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Type: Monograph
Published: 2022-12-06
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Systematics of Opsiphanes Doubleday, [1849] (Lepidoptera: Nymphalidae, Satyrinae, Brassolini): an integrative approach

Departamento de Zoologia, Universidade Federal do Paraná, CP 19020, 81531-980 Curitiba, Paraná, Brazil
Departamento de Zoologia, Universidade Federal do Paraná, CP 19020, 81531-980 Curitiba, Paraná, Brazil
Departamento de Zoologia, Universidade Federal do Paraná, CP 19020, 81531-980 Curitiba, Paraná, Brazil
Lepidoptera Owl butterflies Taxonomy Synonyms Secondary sexual characters Host plants

Abstract

Opsiphanes Doubleday, [1849] (Lepidoptera: Nymphalidae: Satyrinae: Brassolini) is an exclusively Neotropical genus, occurring from Argentina to Mexico. Until the present study, Opsiphanes was considered to contain 14 species, 60 subspecies, and 38 synonyms. The considerable phenotypic variation of species and subspecies of the genus has affected the taxonomy of the group by causing the proliferation of several names that have been proposed to represent their diversity, taxa that have often not been adequately described and/or delimited. The present study analyzed information on the immature stages and morphology, with molecular data and distribution data, in order to provide revised taxonomic hypotheses for Opsiphanes species and subspecies. These analyses of approximately 5,500 specimens and all species known for the genus made it possible to define two groups: “cassiae” and “quiteria”. The “quiteria” group was subdivided into seven subgroups: “boisduvallii”, “camena”, “zelotes”, “sallei”, “quiteria”, “fabricii”, and “invirae”. The statuses of three species and two subespecies are reinstated: Opsiphanes badius Stichel, 1902 stat. rest., Opsiphanes quirinus Godman & Salvin, 1881 stat. rest., Opsiphanes merianae Stichel, 1902 stat. rest., Opsiphanes bogotanus castaneus Stichel, 1904 stat. rest. and Opsiphanes badius cauca Röber, 1906 stat. rest. Six subspecies are here treated as species: Opsiphanes mexicana Bristow, 1991 stat. nov., Opsiphanes zelus Stichel, 1908 stat. nov., Opsiphanes farrago Stichel, 1904 stat. nov., Opsiphanes barkeri Bristow, 1991 stat. nov., Opsiphanes caliensis Bristow, 1991 stat. nov., and Opsiphanes cuspidatus Stichel, 1904 stat. nov. One subjective synonym is treated as a valid subspecies: Opsiphanes invirae pernambucoensis Bristow, 1991 stat. rev. One species is treated as a subspecies: Opsiphanes cassiae tamarindi C. Felder & R. Felder, 1861 stat. nov. Eight new statuses are proposed: Opsiphanes cassiae incolumis Stichel, 1904 stat. nov., Opsiphanes cassiae tamarindi C. Felder & R. Felder, 1861 stat. nov., Opsiphanes badius angostura Bristow, 1979 stat. nov., Opsiphanes fabricii camposi Bristow, 1991 stat. nov., Opsiphanes fabricii numatius Fruhstorfer, 1912 stat. nov., Opsiphanes merianae notanda Stichel, 1904 stat. nov., Opsiphanes periphetes Fruhstorfer, 1912 stat. nov., and Opsiphanes cuspidatus relucens Fruhstorfer, 1907 stat. nov. Seven subjective synonyms are reinstated: Opsiphanes crameri C. Felder & R. Felder, 1862 syn. rest. of Opsiphanes cassiae cassiae (Linnaeus, 1758); Opsiphanes tamarindi latifascia Rothschild, 1916 syn. rest. of Opsiphanes cassiae incolumis Stichel, 1904 stat. nov.; Opsiphanes erebus Röber, 1927 syn. rest. of Opsiphanes quiteria quirinalis Staudinger, 1887; Opsiphanes cassina aequatorialis Stichel, 1902 syn. rest., Opsiphanes invirae pseudophilon Fruhstorfer, 1907 syn. rest., Opsiphanes invirae remoliatus Fruhstorfer, 1907 syn. rest., and Opsiphanes invirae agasthenes Fruhstorfer, 1907 syn. rest. of Opsiphanes invirae invirae (Hübner, [1808]). Twenty-five new synonyms are proposed: Pavonia Godart [1824] syn. nov. of Bia Hübner, [1819]; Opsiphanes bogotanus phrataphernes Fruhstorfer, 1912 syn. nov., and Opsiphanes bogotanus blandini Bristow, 1991 syn. nov. of Opsiphanes bogotanus bogotanus Distant, 1875; Opsiphanes cassiae alajuela Bristow, 1991 syn. nov. of Opsiphanes bogotanus castaneus Stichel, 1904 stat. rest.; Opsiphanes cassiae rubigatus Stichel, 1904 syn. nov., Opsiphanes cassiae strophios Fruhstorfer, 1907 syn. nov., and Opsiphanes tamarindi xiphos Fruhstorfer, 1907 syn. nov. of Opsiphanes cassiae cassiae (Linnaeus, 1758); Opsiphanes tamarindi corrosus Stichel, 1904 syn. nov., and Opsiphanes tamarindi kleisthenes Fruhstorfer, 1912 syn. nov. of Opsiphanes cassiae tamarindi C. Felder & R. Felder, 1861 stat. nov.; Opsiphanes mutatus parodizi Bristow, 1991 syn. nov. of Opsiphanes farrago Stichel, 1904 stat. nov.; Opsiphanes sallei kennerleyi Bristow, 1991 syn. nov. of Opsiphanes sallei colombiana Bristow, 1991; Opsiphanes quiteria talamancensis Bristow, 1991 syn. nov. of Opsiphanes quirinus Godman & Salvin, 1881 stat. rest.; Opsiphanes quiteria quaestor Stichel, 1902 syn. nov., Opsiphanes quiteria bolivianus Stichel, 1902 syn. nov., and Opsiphanes quiteria cardenasi Bristow, 1991 syn. nov. of Opsiphanes quiteria quiteria (Stoll, 1780); Opsiphanes quiteria phylas Fruhstorfer, 1912 syn. nov. of Opsiphanes quiteria quirinalis Staudinger, 1887; Opsiphanes cassina chiriquensis Stichel, 1902 syn. nov. of Opsiphanes fabricii fabricii (Boisduval, 1870); Opsiphanes cassina milesi Bristow, 1991 syn. nov. of Opsiphanes merianae notanda Stichel, 1904 stat. nov.; Opsiphanes cassina aucotti Bristow, 1991 syn. nov. of Opsiphanes periphetes Fruhstorfer, 1912 stat. nov.; Opsiphanes cassina C. Felder & R. Felder, 1862 syn. nov., Opsiphanes invirae intermedius Stichel, 1902 syn. nov., Opsiphanes invirae amplificatus Stichel, 1904 syn. nov., Opsiphanes sticheli Röber, 1906 syn. nov., Opsiphanes invirae roraimaensis Bristow, 1991 syn. nov., and Opsiphanes invirae sieberti Bristow, 1991 syn. nov. of Opsiphanes invirae invirae (Hübner, [1808]). To ensure unambiguous identification of names, nine neotypes were designated for: Opsiphanes bogotanus Distant, 1875, Opsiphanes aurivillii Röber, 1906, Papilio glycerie Fabricius, 1787, Opsiphanes zelotes zelus Stichel, 1908, Opsiphanes badius var. cauca Röber, 1906, Opsiphanes erebus Röber, 1927, Potamis invirae Hübner, [1808], Opsiphanes sticheli Röber, 1906, and Opsiphanes invirae ledon Fruhstorfer, 1912; and nine lectotypes for: Opsiphanes bogotanus phrataphernes Fruhstorfer, 1912, Opsiphanes tamarindi cherocles Fruhstorfer, 1912, Caligo tamarindi Boisduval, 1870, Opsiphanes sallei nicandrus Fruhstorfer, 1912, Opsiphanes quiteria augeias Fruhstorfer, 1912, Opsiphanes quirinus Godman & Salvin, 1881, Opsiphanes quiteria var. meridionalis Staudinger, 1887, Opsiphanes quiteria oresbios Fruhstorfer, 1912, and Opsiphanes quiteria phylas Fruhstorfer, 1912, . The present taxonomic scheme proposed for Opsiphanes includes 23 species, 23 subspecies, and 69 synonyms.

 

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
    Allio, R., Nabholz, B., Wanke, S., Chomicki, G., Pérez-Escobar, O.A., Cotton, A.M., Clamens, A.-L., Kergoat, G.J., Sperling, F.A.H. & Condamine, F.L. (2021) Genome-wide macroevolutionary signatures of key innovations in butterflies colonizing new host plants. Nature Communications, 12 (354), 1–15. https://doi.org/10.1038/s41467-020-20507-3
    Álvarez-Sierra, J.R. & Álvarez-Corral, J.R. (1984) Mariposas diurnas de Venezuela. Introducción a su conocimiento. Editorial Arte, Caracas, 200 pp.
    Aurivillius, P.O.C. (1882) Recensio critica Lepidopterorum Musei Ludovicae Ulricae quae descripsit Carolus a Linné. Kongliga svenska Vetenskaps-Akademiens Handlingar, 19 (5), 1–188. https://doi.org/10.5962/bhl.title.67730
    Austin, G.T., Warren, A.D., Penz, C.M., Llorente, J.E., Luis-Martínez, A.J. & Vargas-Fernández, I. (2007) A new species of Opsiphanes Doubleday, [1849] from Western Mexico (Nymphalidae: Morphinae: Brassolini). Bulletin of the Allyn Museum, 150, 1–20.
    Bacquet, P.M.B., Brattström, O., Wang, H.-L., Allen, C.E., Löfstedt, C., Brakefield, P.M. & Nieberding, C.M. (2015) Selection on male sex pheromone composition contributes to butterfly reproductive isolation. Proceedings of the Royal Society B: Biological Sciences, 282, 1–9. https://doi.org/10.1098/rspb.2014.2734
    Barth, R. (1952) Die Hautdrüsen des Männchens von Opsiphanes invirae isagoras Fruhst. (Lepidoptera, Brassolidae). Zoologische Jahrbücher, Abteilung für Anatomie und Ontogenie der Tiere, 72 (2), 216–230.
    Bates, D.M. (1932) Notes on the metamorphosis of the Brassolidae (Lepidoptera). Bulletin of the Brooklyn entomological Society, 27 (3), 155–163.
    Beccaloni, G.W., Viloria, A.L., Hall, S.K. & Robinson, G.S. (2008) Catalogue of the host plants of the Neotropical butterflies. Catálogo de las plantas huésped de las mariposas neotropicales. Vol. 8. Sociedad Entomológica Aragonesa [u.a.], Zaragoza, 536 pp.
    Becerra, J.X. (2015) On the factors that promote the diversity of herbivorous insects and plants in tropical forests. Proceedings of the National Academy of Sciences, 112 (19), 6098–6103. https://doi.org/10.1073/pnas.1418643112
    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
    Biezanko, C.M. & Freitas, R.G. de (1938) Catálogo dos insetos encontrados na cidade de Pelotas e seus arredores. Fasciculo 1. Lepidópteros (Contribuição ao conhecimento da fisiografia do Rio Grande do Sul). Boletim. Escola de Agronomia e Veterinária “Eliseu Maciel”, Pelotas, 25, 1–30.
    Biezanko, C.M., Ruffinelli, A. & Carbonell, C.S. (1962) Lepidoptera del Uruguay. Notas complementarias. II. Revista de la Facultad de Agronomía. Universidad de la República, Montevideo, 50, 47–117.

  2. Blandin, P. (1977) La distribution géographique des Brassolinae (Lepidoptera, Satyridae). faits et problèmes. Publications du Laboratoire de Zoologie de l’Ecole normale supérieure, Paris, 9, 161–218.
    Blandin, P., Costa, M. & Attal, S. (2015) Opsiphanes sallei Doubleday, [1849], au Vénézuela: une espèce meconnue (Lepidoptera Nymphalidae Morphinae Brassolini). Antenor, 2 (1), 109–121.
    Blandin, P. & Descimon, H. (1975) Contribution à la connaissance des lépidoptères de l’Equateur. Les Brassolinae (Nymphalidae). Annales de la Société entomologique de France, New Series, 11(1), 3–28.
    Boisduval, J.B.A.D. de (1870) Considérations sur des lépidoptères envoyés du Guatemala à M. de l’Orza. Oberthür et fils., Rennes, Paris, 101 pp.
    Bondar, G.G. (1940) Insetos nocivos e moléstias do coqueiro (Cocos nucifera) no Brasil. Boletim. Instituto central de Fomento econômico da Bahia, Salvador, 8, 118–160.
    Brandão, A.D.S., Farias, P.R.S., Dionisio, L.F.S., Tinôco, R.S., Silva, A.G. & Silva, T.A.F. (2017) Spatial and temporal distribution of Opsiphanes invirae (Lepidoptera: Nymphalidae) in oil palm, Pará State, Brazil. Revista Brasileira de Ciências Agrárias—Brazilian Journal of Agricultural Sciences, 12 (4), 464–469. https://doi.org/10.5039/agraria.v12i4a5479
    Briceño, A.J. (1979) Ciclo de vida de Opsiphanes tamarindi Felder (Lepidoptera: Brassolidae) en Venezuela. Revista de la Facultad de Agronomía, Maracay, Alcance, 26, 107–116.
    Bristow, C.R. (1979) Opsiphanes quiteria angostura a new sub-species of Brassolinae (Lepidoptera; Rhopalocera) from western Ecuador, with notes on collectors and localities. Journal of Natural History, 13 (3), 377–384. https://doi.org/10.1080/00222937900770291
    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
    Butler, A.G. (1870] Catalogue of diurnal Lepidoptera described by Fabricius in the collection of the British Museum. Taylor and Francis, London, 303 pp.
    Campos, F. (1921) Estudios sobre la fauna entomológica del Ecuador. 1° Lepidópteros. Revista del Colegio nacional Vicente Rocafuerte, Guayaquil, 4, 16–62.
    Campos, F. (1927) Catálogo preliminar de los lepidópteros del Ecuador. Primera parte. Ropalóceros. Revista del Colegio nacional Vicente Rocafuerte, Guayaquil, 9 (27/28), 3–106.
    Casagrande, M.M. (1979) Sobre Caligo beltrao (Illiger). III. Morfologia externa do adulto—abdome (Lepidoptera, Satyridae, Brassolinae). Revista Brasileira de Biologia, 39 (3), 711–716.
    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. 201­–204.
    Casagrande, M.M. & Mielke, O.H.H. (2005) Larva de quinto estádio e pupa de Opsiphanes quiteria meridionalis Staudinger (Lepidoptera, Nymphalidae, Brassolinae). Revista Brasileira de Entomologia, 49 (3), 421–424. https://doi.org/10.1590/S0085–56262005000300012
    Chacón, I.A. & Montero, J.J. (2007) Mariposas de Costa Rica. Butterflies and moths of Costa Rica. Instituto Nacional de Biodiversidad, Santo Domingo de Heredia, 366 pp.
    Chacón, I.A., Montero-Ramírez, J., Janzen, D.H., Hallwachs, W., Blandin, P., Bristow, C.R. & Hajibabaei, M. (2012) A new species of Opsiphanes Doubleday,[1849] from Costa Rica (Nymphalidae: Morphinae: Brassolini), as revealed by its DNA barcodes and habitus. Bulletin of the Allyn Museum, 166, 1–15.
    Checa, M.F. & Torres, K. (2019) Notes on the natural history of six nymphalid butterfly species from an Ecuadorian dry forest. Tropical Lepidoptera Research, 29(1), 56–61.
    Checa, M.F., Velasco, N. & Mogollón, H.F. (2010) New distributional records of nymphalid species (Lepidoptera: Nymphalidae) for the Chocó region and Western Ecuador. Tropical Lepidoptera Research, 20 (1), 14–18.
    Clerck, C.A. (1759-[1764]) Icones insectorum rariorum cum nominibus eorum trivialibus, locisque a C. Linnaei Arch: R: et Equ: Aur: Syst: Nat: allegatis. s.n., Holmiae, 55 pp.
    Cock, M.J.W. (2020) Field identification of caterpillars and adults of bark butterflies Opsiphanes spp. (Lepidoptera, Nymphalidae) in Trinidad, W.I. Living World, Journal of the Trinidad and Tobago Field Naturalists’ Club, 2020, 89–91.
    Comstock, J.A. & Vázquez, L. (1961) Estudios de los ciclos biológicos en lepidópteros mexicanos. Anales del Instituto de Biología de México, 31 (1/2), 349–448.
    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, 231–257. https://doi.org/10.1086/276835
    Comstock, J.H. & Needham, J.G. (1899) The wings of insects. The American Naturalist 33, 845–860. https://doi.org/10.1086/277462
    Costa, M., Viloria, A.L., Attal, S., Blandin, P., Neild, A.F.E. & Benmesbah, M. (2020) Lepidoptera del Pantepui. Parte IX. Nuevos Nymphalidae (Satyrinae) y Riodinidae (Riodininae). Antenor, 7 (1), 19–41.
    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, Utrecht and Barthelemy Wild, Amsteldam, 151 pp., pls. 97–192.
    D’Abrera, B.L. (1987) Butterflies of the Neotropical Region. Part III. Brassolidae, Acraeidae & Nymphalidae (partim). Hill House, Victoria, 385 pp.
    Dayrat, B. (2005) Towards integrative taxonomy. Biological Journal of the Linnean Society, 85, 407–415. https://doi.org/10.1111/j.1095-8312.2005.00503.x
    DeVries, P.J. (1987) The butterflies of Costa Rica and their natural history. Papilionidae, Pieridae, Nymphalidae. Princeton University Press, Princeton, 327 pp.
    Dewitz, H. (1878) Entwickelung einiger venezuelanischer Schmetterlinge nach Beobachtungen von Göllmer. Archiv für Naturgeschichte, 44 (1), 1–36.
    Doubleday, E. ([1849]) s.n. In: Westwood, J.O. (Ed.), 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. 45–52 + 56–58 + 60–62 + 64.
    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. https://doi.org/10.1111/j.1365-2311.1929.tb00675.x
    Fabricius, J.C. (1775) Systema entomologiae, sistens insectorvm classes, ordines, genera, species, adiectis synonymis, locis, descriptionibvs, observationibvs. Flensburgi et Lipsiae, Korte, 832 pp. https://doi.org/10.5962/bhl.title.36510
    Fabricius, J.C. (1807) Rechenschaft an das Publicum über seine Classification der Glossaten. Joh. Christ. Fabricii Systema Glossatorum. Zeitung für Literatur und Kunst in den königlichen dänischen Staaten, 21, 81–84.
    Fassl, A.H. (1909) Jugendzustände tropischer Tagfalter. Societas entomologica, 24 (15), 115–117.
    Fassl, A.H. (1910) Tropische Reisen. II. Ueber den Quindiupass. Entomologische Zeitschrift, 24 (21), 113–114.
    Fassl, A.H. (1914a) Tropische Reisen. V. Das obere Caucatal und die Westcordillere. Entomologische Rundschau, 31 (8), 42–46.
    Fassl, A.H. (1914b) Tropische Reisen. VI. Die Hochkordillere von Bogota. Entomologische Rundschau, 31 (17), 97–100.
    Fassl, A.H. (1920) Meine Bolivia-Reise. Entomologische Rundschau, 37 (12), 45–48.
    Felder, C. & Felder, R. (1862) Specimen faunae lepidopterologicae riparum fluminis Negro superioris in Brasilia septentrionali. Wiener entomologische Monatschrift, 6 (3), 65–80.
    Froriep, F. von (1836) Wörterbuch der Naturgeschichte. Vol. 10. Landes-Industrie-Comptoir, Weimar, 582 pp.
    Fruhstorfer, H. (1907a) 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. (1907b) Zwei neue Brassoliden aus Brasilien. Internationale entomologische Zeitschrift, 1 (5), 29–30.
    Fruhstorfer, H. (1912) Brassolidae. In: Seitz, A. (Ed.), The Macrolepidoptera of the World (Gross-Schmett. Erde). Fritz Lehmann Verlag, Stuttgart, pp. 285–332.
    Gernaat, H.B.P.E., Beckles, B.G. & Andel, T.V. (2012) Butterflies of Suriname. A natural history. KIT Publishers, Amsterdam, 680 pp.
    Gistel, J. von N.F.X. (1848) Naturgeschichte des Thierreichs für höhere Schulen. 1st Edition. Hoffmann, Stuttgart, 216 pp.
    Godart, J.B. ([1824]) Papillons. In: Godart, J.B. & Latreille, P. (Eds.), Encyclopédie Méthodique. 9 (2). Veuve Agasse, Paris, pp. 329–828.
    Godman, F.D.C. & Salvin, O. (1881) Biologia Centrali-Americana. Vol. 1. Dulau & Co., Bernard Quaritch, London, pp. 89–168, pls. 9–18.
    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, 307–321. https://doi.org/10.1093/sysbio/syq010
    Hallberg, E. & Poppy, G. (2003) Exocrine Glands: Chemical communication and chemical defense. In: Kristensen, N.P. (Ed.), Lepidoptera, Moths and Butterflies: Evolution, Systematics and Biogeography. Vol. 4. Walter de Gruyter, Berlin, pp. 361–375. https://doi.org/10.1515/9783110893724.361
    Harrison, J.O. (1963) On the Biology of Three Banana Pests in Costa Rica (Lepidoptera: Limacodidae, Nymphalidae). Annals of the Entomological Society of America, 56 (1), 87–94. https://doi.org/10.1093/aesa/56.1.87
    Hayward, K.J. (1951) Catálogo sinonímico de los ropalóceros argentinos excluyendo “Hesperiidae.” Acta zoologica Lilloana, 9, 85–281.
    Hayward, K.J. (1969) Datos para el estudio de la ontogenia de lepidópteros argentinos. Miscelánea. Instituto Miguel Lillo. Universidad nacional de Tucumán, 31, 1–142.
    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: Biological Sciences, 270, 313–321. https://doi.org/10.1098/rspb.2002.2218
    Hemming, F. (1943) Notes on the generic nomenclature of the Lepidoptera Rhopalocera, II. Proceedings of the Royal Entomological Society of London, Series B, Taxonomy, 12 (2), 23–30. https://doi.org/10.1111/j.1365-3113.1943.tb00739.x
    Herbst, J.F.W. (1793) Der Schmetterlinge. In: Jablonsky, C.G. (Ed.), Natursystem aller bekannten in- und ausländischen Insekten als eine Fortsetzung der von Buffonschen Naturgeschichte. Nach dem System des Ritters Carl von Linné. Joachim Pauli, Berlin, pp. 6 + 1–162, pls. 118–153. https://doi.org/10.1515/9783112444245-002
    Hewitson, W.C. (1873) s.n. In: Illustrations of new species of exotic butterflies. 5 (86). John Van Voorst, London, pp. [11–12] + [45–46] + [61–62], pls. [6] + [24] + [33].
    Hoffmann, F. (1930) Beiträge zur Naturgeschichte brasilianischer Schmetterlinge. I. Zeitschrift für wissenschaftliche Insektenbiologie, 25 (6/7), 92–112.
    Hoffmann, F. (1932) Beiträge zur Naturgeschichte brasilianischer Schmetterlinge. III. Zeitschrift für wissenschaftliche Insektenbiologie, 26 (4/6), 109–124.
    Honey, M.R. & Scoble, M.J. (2001) Linnaeus’s butterflies (Lepidoptera: Papilionoidea and Hesperioidea). Zoological Journal of the Linnean Society, 132 (3), 277–399. https://doi.org/10.1111/j.1096-3642.2001.tb01326.x
    Horn, W.H.R. & Kahle, I. (1937) Nachtrag (Supplemente: Addenda et Corrigenda). In: Über entomologische Sammlungen. (Ein Beitrag zur Geschichte der Entomo-Museologie). Entomologische Beihefte aus Berlin-Dahlem. Gobler, Berlin-Dahlem, pp. 297–536 + i–vi, pls. 27–38.
    Hübner, J. ([1808]) s.n. In: Sammlung exotischer Schmetterlinge. Vol. 1. Jacob Hübner, Augsburg, pl. [76].
    Hübner, J. ([1819]) s.n. In: Verzeichniss bekannter Schmettlinge. Vol. 2–8. Jacob Hübner, Augsburg, pp. 17–128.
    Hughes, R.A. (1959) Notes on the butterfly fauna of Tingo Maria, East-central Peru. Entomologist, 92 (1151), 80–87.
    Janzen, D.H. & Hallwachs, W. (2009) Dynamic database for an inventory of the Macrocaterpillar fauna, and its food plants and parasitoids, of Área de Conservacion Guanacaste (ACG), Northwestern Costa Rica. Available from: http://janzen.sas.upenn.edu (accessed 29 May 2021)
    Katoh, K. & Standley, D.M. (2013) MAFFT Multiple Sequence Alignment Software Version 7: Improvements in Performance and Usability. Molecular Biology and Evolution, 30, 772–780. https://doi.org/10.1093/molbev/mst010
    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
    Kirby, W.F. (1871) A Synonymic Catalogue of Diurnal Lepidoptera. John Van Voorst, London, 690 pp. https://doi.org/10.5962/bhl.title.23779
    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, 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.
    Krüger, E. (1920) Ueber sekundäre sexuelle Organe der Morphiden und anderer Tagfalterfamilien nebst Angaben über den Geruch einiger kolumbianischer Tagfalter. Entomologische Rundschau, 37 (10), 37–39.
    Kumar, S., Stecher, G., Li, M., Knyaz, C., Tamura, K. & Battistuzzi, F.U. (2018) MEGA X: Molecular Evolutionary Genetics Analysis across Computing Platforms. Molecular Biology and Evolution, 35, 1547–1549. https://doi.org/10.1093/molbev/msy096
    Lamas, G. (1976) A gazetteer of Peruvian entomological stations (based on Lepidoptera). Revista peruana de Entomología, 19 (1), 17–25.
    Lamas, G. (2021) An Annotated Bibliography of the Neotropical Butterflies and Skippers (Lepidoptera: Papilionoidea and Hesperioidea). Available from: http://www.butterfliesofamerica.com/L/Biblio.htm (accessed 29 April 2021)
    Link, D. & Alvarez-Filho, A. (1979) Palmeiras atacadas por lagartas de Brassolidae (Lepidoptera) em Santa Maria, RS. Revista do Centro de Ciências rurais, Santa Maria, 9 (2), 221–225.
    Linnaeus, C. (1758) Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synomymis, locis. 10th Edition. Laurentius Salvius, Holmiae, 823 pp. https://doi.org/10.5962/bhl.title.542
    Linnaeus, C. (1764) Museum S:ae R:ae M:tis Ludovicae Ulricae Reginae Svecorum, Gothorum, Vandalorumque &c. &c. &c. In quo animalia rariora, exotica, imprimis Insecta & Conchilia describuntur & determinantur. Prodromi instar editum. Laurentius Salvius, Holmiae, 720 pp. https://doi.org/10.5962/bhl.title.51532
    Llorente-Bousquets, J.E., Luis-Martínez, A., Vargas-Fernández, I. & Warren, A.D. (1996) Lista de las mariposas del Estado de Jalisco, México. Revista de la Sociedad mexicana de Historia natural, 46, 35–48.
    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 E., R.G. de la & Maza E., J. de la (1993) Mariposas de Chiapas. Gobierno del Estado de Chiapas, México, 224 pp.
    Maza E., R.G. de la & Maza E., J. de la (2017) Las Mariposas Diurnas (Papilionoidea y Hesperioidea) de los Altos de Chiapas, Mexico, Distribución, Composición, Origen y Evolución. Revista de la Sociedad mexicana de Lepidopterología, Nueva Serie, 5(1), 9–40.
    Maza R., R.F. de la (1987) Mariposas mexicanas. Guía para su colecta y determinación. Fondo de Cultura Económica, México City, 304 pp.
    Merian, M.S. (1705) Metamorphosis insectorum surinamensium. Gerard Valk., Amstelodami, 60 pp., 60 pls.
    Miller, J.C., Janzen, D.H. & Hallwachs, W. (2006) 100 caterpillars: portraits from the tropical forests of Costa Rica. Belknap Press of Harvard University Press, Cambridge, 264 pp.
    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 20 May 2021)
    Minh, B.Q., Nguyen, M.A.T. & von Haeseler, A. (2013) Ultrafast Approximation for Phylogenetic Bootstrap. Molecular Biology and Evolution, 30, 1188–1195. https://doi.org/10.1093/molbev/mst024
    Minh, B.Q., Schmidt, H.A., Chernomor, O., Schrempf, D., Woodhams, M.D., von Haeseler, A. & Lanfear, R. (2020) IQ-TREE 2: New Models and Efficient Methods for Phylogenetic Inference in the Genomic Era. Molecular Biology and Evolution, 37, 1530–1534. https://doi.org/10.1093/molbev/msaa015
    Monte, O. (1934) Borboletas que vivem em plantas cultivadas. Boletim de Agricultura, Zootechnia e Veterinaria, Belo Horizonte, 7 (3), 145–165.
    Müller, W. (1886) Südamerikanische Nymphalidenraupen Versuch eines natürlichen Systems der Nymphaliden. G. Fischer, Jena, 255 pp. https://doi.org/10.5962/bhl.title.109410
    Neves, D.A. & Paluch, M. (2016) Biology and External Morphology of Immatures of Opsiphanes quiteria meridionalis Staudinger (Lepidoptera: Nymphalidae). Neotropical Entomology, 45 (1), 13–21. https://doi.org/10.1007/s13744-015-0323-4
    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
    Nieberding, C.M., de Vos, H., Schneider, M.V., Lassance, J.-M., Estramil, N., Andersson, J., Bång, J., Hedenström, E., Löfstedt, C. & Brakefield, P.M. (2008) The male sex pheromone of the butterfly Bicyclus anynana: Towards an evolutionary analysis. PLoS ONE, 3, 1–13. https://doi.org/10.1371/journal.pone.0002751
    Nijhout, H.F. (1991) The development and evolution of butterfly wing patterns. Smithsonian Institution Press, Washington, 297 pp.
    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. Springer Singapore, Singapore, pp. 3–19. https://doi.org/10.1007/978-981-10-4956-9_1
    Núñez-Bustos, E.O. (2010) Mariposas de la ciudad de Buenos Aires y alrededores. Vázquez Mazzini Editores, Buenos Aires, 264 pp.
    Orlandin, E., Piovesan, M. & Carneiro, E. (2020) Borboletas do Meio—Oeste de Santa Catarina: História Natural e Guia de Identificação. 1st Edition. Authors, Joaçaba, 399 pp.
    Otero, L.S. & Marigo, L.C. (1990) Butterflies. Beauty and behavior of Brazilian species. Marigo Comunicação Visual, Rio de Janeiro, 128 pp.
    Padial, J.M., Miralles, A., De la Riva, I. & Vences, M. (2010) The integrative future of taxonomy. Frontiers in Zoology, 7, 16. https://doi.org/10.1186/1742-9994-7-16
    Pastrana, J.A. (2004) Los lepidópteros argentinos. Sus plantas hospedadoras y otros sustratos alimenticios. Sociedad Entomológica Argentina, Buenos Aires, 334 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., Aiello, A. & Srygley, R. (1999) Early stages of Caligo illioneus and C. idomeneus (Nymphalidae, Brassolinae) from Panama, with remarks on larval food plants for the subfamily. Journal of the Lepidopterists’ Society, 53 (4), 142–152.
    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., Mohammadi, N. & Wahlberg, N. (2011) Neotropical Blepolenis butterflies: wing pattern elements, phylogeny, and Pleistocene diversification (Lepidoptera, Nymphalidae). Zootaxa, 289 (1), 1–17. https://doi.org/10.11646/zootaxa.2897.1.1
    Pulido, J.I. & Cárdenas, R. (1979) Biología del gusano cabra de las hojas del plátano Opsiphanes envirae Hübner (Lepidoptera: Brassolidae). Revista colombiana de Entomología, 5 (3/4), 45–51. https://doi.org/10.25100/socolen.v5i3-4.10363
    Raguso, R.A. & Gloster, O. (1996) Preliminary checklist and field observations of the butterflies of the Maquipucuna Field Station (Pichincha Province, Ecuador). Journal of Research on the Lepidoptera, 32, 135–161. https://doi.org/10.5962/p.266610
    Ratnasingham, S. & Hebert, P.D.N. (2007) BARCODING: bold: The Barcode of Life Data System (http://www.barcodinglife.org). Molecular Ecology Notes, 7, 355–364. https://doi.org/10.1111/j.1471-8286.2007.01678.x
    Ribeiro, R.C., Lemos, W.P., Bernardino, A.S., Buecke, J. & Müller, A.A. (2010) Primeira ocorrência de Alcaeorrhynchus grandis (Dallas) (Hemiptera: Pentatomidae) predando lagartas desfolhadoras do dendezeiro no Estado do Pará. Neotropical Entomology, 39 (1), 131–132. https://doi.org/10.1590/S1519-566X2010000100018
    Röber, J.K.M. (1906) Neue Brassoliden. Societas entomologica, 21 (3), 18–21.
    Röber, J.K.M. (1927) Neue exotische Falter. Internationale entomologische Zeitschrif, 21 (13), 97–100.
    Rodríguez-González, G., Silva, R., Cásares, R., Barrios, R., Díaz, A. & Fariñas, J. (2012) Tecnología agronómica de la palma aceitera (Elaeis guineensis Jacq.) y manejo integrado de su defoliador Opsiphanes cassina Felder (Lepidoptera: Brassolidae) en plantaciones comerciales del estado Monagas, Venezuela. UDO agrícola, 12 (3), 584–598.
    Ross, G.N. (1976) An ecological study of the butterflies of the Sierra de Tuxtla in Veracruz, Mexico. Journal of Research on the Lepidoptera, 15 (1), 41–60. https://doi.org/10.5962/p.333707
    Rothschild, L.W. (1916) Notes on Amathusiidae, Brassolidae, Morphidae, etc., with descriptions of new forms. Novitates zoologicae, 23 (3), 299–318.
    Schneider, C.A., Rasband, W.S. & Eliceiri, K.W. (2012) NIH Image to ImageJ: 25 years of image analysis. Nature Methods, 9, 671–675. https://doi.org/10.1038/nmeth.2089
    Scoble, M.J. (1995) Hearing, sound, and scent. In: Scoble, M.J. (Ed.), The lepidoptera: form, function, and diversity. Oxford University Press, Oxford, pp. 160–169.
    Scudder, S.H. (1875) Historical Sketch of the generic names proposed for butterflies, A contribution to systematic nomenclature. Proceedings of the American Academy of Arts and Sciences, 10 (2), 91–293. https://doi.org/10.5962/bhl.title.9688
    Shorthouse, D.P. (2010) SimpleMappr, an online tool to produce publication—quality point maps. Available from: http://www.simplemappr.net (accessed 27 November 2020)
    Silva, A.G. d’Araújo, Gonçalves, C.R., Galvão, D.M., Gonçalves, A.J.L., Gomes, J., Silva, M.N. & Simoni, L. (1968) Quarto catálogo dos insetos que vivem nas plantas do Brasil seus parasitos e predadores. Edição ampliada do “3° catálogo dos insetos que vivem nas plantas do Brasil” de autoria do Prof. A. M. da Costa Lima. Parte II. Insetos, hospedeiros e inimigos naturais. Indice de insetos e índice de plantas. Ministério da Agricultura, Rio de Janeiro, 622 + 265 pp.
    Smart, P.E. (1975) The illustrated encyclopedia of the butterfly world in colour. Salamander Books, London, 275 pp.
    Specht, M.J. da S., Borges, L.R. & Paluch, M. (2011) Borboletas associadas à Heliconia velloziana L. Emygd. (Zingiberales: Heliconiaceae) no Estado de Pernambuco. