrine butterflies from Colombia and Venezuela (Lepidoptera: Nymphalidae). Anartia, 28, 26–46.

    Aurivillius, P.O.C. (1929) Wissenschaftliche Ergebnisse der schwedischenentomologischen Reisen des Herrn Dr. A. Roman in Amazonas 1914–1915 und 1923–1924. 13. Rhopalocera. Entomologisk Tidskrift, 50 (3/4), 153–168.

    Austin, G.T., Haddad, N.M., Méndez, C., Sisk, T.D., Murphy, D.D., Launer, A.E. & Ehrlich, P.R. (1996) Annotated checklist of the butterflies of the Tikal National Park area of Guatemala. Tropical Lepidoptera, 7 (1), 21–37.

    Austin, G.T. & Mielke, O.H.H. (2008) Hesperiidae of Rondônia, Brazil: Porphyrogenes Watson (Lepidoptera: Pyrginae: Eudamini), with descriptions of new species from Central and South America. Insecta Mundi, 44, 1–56.

    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 & Evolution, 15 (3), 577–589.

    https://doi.org/10.1007/s13127-015-0221-y

    Barcant, M. (1970) Butterflies of Trinidad and Tobago. Collins, London, 314 pp.

    Beccaloni, G.W., Viloria, Á.L., Hall, S.K. & Robinson, G.S. (2008) Catalogue of the hostplants of the Neotropical butterflies. Monografias 3ercer Milenio, 8, 1–536.

    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.

    Benson, D.A., Cavanaugh, M., Clark, K., Karsch-Mizrachi, I., Lipman, D.J., Ostell, J. & Sayers, E.W. (2013) Genbank. Nucleic Acids Research, 41, 36–42.

    https://doi.org/10.1093/nar/gks1195

    Brévignon, C. (2005) Description de nouveau Satyrinae provenant de Guyane française (Lepidoptera, Nymphalidae). Lambillionea, 105 (3), (1), 393–404.

    Brévignon, C. (2008) Inventaire des Satyrinae de Guyane française (Lepidoptera: Nymphalidae). In: Lacomme, D. & Manil, L. (Eds.), Lépidoptères de Guyane. Tome 3. Rhopalocères 2. Association des Lépidoptéristes de France, Paris, pp. 62–94.

    Brévignon, C. & Benmesbah, M. (2012) Complément à l’inventaire des Satyrinae de Guyane (Lepidoptera: Nymphalidae). In: Lacomme, D. & Manil, L. (Eds.), Lépidoptères de Guyane. Tome 7. Nymphalidae Association des Lépidoptéristes de France, Paris, pp. 36–52.

    Brown, K.S., Freitas, A.V.L., von Schoultz, B., Saura, A.O. & Saura, A. (2007) Chromosomal evolution of South American frugivorous butterflies in the satyroid clade (Nymphalidae: Charaxinae, Morphinae and Satyrinae). Biological Journal of the Linnean Society, 92 (3), 467–481.

    https://doi.org/10.1111/j.1095-8312.2007.00872.x

    Bryk, F. (1953) Lepidoptera aus dem Amazonasgebiete und aus Peru gesammelt von Dr. Douglas Melin und Dr. Abraham Roman. Arkiv för Zoologi, New Series, 5 (1), 1–268.

    Butler, A.G. (1867a) A monograph of the genus Euptychia, a numerous race of butterflies belonging to the family Satyridae; with descriptions of sixty species new to science, and notes to their affinities, etc. Proceedings of the Zoological Society of London, 1866 (3), 458–504.

    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.

    Butler, A.G. (1868) Catalogue of diurnal Lepidoptera of the family Satyridae in the collection of the British Museum. Taylor and Francis, London, 146 pp.

    Butler, A.G. (1870) On new or recently described species of diurnal Lepidoptera. Entomologist’s Monthly Magazine, 6 (71), 250–252.

    Butler, A.G. (1871) Descriptions of five new species, and a new genus, of Diurnal Lepidoptera, from Shanghai. Transactions of the Entomological Society of London, 1871 (3), 401–403.

