Abstract
Opoptera Aurivillius, 1882 (Lepidoptera: Nymphalidae, Brassolini) is a Neotropical genus, with species occurring from Mexico to Argentina. Although previous phylogenetic hypotheses involving Opoptera species have been proposed, an investigation of the species relationships based on molecular data, supported by a systematic review, has not been carried out to date. This study presents a taxonomic revision of Opoptera, which currently comprises eight species and two subspecies, proposing a hypothesis of relationships among the taxa based on DNA, morphology, geographic distribution, and natural history data. To carry out this study, 530 specimens of all species of the genus were examined, including type material of all proposed taxa. For the phylogenic hypothesis, the maximum likelihood method was used, and five molecular markers: one mitochondrial, cytochrome c oxidase subunit I (COI), and four nuclear, elongation factor 1-alpha (EF-1α), ribosomal protein S5 (RpS5), wingless (wg), and glyceraldehyde-3-phosphate dehydrogenase (GAPDH). Opoptera was recovered as monophyletic and the relationships between species corroborate the previous hypotheses. From the taxonomic review, two synonyms are proposed: Opoptera hilaris colombiana (Rothschild, 1916) syn. nov. and Opoptera hilaris fuscata Stichel, 1908 syn. nov. of Opoptera hilaris Stichel, 1901. A new status is proposed: Opoptera mexicana J. Maza & R. G. Maza, 1989 stat. nov. A new species is described: Opoptera iracildae sp. nov. In order to stabilize the nomenclature of the group, three neotypes are designated for Caligo syme Hübner, [1821], Opsiphanes arsippe Hopffer, 1874, and Opoptera aorsa hilaris Stichel, 1901; and ten lectotypes for Morpho aorsa Godart, [1824], Opoptera aorsa litura Fruhstorfer, 1907, Opoptera arsippe bracteolata Stichel, 1901, Opoptera aorsa fuscata Stichel, 1908, Opsiphanes sulcius Staudinger, 1887, Opsiphanes sulcius var. sodalis Röber, 1906, Papilio alcimedon Dalman, 1823, Morpho acadina Godart, [1824], Opoptera syme fumosa Stichel, 1902, and Opsiphanes staudingeri Godman & Salvin, 1894. This study recognizes ten valid species and eight synonyms.
References
- Aguiar, T.M.C., Gueratto, P.E., Machado, P.A., Santos, J.P., Carreira, J.Y.O., Moraes, S.S. & Freitas, A.V.L. (2024) Immature Stages, Natural History and Population Biology of Opoptera syme (Hübner, 1821), (Lepidoptera: Nymphalidae: Satyrinae) from Southeastern Brazil. Neotropical Entomology, 53 (1), 101–109. https://doi.org/10.1007/s13744-023-01093-7
- Aurivillius, P.O.C. (1882) Recensio critica Lepidopterorum Musei Ludovicae Ulricae quae descripsit Carolus a Linné. Kongliga svenska Vetenskaps-Akademiens Handlingar, Ny Följd, 19 (5), 1–188. https://doi.org/10.5962/bhl.title.67730
- Benson, D.A., Karsch-Mizrachi, I., Lipman, J., Ostell, J. & Wheeler, D. (2005) GenBank. Nucleic Acids Research, 33, D34–D38. https://doi.org/10.1093/nar/gki063
- Biedermann, W. (1928) Descriptions et note diverses. Encyclopédie entomologique, Series B, 3 (1), 1–3, pls. 1–3.
- Boisduval, J.B.A.D. (1836) Suites à Buffon. Histoire naturelle des Insectes. Species général des Lépidoptères. Librairie Encyclopédique de Roret, Suites à Buffon, Paris, 690 pp., 24 pls.
- Bristow, C.R. (1991) A revision of the brassoline genus Opsiphanes (Lepidoptera: Rhopalocera). Zoological Journal of the linnean society, 101 (3), 203–293. https://doi.org/10.1111/j.1096-3642.1991.tb00282.x
- Burmeister, H.C.C. (1873) Description de Morphonides brésiliens. Revue et Magasin de Zoologie, Series 3, 1 (1), 17–47.
- Casagrande, M.M. (1979a) Sobre Caligo beltrao (Illiger). III. Morfologia externa do adulto—Tórax (Lepidoptera, Satyridae, Brassolinae). Revista brasileira de Biologia, 39 (2), 347–355.
- Casagrande, M.M. (1979b) Sobre Caligo beltrao (Illiger). III [sic]. Morfologia externa do adulto—abdôme (Lepidoptera, Satyridae, Brassolinae). Revista brasileira de Biologia, 39 (3), 711–716.
- Casagrande, M.M. (1982) Quatro gêneros novos de Brassolinae (Lepidoptera, Nymphalidae). Revista brasileira de Entomologia, 26 (3/4), 355–356.
- Casagrande, M.M. (1996) Notas sistemáticas sobre Brassolinae. I. Tribos (Lepidoptera, Nymphalidae). Revista brasileira de Zoologia, 12 (3), 671–699. https://doi.org/10.1590/S0101-81751995000300023
- Casagrande, M.M. (2002) Naropini Stichel, taxonomia e imaturos (Lepidoptera, Nymphalidae, Brassolinae). Revista brasileira de Zoologia, 19 (2), 467–569. https://doi.org/10.1590/S0101-81752002000200012
- Casagrande, M.M. (2004) Tribe Brassolini. In: Lamas, G. (Ed.), Checklist: Part 4A, Hesperioidea—Papilionoidea. In: Heppner, J.B. (Ed.), Atlas neotroprical lepidoptera 5A. Association for Tropical Lepidoptera, Scientific Publishers, Gainesville, Florida, pp. 201–205.
- Comstock, J.H. (1918) The wings of insects: an exposition of the uniform terminology of the wing-veins of insects and a discussion of the more general characteristics of the wings of the several orders of insects. Comstock Publishing Company, Ithaca, New York, 470 pp. https://doi.org/10.5962/bhl.title.54605
- Comstock, J.H. & Needham, J.G. (1898) The wings of insects. Chapter III. The specialization of wings by reduction. The american Naturalist, 32 (376), 231–257. https://doi.org/10.1086/276835
- Comstock, J.H. & Needham, J.G. (1899) The wings of insects. Chapter V. The development ow wings. The american Naturalist 33 (395), 845–860. https://doi.org/10.1086/277462
- Cramer, P. (1777) De uitlandsche Kapellen voorkomende in de drie Waereld-Deelen Asia, Africa en America. Papillons exotiques des trois parties du monde l’Asie, l’Afrique et l’Amérique. Vol. 2. S. J. Baalde, Amsteldam and Barthelemy Wild, Utrecht, 151 pp., pls. 97–192.
- Dalman, J.W. (1823) Analecta entomologica. Lindhian, Holmiae, [2] + vii + 104 + [4] pp., 4 pls. https://doi.org/10.5962/bhl.title.66069
- Doubleday, E. ([1845]) List of the specimens of lepidopterous insects in the collection of the British Museum. Part I. Edward Newman, London, v + 150 pp.
- Doubleday, E. ([1848–1849]) The genera of diurnal Lepidoptera: comprising their generic characters, a notice of their habits and transformations, and a catalogue of the species of each genus. Vol. 2. Longman, Brown, Green, and Longmans, London, pls. 31–52 + 56–58 + 60–62 + 64.
- DeVries, P.J. (1987) The Butterflies of Costa Rica and their Natural History: Papilionidae, Pieridae, Nymphalidae. Princeton University Press, Princeton, New Jersey, 327 pp.
- Eltringham, H. (1929) On the scent organs of Opsiphanes cassiae lucullus, Fruhst. (Lepidoptera, Brassolidae). Transactions of the Royal entomological society of London, 77 (1), 1–4., pl. 1. https://doi.org/10.1111/j.1365-2311.1929.tb00675.x
- Fruhstorfer, H. (1907) Verzeichnis der von Herrn Dr. Theodor Koch-Grünberg am oberen Waupes 1903–1905 gesammelten Rhopaloceren mit Besprechung verwandter Arten. Stettiner Entomologische Zeitung, 68 (1), 117–164.
- Fruhstorfer, H. (1911-12) 5. Familie: Brassolidae. In: Seitz, A. (Ed.), Die Gross-Schmetterlinge der Erde. Vol. 5. Alfred Kernen, Stuttgart, pp. 285–332, pls. 61–67.
- Glaser, L. (1887) Catalogus etymologicus Coleopterorum et Lepidopterorum. Erklärendes und verdeutschendes Namenverzeichniss der Käfer und Schmetterlinge für Liebhaber und wissenschaftliche Sammler systematisch und alphabetisch zusammengestellt. R. Friedländer & Sohn, Berlin, iv + 396 pp.
- Godart, J.B. ([1824]) Papillons. In: Godart, J.B. & Latreille, P. (Eds.), Encyclopédie Méthodique. Histoire naturelle. Entomologie, ou histoire naturelle dês crustacés, dês arachnides et dês insectes. Vol. 9 (2). Veuve Agasse, Paris, pp. 329–828.
- Godman, F.D. & Salvin, O. (1894) Descriptions of new species of Rhopalocera from Mexico and Central America. Annals and Magazine of natural History, Series 6, 14 (80), 95–98. https://doi.org/10.1080/00222939408677774
- Godman, F.D. (1901) Biologia Centrali Americana. Vol. II. Dulau and Co., Bernard Quaritch, London, 2 pp. [pp. 664–665]
- Griffith, E. & Pidgeon, W. (1832) The Class Insecta arranged by the Baron Cuvier, with supplementary additions to each order. In: Griffith, E. (Ed.), The Animal Kingdom arranged in conformity with its organization, by the Baron Cuvier, member of the Institute of France, &c. &c. &c. with supplementary additions to each Order. Vol. 15. Whittaker, Treacher, and Co., London, pp. 1–796.
- Guérin-Méneville, F.E. (1832) Iconographie du Règne Animal de G. Cuvier, ou représentation d’après nature de l’une des espèces les plus rémarquables et souvent non encore figurées de chaque genre d’animaux. Avec un texte descriptif mis au courant de la science. Ouvrage pouvant servir d’atlas a tous les traités de zoologie. Insectes. J. B. Baillière, Paris, 576 pp.
- Guindon, 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 (3), 307–321. https://doi.org/10.1093/sysbio/syq010
- Hebert, P.D.N., Cywinska, A., Ball, S.L. & deWaard, J.R. (2003) Biological identifications through DNA barcodes. Proceedings of the Royal Society of London, Series B, 270 (1512), 313–321. https://doi.org/10.1098/rspb.2002.2218
- Hemming, F. (1967) The generic names of the butterflies and their type-species (Lepidoptera: Rhopalocera). Bulletin of the British Museum Natural History, Entomology, Supplement 9, 1–509. https://doi.org/10.5962/p.78596
- Hernández, B., Lapeze, J. & Maes, J.M. (2018) Opoptera staudingeri (Godman & Salvin, 1894) (Lepidoptera: Nymphalidae) confirmación para la fauna de Nicaragua. Revista nicaragüense de Entomología, 152, 1–10.
- Hernández-Roldán, J.L., Bofill, R., Dapporto, L., Munguira, M.L. & Vila, R. (2014) Morphological and chemical analysis of male scent organ in the butterfly genus Pyrgus (Lepidoptera: Hesperiidae). Organisms Diversity & Evolution, 14 (3), 269–278. https://doi.org/10.1007/s13127-014-0170-x
- Herrich-Schäffer, G.A.W. (1865) Lepidopterorum Index systematicus. Correspondenz-Blatt des zoologisch-mineralogischen Vereines in Regensburg, 19 (5), 63–76.
- Hoffmann, F. (1937) Beiträge zur Naturgeschichte brasilianischer Schmetterlinge. II. Internationale entomologische Zeitschrift, 51 (5), 49–52.
- Hopffer, C.H. (1874) Neue Lepidopteren von Peru und Bolivia. Stettiner entomologische Zeitung, 35 (10/12), 329–371.
- Hübner, J. ([1821]) Sammlung exotischer Schmetterlinge, Vol. II, Jacob Hübner, Augsburg, pls. 70–71.
- Katoh, K. & Standley, D.M. (2013) MAFFT Multiple Sequence Alignment Software Version 7: Improvements in Performance and Usability. Molecular Biology and Evolution, 30 (4), 772–780. https://doi.org/10.1093/molbev/mst010
- Kirby, W.F. (1871) A Synonymic Catalogue of Diurnal Lepidoptera. John Van Voorst, London, vii + 690 pp. https://doi.org/10.5962/bhl.title.23779
- Kirby, W.F. (1877) A Synonymic Catalogue of Diurnal Lepidoptera, Supplement, John Van Voorst, London, viii + 193 pp. [pp. i–viii, 691–883] https://doi.org/10.5962/bhl.title.11413
- 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.
- Köhler, P.E. (1923) Fauna Argentina. Lepidoptera e collectione Alberto Breyer. I. Teil. Rhopalocera. Systematischer Katalog und Studien, Berichtigungen u. Neubeschreibungen. Zeitschrift für wissenschaftliche Insektenbiologie, 18 (12), Sonderbeilage, 1–34, 3 pls.
- Kumar, S., Stecher, G., Li, M., Knyaz, C., Tamura, K. (2018) MEGA X: Molecular Evolutionary Genetics Analysis across Computing Platforms. Molecular Biology and Evolution, 35 (6), 1547–1549. https://doi.org/10.1093/molbev/msy096
- Lamas, G. (2004) Checklist: Part 4A. Hesperioidea-Papilionoidea. Vol. 5A. Association for Tropical Lepidoptera ⁄Scientific Publishers, Gainesville, Florida, xxxvi + 439 pp.
- Le Cerf, F.L. (1926) Contribution à l’étude des organes sensoriels des lépidoptères. Encyclopédie Entomologique, Series B, 1 (3), 133–158.
- Lucas, P.H. (1851) Papillons. In: Chenu, J.C. (Ed.), Encyclopédie d’Histoire Naturelle ou traité complet de cette Science, d’après les travaux des naturalistes les plus éminents de tous les pays et de toutes les époques. Buffon, Daubenton, Lacépède, G. Cuvier, F. Cuvier, Geoffroy Saint-Hilaire, Latreille, de Jussieu, Brongniart, etc., etc. Ouvrage résumant les observations des auteurs anciens et comprenant toutes les découvertes modernes jusqu’à nos jours. Vol. 1. Marescq et Compagnie; Gustave Havard, Paris, 310 pp., 40 pls.
- Mabilde, A. (1896) Borboletas do Estado do Rio Grande do Sul. Guia Practica para os principiantes colleccionadores de insectos contendo a descripção fiel de perto de l000 borboletas com 280 figuras lythographadas em tamanho, formas e dezenhos conforme o natural. Estudo sobre a vida de insectos do Rio Grande do Sul e sobre a caça, classificação e a conservação de uma collecção, mais ou menos regular. Typographia Gundlach 8 Schuldt, Pôrto Alegre, 240 pp., 24 pls.
- Matos-Maraví, P., Wahlberg, N., Freitas, A.V.L., DeVries, P., Antonelli, A. & Penz, C.M. (2021) Mesoamerica is a cradle and the Atlantic Forest is a museum of Neotropical butterfly diversity: insights from the evolution and biogeography of Brassolini (Lepidoptera: Nymphalidae). Biological Journal of the Linnean Society, 133 (3), 704–724. https://doi.org/10.1093/biolinnean/blab034
- Maza-Elvira., J. de la & Maza-Elvira, R.G. de la (1989) Una nueva subespecie de Opoptera staudingeri (G. y S. 1894) de Mexico (Nymphalidae: Brassolinae). Revista de la Sociedad mexicana de Lepidopterología, 13 (1), 25–28.
- Ménétriés, E. (1855) Enumeratio corporum animalium Musei Imperialis Academiae Scientiarum Petropolitanae. Classe insectorum, Ordo lepidopterorum. Pars I. Lepidoptera Diurna. Typis Academiae Scientiarum Imperialis, Petropoli, xv + 99 pp. ], 6 pls. [pp. i–xv + 1–66 + 67–97 + (2) pp., pls. 1–6] https://doi.org/10.5962/bhl.title.12490
- Minh, B.Q., Lanfear, R., Trifinopoulos, J., Schrempf, D. & Schmidt, H.A. (2021) IQ-TREE Version 2.1.2. Tutorials and Manual Phylogenomic software by maximum likelihood. Available from: http://www.iqtree.org/doc/iqtree-doc.pdf (accessed 8 February 2023)
- Minh, B.Q., Nguyen, M.A.T. & von Haeseler, A. (2013) Ultrafast Approximation for Phylogenetic Bootstrap. Molecular Biology and Evolution, 30 (5), 1188–1195. https://doi.org/10.1093/molbev/mst024
- Miranda-Ribeiro, V. (1931) Lepidopteros de Matto Grosso. Boletim do Museu Nacional, 7 (1), 31–52.
- 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 (1), 268–274. https://doi.org/10.1093/molbev/msu300
- Nijhout, H.F. (1991) The development and evolution of butterfly wing patterns. Smithsonian Institution Press, Washington, D.C., xvi + 297 pp., 8 pls.
- Nijhout, H.F. (2017) The Common Developmental Origin of Eyespots and Parafocal Elements and a New Model Mechanism for Color Pattern Formation. In: Sekimura, T. & Nijhout, H.F. (Eds.), Diversity and evolution of Butterfly Wing Patterns. An integrative approach. Springer Nature Singapore, Singapore, pp. 3–19. https://doi.org/10.1007/978-981-10-4956-9_1
- Orlandin, E., Piovesan, M. & Carneiro, E. (2020) Borboletas do Meio—Oeste de santa Catarina: História natural e Guia de Identificação. Authors, Joaçaba, 398 pp.
- Penz, C.M. (2007) Evaluating the monophyly and phylogenetic relationships of Brassolini genera (Lepidoptera, Nymphalidae). Systematic Entomology, 32 (4), 668–689. https://doi.org/10.1111/j.1365-3113.2007.00391.x
- Penz, C.M. (2009a) The phylogeny of Opoptera butterflies, and an assessment of the systematic position of O. staudingeri (Lepidoptera, Nymphalidae). Zootaxa, 1985 (1), 1–20. https://doi.org/10.11646/zootaxa.1985.1.1
- Penz, C.M. (2009b) Phylogeny of Dasyophthalma butterflies (Lepidoptera, Nymphalidae, Brassolini). Insecta Mundi, 69, 1–12. https://doi.org/10.5281/zenodo.5405005
- Penz, C.M., Casagrande, M.M., DeVries, P.J. & Simonsen, T.J. (2017) Documenting diversity in the Amazonian butterfly genus Bia (Lepidoptera, Nymphalidae). Zootaxa, 4258 (3), 201–237. https://doi.org/10.11646/zootaxa.4258.3.1
- Penz, C.M., Freitas, A., Kaminski, L., Casagrande, M.M. & DeVries, P. (2013) Adult and early‐stage characters of Brassolini contain conflicting phylogenetic signal (Lepidoptera, Nymphalidae). Systematic Entomology, 38 (2), 316–333. https://doi.org/10.1111/syen.12000
- Penz, C.M. & Heine, K.B. (2016) Did adult diurnal activity influence the evolution of wing morphology in Opoptera butterflies? Neotropical Entomology, 45 (1), 50–57. https://doi.org/10.1007/s13744-015-0338-x
- Penz & Mohammadi, N. (2013). Wing pattern diversity in Brassolini butterflies (Nymphalidae, Satyrinae). Biota neotropica, 13 (3), 154–180. https://doi.org/10.1590/S1676-06032013000300020
- Penz, C.M., Mohammadi, N. & Wahlberg, N. (2011) Neotropical Blepolenis butterflies: wing pattern elements, phylogeny, and Pleistocene diversification (Lepidoptera, Nymphalidae). Zootaxa, 2897 (1), 1–17. https://doi.org/10.11646/zootaxa.2897.1.1
- Piovesan, M., Casagrande, M.M. & Mielke, O.H.H. (2022) Systematics of Opsiphanes Doubleday, [1849] (Lepidoptera: Nymphalidae, Satyrinae, Brassolini): an integrative approach. Zootaxa, 5216 (1), 1–278. https://doi.org/10.11646/zootaxa.5216.1.1
- Prittwitz, O.F.W.L. (1865) Beitrag zur Fauna des Corcovado. Stettiner entomologische Zeitung, 26 (4/5), 123–143.
- QGIS Development Team (2023). QGIS Geographic Information System. Open Source Geospatial Foundation Project. Disponível em: http://qgis.osgeo.org (accessed 11 July 2023)
- Rambaut, A. (2018) Fig Tree. Version 1.4.4. Available from: http://tree.bio.ed.ac.uk/software/figtree/ (accessed 8 February 2023)
- Ratnasingham, S. & Hebert, P.D.N. (2007) BARCODING: bold: The Barcode of Life Data System (http://www.barcodinglife. org). Molecular Ecology Notes, 7 (3), 355–364. https://doi.org/10.1111/j.1471-8286.2007.01678.x
- Röber, J.K.M. (1896) Opsiphanes fruhstorferi n. sp. Entomologische Nachrichten, 22 (21), 323–324. https://doi.org/10.5962/bhl.part.1950
- Röber, J.K.M. (1906) Neue Schmetterlinge. Societas entomologica, 20 (23), 177–178.
- Rothschild, L.W. (1916) Notes on Amathusiidae, Brassolidae, Morphidae, etc., with descriptions of new forms. Novitates zoologicae, 23 (3), 299–318, pls. 3–6.
- Schneider, C.A., Rasband, W.S. & Eliceiri, K.W. (2012) NIH Image to ImageJ: 25 years of image analysis. Nature Methods, 9 (7), 671–675. https://doi.org/10.1038/nmeth.2089
- Schulz, S., Francke, W. & Boppré, M. (1988) Carboxylic acids from hairpencil of male Amauris butterflies (Lep.: Danainae). Biological Chemistry, 369 (2), 633–638. https://doi.org/10.1515/bchm3.1988.369.2.633
- Scoble, M.J. (1992) Hearing, sound, and scent. In: The Lepidoptera: form, function, and diversity. Oxford University Press, Oxford, pp. 160–169.
- Schwanwitsch, B.N. (1924) On the groundplan of wing-pattern in nymphalids and certain other families of Rhopalocerous Lepidoptera. Proceedings of the Zoological Society of London, 34 (2), 509–528. https://doi.org/10.1111/j.1096-3642.1924.tb01511.x
- Séguy, E. ([1924]) Papillons. Vingt planches en phototypie coloriées au patron donnant 81 papillons et 16 compositions décoratives. Tolmer, Paris, [iv] pp., 20 pls.
- Staudinger, O. (1887) I. Theil. Exotische Tagfalter in systematischer Reihenfolge mit Berücksichtigung neuer Arten. In: Staudinger, O. & Schatz, E. (Eds.), Exotische Schmetterlinge. G. Löwensohn, Fürth, pp. 195–234, pls. 86–95. https://doi.org/10.5962/bhl.title.46867
- Stichel, H. (1901) [A note]. Insekten-Börse, 18 (52), 413.
- Stichel, H. (1902) Aufteilung der Gattung Opsiphanes Westw., Beschreibung neuer Brassoliden und synonymische Notizen. Berliner entomologische Zeitschrift, 46 (4), 487–524, pls. 8–9.
- Stichel, H. (1904) Lepidoptera Rhopalocera, Fam. Nymphalidae, Subfam. Brassolinae. Genera Insectorum, 20, 1–48, 5 pls.
- Stichel, H. (1908) Brassoliden—Studien. Berliner entomologische Zeitschrift, 52 (3), 160–179. https://doi.org/10.1002/mmnd.47919071006
- Stichel, H. (1909) Brassolidae. Das Tierreich, 25, i–xiv + 1–244
- Stichel, H. (1925) Zur Systematik der Brassolidae (Lep. Rhop.). Neue Beiträge zur systematischen Insektenkunde, 3 (8/9), 61–69.
- Stichel, H. (1932) Brassolidae. Lepidopterorum Catalogus, 51, 1–115. https://doi.org/10.5962/bhl.title.124190
- Strand, E. (1918) Nachtrag zum Zweiten Teil meiner “Lepidoptera Niepeltiana”. Societas entomologica, 33 (5), 19–20.
- Uehara-Prado, M., Freitas, A.V.L., Francini, R.B. & Brown, Jr. K.S. (2004) Guia das borboletas frugívoras da Reserva Estadual do Morro Grande e região de Caucaia do Alto, Cotia (São Paulo). Biota neotropica, 4 (1), 1–25, 18 pls. https://doi.org/10.1590/S1676-06032004000100007
- Vane-Wright, R.I. (1972) Scent organs of male butterflies. Report of the British Museum (Natural History), 1969/71, 31–35, 4 pls.
- Vane-Wright, R.I. & Boppré, M. (2005) Adult morphology and the higher classification of Bia Hübner (Lepidoptera: Nymphalidae). Bonner zoologische Beiträge, 53 (1/2), 235-254.
- Warren, A.D., Ogawa, J.R. & Brower, A.V.Z. (2009) Revised classification of the family Hesperiidae (Lepidoptera: Hesperioidea) based on combined molecular and morphological data. Systematic Entomology, 34 (3), 467–523. https://doi.org/10.1111/j.1365-3113.2008.00463.x
- Wolfe, K.V. (2017) Probable last-instar larvae of Opoptera staudingeri mexicana (Nymphalidae, Satyrinae, Brassolini). News of The Lepidopterists’ Society, 59 (1), 28–29.
- Young, A.M. & Muyshondt, A. (1975) Studies on the Natural History of Central American Butterflies in the Family Cluster Satyridae‐Brassolidae‐Morphidae (Lepidoptera: Nymphaloidea). III. Opsiphanes tamarindi and Opsiphanes cassina in Costa Rica and El Salvador. Studies on Neotropical Fauna, 10 (1), 19–55. https://doi.org/10.1080/01650527509360481
- Zhang, D., Gao, F., Jakovlić, I., Zou, H., Zhang, J., Li, W.X. & Wang, G.T. (2020) PhyloSuite: An integrated and scalable desktop platform for streamlined molecular sequence data management and evolutionary phylogenetics studies. Molecular ecology Resources, 20 (1), 348–355. https://doi.org/10.1111/1755–0998.13096
- Zikán, J.F. (1928) Die Macro-Lepidoptera des Itatiaya (Südabhang bei Campo-Bello). Entomologische Rundschau, 45 (3), 10–11.
- Zikán, J.F. & Zikán, W. (1968) Inseto-fauna do Itatiaia e da Mantiqueira, 3: Lepidoptera. Pesquisa agropecuária brasileira, Agronomia, 3, 45–109.