    Butler, A.G. (1877) On new species of the genus Euptychia, with a tabular view of those hitherto recorded. Journal of the Linnean Society of London, Zoology, 13 (67), 116–128.

    Casagrande, M.M., Mielke, O.H.H., Carneiro, E., Rafael, J.A. & Hutchings, R.W. (2012) Hesperioidea e Papilionoidea (Lepidoptera) coligidos em expedição aos Rios Nhamundá e Abacaxis, Amazonas, Brasil: novos subsídios para o conhecimento da biodiversidade da Amazônia Brasileira. Revista Brasileira de Entomologia, 56 (1), 23–28.

    https://doi.org/10.1590/S0085-56262012005000012

    Chernomor, O., von Haeseler, A. & Minh, B.Q. (2016) Terrace aware data structure for phylogenomic inference from supermatrices. Systematic Biology, 65, 997–1008.

    https://doi.org/10.1093/sysbio/syw037

    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.

    Comstock, J.H. (1918) The Wings of Insects. Comstock Publishing Company, New York, 430 pp.

    Comstock, J.H. & Needham, J.G. (1898–99) The wings of insects. American Naturalist, 32 (373, 374, 476, 377, 378, 380, 382, 384) & 33 (386, 391, 395), 43–48, 81–89, 231–340, 335–340, 413–424, 561–565, 769–777, 903–911 & 117–126, 573–582, 845–860.

    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.

    D’Abrera, B.L. (1988) Butterflies of the Neotropical Region—Part V. Nymphalidae (Concl.) & Satyridae. Black Rock, Hill House, Victoria, pp. 679–877.

    DeVries, P.J. (1987) The butterflies of Costa Rica and their Natural History. Vol.1. Princeton University Press, Princeton, 288 pp.

    Dias, M.M. (2011) Taxonomic considerations on the genera Moneuptychia Forster and Carminda Dias, reval. (Lepidoptera, Nymphalidae, Satyrinae). Brazilian Journal of Biology, 71 (2), 537–540.

    https://doi.org/10.1590/S1519-69842011000300026

    Distant, W.L. (1876) Remarks on the Rhopalocera of Costa Rica. Proceedings of the Entomological Society of London, 1876 (3), 10–14.

    Ehrlich, P.R., Sparrow, H.R., Sisk, T.D. & Daily, G.C. (1994) Notes on butterfly distributions in southern Costa Rica. Tropical Lepidoptera, 5 (1), 21–23.

    Espeland, M., Breinholt, J.W., Barbosa, E.P., Casagrande, M.M., Huertas, B., Lamas, G., Marín, M.A., Mielke, O.H.H., Miller, J.Y., Nakahara, S., Tan, D., Warren, A.D., Zacca, T., Kawahara, A.Y., Freitas, A.V.L. & Willmott, K.R. (2019) 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.

    https://doi.org/10.1016/j.ympev.2018.10.039

    Ferris, C.D. & Brown, F.M. (1981) Butterflies of the Rocky Mountain States. University of Oklahoma, Oklahoma, 442 pp.

    Figueroa-Fernandéz, A.L., Mélendez-Herrada, A., Luis-Martínez, A. & Vargas-Fernández, I. (2014) Diversity of diurnal butterflies (Lepidoptera: Hesperioidea and Papilionoidea) of Laguna Potosí and surrounding area, Guerrero, Mexico. Southwestern Entomologist, 39 (1), 57–75.

    https://doi.org/10.3958/059.039.0107

    Forster, W. (1964) Beiträge zur Kenntnis der Insektenfauna Boliviens XIX. Lepidoptera III. Satyridae. Veröffentlichungen der zoologischen Staatssammlung München, 8, 51–188, pls. 27–35.

    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, 30 (6), 661–668.

    https://doi.org/10.1590/S1984-46702013005000005

    Freitas, A.V.L., Barbosa, E.P., Siewert, R.R., Mielke, O.H.H., Zacca, T. & Azeredo-Espin, A.M. (2015) Four new species of Moneuptychia (Lepidoptera: Satyrinae: Euptychiina) from Brazil. Zootaxa, 3981 (4), 521–541.

    https://doi.org/10.11646/zootaxa.3981.4.4

    Freitas, A.V.L., Barbosa, E.P., 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, 60 (2), 157–165.

    https://doi.org/10.1016/j.rbe.2016.01.004

    Freitas, A.V.L., Barbosa, E.P., Zacca, T., Marín, M.A., Beirão, M.V., Silva, A.R.M., 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 (2), 148–158.

    https://doi.org/10.1016/j.rbe.2018.02.003

    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 (3), 254–261.

    https://doi.org/10.1016/j.rbe.2019.05.004

    Gaede, M. (1931) Familia Satyridae. Lepidopterum Catalogus, 46, 321–544.

    Garwood, K. & Jaramillo, J.G. (2016) Catálogo de mariposas Cañón del Río Claro, San Luis, Antioquia. Available from: http:// http://www.neotropicalbutterflies.com/colombiapdf/PDFs/PDF07_Catalogo_RioClaro_mar13_2016.pdf. (accessed 01 June 2016)

    Garwood, K., Geale, D. & Jaramillo, J.G. (2016) Catálogo de mariposas Parque Nacional Natural Tatamá, Carretera Montezuma. Available from: http://www.butterflycatalogs.com/uploads/1/0/3/2/103240120/aa1_cl_mogntezuma_117.pdf (accessed 1 June 2016)

    GIMP Team (2016) GIMP—GNU Image Manipulation Program. Available from: https://www.gimp.org (accessed 1 June 2016)

    Godman, F.D. & Salvin, O. (1880a) s.n. In: Biologia Centrali-Americana. Insecta. Lepidoptera-Rhopalocera. 1 (6). Dulau & Co., Bernard Quaritch, London, pp. 73–88, pl. 8.

    Godman, F.D. & Salvin, O. (1880b) A list of diurnal Lepidoptera collected in the Sierra Nevada de Santa Marta, Colombia, and the vicinity. Transactions of the Entomological Society of London, 1880 (3), 119–132, pls. 3–4.

    Godman, F.D. & Salvin, O. (1889) Descriptions of new species of Rhopalocera from Mexico and Central America. Annals and Magazine of Natural History, Series 6, 3 (16), 351–358.

    Guidon, S., Dufayard, J.-F., Lefort, V., Anisimova, M., Hordijk, W. & Gascuel, O. (2010) New algorithms and methods to estimate Maximum-Likelihood phylogenies: Assessing the performance of PhyML 3.0. Systematic Biology, 59, 307–321.

    https://doi.org/10.1093/sysbio/syq010

    Harvey, C.A., Villanueva, C., Villacís, J., Chacón, M., Muñoz, D., López, M., Ibrahim, M., Gómez, R., Taylor, R., Martínez, J., Navas, A., Sáenz, J., Sánchez, D., Medina, A., Vílchez, S., Hernández, B., Pérez, A., Ruiz, F., López, F., Lang, I. & Sinclair, F.L. (2005) Contribution of live fences to the ecological integrity of agricultural landscapes. Agriculture, Ecosystems and Environment, 111 (1/4), 200–230.

    https://doi.org/10.1016/j.agee.2005.06.011

    Henao-Bañol, E.R. & Gantiva-Q, C.H. (2020) Mariposas (Lepidoptera: Hesperioidea-Papilionoidea) del bosque seco tropical (BST) en Colombia. Conociendo la diversidad em um ecosistema amenazado. Boletín Científico del Museo de Historia Natural de la Universidade de Caldas, 24 (1), 150–196.

    Hernández-Mejía, C., Llorente-Bousquets, J., Vargas-Fernández, I. & Luis-Martínez, A. (2008) Las mariposas (Hesperioidea y Papilionoidea) de Malinalco, Estado de Mexico. Revista Mexicana de Biodiversidad, 79 (1), 117–130.

    https://doi.org/10.22201/ib.20078706e.2008.001.538

    Hoang, D.T., Chernomor, O., von Haeseler, A., Minh, B.Q. & Vinh, L.S. (2018) UFBoot2: Improving the ultrafast bootstrap approximation. Molecular Biology and Evolution, 35, 518–522.

    https://doi.org/10.1093/molbev/msx281

    Huertas, B. & Fagua, G. (2011) Distribution and conservation status of nominate Euptychia cesarense in the East Andes of Colombia. Zootaxa, 2906 (1), 66–68.

    https://doi.org/10.11646/zootaxa.2906.1.5

    Huertas, B., Lamas, G., Fagua, G., Mallet, J., Nakahara, S. & Willmott, K. (2016) A remarkable new butterfly species from western Amazonia (Lepidoptera: Nymphalidae: Satyrinae). Conservación Colombiana, 24, 5–11.

    Kalyaanamoorthy, S., Minh, B.Q., Wong, T.K.F., von Haeseler, A. & Jermiin, L.S. (2017) ModelFinder: Fast Model Selection for Accurate Phylogenetic Estimates. Nature Methods, 14, 587–589.

    https://doi.org/10.1038/nmeth.4285

    Kaye, W.J. (1921) A catalogue of the Trinidad Lepidoptera Rhopalocera. Memoirs of the Department of Agriculture of Trinidad and Tobago, 2, 13–163.

    Kirby, W.F. (1871) A Synonymic Catalogue of Diurnal Lepidoptera. John Van Voorst, London, 690 pp.

    Kirby, W.F. (1879) Catalogue of the collection of diurnal Lepidoptera formed by the late William Chapman Hewitson of Oatlands, Walton-on-Thames; and bequeathed by him to the British Museum. John Van Voorst, London, iv + 246 pp.

    Klots, A.B. (1970) Lepidoptera. In: Tuxen, S.L. (Ed.), Taxonomist’s Glossary of Genitalia in Insects. Munksgaard, Copenhagen, pp. 115–130.

    Lamas, G. (1981) La fauna de mariposas de la Reserva de Tambopata, Madre de Dios, Perú (Lepidoptera, Papilionoidea y Hesperioidea). Revista de la Sociedad Mexicana de Lepidopterología, 6 (2), 23–24.

    Lamas, G. (1994) Appendix 12. List of Butterflies from Tambopata Explorer Inn Reserve, 6. In: The Tambopata-Candamo Reserved Zone of Southeastern Peru, A biological assessment. Conservation International RAP Working Papers, 1994, pp. 162–184.

    Lamas, G. (2004) Nymphalidae. Satyrinae. Tribe Satyrini. Subtribe Euptychiina. In: Lamas, G. (Ed.), Checklist: Part 4A. Hesperioidea—Papilionoidea. In: Heppner, J.B. (Ed.), Atlas of Neotropical Lepidoptera. Vol. 5A. Association for Tropical Lepidoptera, Gainesville, pp. 217–223.

    Lewis, H.L. (1973) Butterflies of the World. Follett, Chicago, Illinois, 312 pp.

    Llorente-Bousquets, J., Garcés, A.R. & Luis, M.A. (1986) Las mariposas de Jalapa-Teocelo, Veracruz. Revista Teocelo, 4, 14–37.

    Llorente-Bousquets, J., Luis-Martínez, A., Vargas-Fernández, I. & Warren, A.D. (1996) Lista de las mariposas del Estado de Jalisco, Mexico. Revista de la Sociedad Mexicana de Historia Natural, 46, 35–48.

    Luis-Martínez, A., Hernández-Mejía, B., Trujano-Ortega, M., Warren, A., Salinas-Gutiérrez, J.L., Ávalos-Hernández, O., Vargas-Fernández, I. & Llorente-Bousquets, J. (2016) Avances faunísticos en los Papilionoidea (Lepidoptera) sensu lato de Oaxaca, México. Southwestern Entomologist, 41 (1), 171–224.

    https://doi.org/10.3958/059.041.0119

    Luis-Martínez, A., Llorente-Bousquets, J., Vargas-Fernández, I. & Warren, A.D. (2003) Biodiversity and biogeography of Mexican butterflies (Lepidoptera: Papilionoidea and Hesperioidea). Proceedings of the Entomological Society of Washington, 105 (1), 209–224.

    Luis-Martínez, A., Llorente-Bousquets, J., Vargas-Fernández, I. & Hernández-Baz, F. (2011) Apéndice VIII.25 Listado de especies de Papilionoidea y Hesperioidea del Estado de Veracruz. In: La Biodiversidad en Veracruz: Estudio de Estado. Comisión Nacional para el Conocimiento y Uso de La Biodiversidad, Gobierno del Estado de Veracruz, Universidad Veracruzana, Instituto de Ecología, A.C. México, pp. 1–29.

    Luis-Martínez, A., Llorente-Bousquets, J., Warren, A.D. & Vargas-Fernández,.I. (2004) Los lepidópteros: papilionoideos y hesperioideos. In: García-Mendoza, A.J., Ordóñez, M.J. & Briones-Salas, M.A. (Eds.), Biodiversidad de Oaxaca. Instituto de Biología, UNAM-Fondo Oaxaqueño para la Conservación de la Naturaleza-WWF, México, D.F, pp. 335–355.

    Luis-Martínez, A., Vargas-Fernández, I. & Llorente-Bousquets, J. (1996) Síntesis de los Papilionoidea (Lepidoptera: Rhopalocera) del Estado de Veracruz. Folia Entomológica Mexicana, 93, 91–133.

    Marín, M.A. & Uribe, S.I. (2009) Actualización sobre Euptychiina (Lepidoptera: Satyrinae) representadas en la colección del Museo Entomológico Francisco Luis Gallego. Boletín del Museo Entomológico Francisco Luis Gallego, 1 (2), 22–32.

    Marín, M.A., Peña, C., Freitas, A.V.L., Wahlberg, N. & Uribe, S.I. (2011) From the phylogeny of the Satyrinae butterflies to the systematics of Euptychiina (Lepidoptera: Nymphalidae): History, Progress and Prospects. Neotropical Entomology, 40 (1), 1–13.

    https://doi.org/10.1590/S1519-566X2011000100001

    Marín, 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 (4), 768–785.

    https://doi.org/10.1111/syen.12245

    Meerman, J.C. & Boomsma, T. (1993) Checklist of the butterflies of the Shipstern Nature Reserve. Occasional Papers of the Belize Natural History Society, 2 (3), 37–46.

    Michán, L., Llorente-Bousquets, J., Luis-Martínez, A. & Castro, D.J. (2005) Breve historia de la taxonomía de Lepidoptera en México durante el siglo XX. Revista de la Academia Colombiana de Ciencias, 29 (110), 101–132.

    Miller, L.D. (1968) The higher classification, phylogeny and zoogeography of the Satyridae (Lepidoptera). Memoirs of the American Entomological Society, 24, 1–174.

    Miller, L.D. (1974) Revision of the Euptychiini (Satyridae). 2. Cyllopsis R. Felder. Bulletin of the Allyn Museum, 20, 1–98.

    Miller, L.D. (1976) Revision of the Euptychiini (Satyridae). 3. Megisto Hübner. Bulletin of the Allyn Museum, 33, 1–23.

    Miller, J.Y., Matthews, D.L., Warren, A.D., Solis, M.A., Harvey, D.J., Gentili-Poole, P., Lehman, R., Emmel, T.C. & Covell, C.V. (2012) An annotated list of the Lepidoptera of Honduras. Insecta Mundi, 0205, 1–72.

    Montero-Abril, F., Moreno-P., M. & Gutiérrez-M., L.C. (2009) Mariposas (Lepidoptera: Hesperioidea y Papilionoidea) asociadas a fragmentos de bosque seco tropical en el departamento del Atlántico, Colombia. Boletín Científico del Museo de Historia Natural de la Universidad de Caldas, 13 (2), 157–173.

    Montero-Abril, F. & Ortiz-Pérez, M. (2013) Estados inmaduros e historia natural de algunas especies de la subtribu Pronophilina (Nymphalidae: Satyrinae) presentes em el Páramo del Tablazo, Colombia III. Neopedaliodes zipa (Adams, 1986). Tropical Lepidoptera Research, 23 (1), 54–61.

    Murray, D.L. & Prowell, D.P. (2005) Molecular phylogenetics and evolutionary history of the neotropical satyrine subtribe Euptychiina (Nymphalidae: Satyrinae). Molecular Phylogenetics and Evolution, 34 (1), 67–80.

    https://doi.org/10.1016/j.ympev.2004.08.014

    Nakahara, S., Marín, M.A. & Mitsuhashi, W. (2012) First records for Cissia themis (Lepidoptera: Nymphalidae) from Colombia and Venezuela. Revista Colombiana de Entomologia, 38 (2), 359–362.

    Nakahara, S., Tan, D., Lamas, G., Parus, A. & Willmott, K.R. (2016a) A distinctive new species of Hermeuptychia Forster, 1964 from the eastern tropical Andes (Lepidoptera: Nymphalidae: Satyrinae). Tropical Lepidoptera Research, 26 (2), 77–84.

    Nakahara, S., Barbosa, E.P., Marín, M.A., Freitas, A.V.L. & Willmott, K.R. (2016b) Graphita gen. nov., a new genus for Neonympha griphe C. Felder & R. Felder, 1867 (Lepidoptera, Nymphalidae, Satyrinae). Neotropical Entomology, 45 (6), 675–691.

    https://doi.org/10.1007/s13744-016-0414-x

    Nakahara, S., Zacca, T., Huertas, B., Neild, A.F.E., Hall, J.P.W., Lamas, G., Holian, L.A., Espeland, M. & Willmott, K.R. (2018a) 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 & Evolution, 49 (2), 130–182.

    https://doi.org/10.1163/1876312X-00002167

    Nakahara, S., Willmott, K.R., Mielke, O.H.H., Schwartz, J., Zacca, T., Espeland, M. & Lamas, G. (2018b) Seven new taxa from the butterfly subtribe Euptychiina (Lepidoptera: Nymphalidae: Satyrinae) with revisional notes on Harjesia Forster, 1964 and Pseudeuptychia Forster, 1964. Insecta Mundi, 639, 1–38.

    Nakahara, S., Lamas, G., Tyler, S., Marín, M.A., 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.

    https://doi.org/10.3897/zookeys.821.31782

    Neild, A.F.E. (1996) The butterflies of Venezuela. Part 1: Nymphalidae I (Limenitidinae, Apaturinae, Charaxinae). A comprehensive guide to the identification of adult Nymphalidae, Papilionidae, and Pieridae. Meridian Publications, Greenwich, London, 144 pp.

    Nguyen, L.-T., Schmidt, H.A., von Haeseler, A. & Minh, B.Q. (2015) IQ-TREE: A fast and effective stochastic algorithm for estimating maximum likelihood phylogenies. Molecular Biology and Evolution, 32, 268–274.

    https://doi.org/10.1093/molbev/msu300

    Peña, C., Nylin, S., Freitas, A.V.L. & Wahlberg, N. (2010) Biogeographic history of the butterfly subtribe Euptychiina (Lepidoptera, Nymphalidae, Satyrinae). Zoologica Scripta, 39 (3), 243–258.

    https://doi.org/10.1111/j.1463-6409.2010.00421.x

    Peña, C., Wahlberg, N., Weingartner, E., Kodandaramaiah, U., Nylin, S., Freitas, A.V.L. & Brower, A.V.Z. (2006) Higher level phylogeny of Satyrinae butterflies (Lepidoptera: Nymphalidae) based on DNA sequence data. Molecular Phylogenetics and Evolution, 40 (1), 29–49.

    https://doi.org/10.1016/j.ympev.2006.02.007

    Petersen, W. (1904) Die Morphologie der Generationsorgane der Schmetterlinge und ihre Bedeutung für die Artbildung. Mémoires de l’Académie Impériale des Sciences de St. Pétersbourg, Series 8, 16 (8), 1–84.

    Pierce, F.N. (1914) The genitalia of the group Geometridae of the Lepidoptera of the Britsh Islands. The Elms, Dingle, Liverpool, 230 pp.

    Pozo, C., Luis-Martínez, A., Llorente-Bousquet, J., Salas-Suárez, N.S., Maya-Martínez, A., Vargas-Fernández, I. & Warren, A.D. (2008) Seasonality and phenology of the butterflies (Lepidoptera: Papilionoidea and Hesperioidea) of Mexico’s Calakmul region. Florida Entomologist, 91 (3), 407–422.

    https://doi.org/10.1653/0015-4040(2008)91[407:SAPOTB]2.0.CO;2

    Pozo, C., Luis-Martínez, A., Tescum, S., Salas-Suárez, N. & Maya-Martínez, A. (2003) Butterflies (Papilionoidea and Hesperioidea) of Calakmul, Campeche, México. Southwestern Naturalist, 48 (4), 505–525.

    https://doi.org/10.1894/0038-4909(2003)048<0505:BPAHOC>2.0.CO;2

    Pulido, H.W., Andrade, M.G., Peña, C. & Lamas, G. (2011) Two new taxa of Euptychia Hübner, 1818 (Lepidoptera: Nymphalidae: Satyrinae). Zootaxa, 2906 (1), 43–51.

    https://doi.org/10.11646/zootaxa.2906.1.2

    QGIS Development Team (2016) QGIS Geographic Information System. Open Source Geospatial Foundation Project. Available from: http.//qgis.osgeo.org (accessed 1 June 2016)

    Racheli, T. & Racheli, L. (1998) Lepidoptera diversity of an Ecuadorian lowland rain forest (Lepidoptera: Papilionidae, Pieridae, Nymphalidae, Saturniidae, Sphingidae). Neue Entomologische Nachrichten, 41, 95–117.

    Raguso, R.A. & Llorente-Bousquets, J. (1990) The butterflies (Lepidoptera) of the Tuxtlas Mts., Veracruz, Mexico, revisited: species-richness and habitat disturbance. Journal of Research on the Lepidoptera, 29 (1/2), 105–133.

    Ramos, F.A. (2000) Nymphalid butterfly communities in an Amazonian forest fragment. Journal of Research on the Lepidoptera, 35, 29–41.

    Riley, N.D. & Gabriel, A.G. (1924) Catalogue of the type specimens of Lepidoptera Rhopalocera in the British Museum. Part I. Satyridae. Oxford University Press, London, 62 pp.

    Robbins, R.K., Lamas, G., Mielke, O.H.H., Harvey, D.J. & Casagrande, M.M. (1996) Taxonomic composition and ecological structure of the species-rich butterfly community at Pakitza, Parque Nacional del Manu, Perú. In: Wilson, D.E. & Sandoval, A. (Eds.), Manu. The biodiversity of southeastern Peru. Smithsonian Institution, Washington, D.C., pp. 217–252.

    Santin, A. (2004) Répertoire des plantes-hôtes et de substitution des chenilles de Lépidoptères du monde. Vol. 1. Office pour la Information Eco-entomologique, Guyancourt, 1200 pp.

    Shuey, J.A., Giles, V., Meerman, J., Labus, P., Schutte, C.W. & Kovarik, P. (2005) New additions to the butterfly fauna of Belize. Journal of the Lepidopterists’ Society, 59 (2), 83–88.

    Siewert, R.R., Zacca, T., Dias, F.M.S., Freitas, A.V.L., Mielke, O.H.H. & Casagrande, M.M. (2013) The “Taygetis yphthima species group” (Lepidoptera, Nymphalidae, Satyrinae): taxonomy, variation and description of a new species. Zookeys, 356, 11–29.

    https://doi.org/10.3897/zookeys.356.6481

    Silva-Brandão, K.L., Freitas, A.V.L., Brower, A.V.Z. & Solferini, V.N. (2005) Phylogenetic relationships of the New World Troidini swallowtails (Lepidoptera:Papilionidae) based on COI, COII, and EF-1 genes. Molecular Phylogenetics and Evolution, 36 (3), 468–483.

    https://doi.org/10.1016/j.ympev.2005.04.007

    Singer, M.C., DeVries, P.J. & Ehrlich, P.R. (1983) The Cissia confusa species-group in Costa Rica and Trinidad (Lepidoptera: Satyrinae). Zoological Journal of the Linnean Society, 79 (2), 101–119.

    https://doi.org/10.1111/j.1096-3642.1983.tb01162.x

    Singer, M.C. & Ehrlich, P.R. (1991) Host specialization of satyrine butterflies, and their responses to habitat fragmentation in Trinidad. Journal of Research on the Lepidoptera, 30 (3/4), 248–256.

    Staudinger, O. (1884–1888) Exotische Tagfalter in systematischer Reihenfolge mit Berücksichtigung neuer Arten. Vol. 1. In: Staudinger, O. & Schatz, E. (Eds.), Exotische Schmetterlinge. G. Löwensohn, Fürth, pp. 1–333, 100 pls.

    Tietz, H.M. (1972) An index of the described life-histories, early stages and hosts of the Macrolepidoptera of continental United States and Canada. Vol. 2. A. C. Allyn, Sarasota, Florida, 1041 pp.

    Viloria, Á.L. & Luis-Martínez, A. (2019) Llorenteana, a new butterfly genus from the American Continent (Lepidoptera: Nymphalidae: Satyrinae). Dugesiana, 26 (2), 167–172.

    Wahlberg, N., Leneveu, J., Kodandaramaiah, U., Peña, C., Nylin, S., Freitas, A.V.L. & Brower, A.V.Z. (2009) Nymphalid butterflies diversity following near demise at the Cretaceous/Tertiary boundary. Proceedings of the Royal Society of London, Series B: Biological Sciences, 276 (1677), 4295–4302.

    https://doi.org/10.1098/rspb.2009.1303

    Wahlberg, N. & Wheat, C.W. (2008) Genomic outposts serve the phylogenomicpioneers: designing novel nuclear markers for genomic DNA extractions of Lepidoptera. Systematics Biology, 57 (2), 231–242.

    https://doi.org/10.1080/10635150802033006

    Wesley, D.J. & Emmel, T.C. (1975) The chromosomes of Neotropical butterflies from Trinidad and Tobago. Biotropica, 7 (1), 24–31.

    https://doi.org/10.2307/2989796

    Weymer, G. (1910–1913) 4. Familie: Satyridae. In: Seitz, A. (Ed.), Die Gross-Schmetterlinge der Erde. 5. A. Kernen, Stuttgart, pp. 173–283.

    Willmott, K.R., Marín, M.A., Nakahara, S., Pomerantz, T., Lamas, G., Huertas, B., Espeland, M., Xiao, L., Hall, J.P.W., Willmott, J.I.R. & 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.

    Zacca, T., Barbosa, E.P. & Freitas, A.V.L. (2018a) Checklist Euptychiina. Available from: https://www2.ib.unicamp.br/labor/site/?page_id=1020 (accessed 1 August 2020).

    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 & Evolution. [published online]

    https://doi.org/10.1163/1876312X-00001027

    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. (2018b) Systematics of the butterfly genus Cissia Doubleday, 1848 (Lepidoptera: Nymphalidae, Satyrinae) using an integrative approach. Arthropod Systematics and Phylogeny, 76 (2), 349–376.

    Zacca, T., Paluch, M., Siewert, R.R., Freitas, A., Barbosa, E., Mielke, O. & Casagrande, M. (2017) Revision of Godartiana Forster (Lepidoptera: Nymphalidae), with the description of a new species from northeastern Brazil. Austral Entomology, 56 (2), 169–190.

    https://doi.org/10.1111/aen.12223

    Zimsen, E. (1964) The Type Material of I. C. Fabricius. Munksgaard, Copenhagen, 656 pp.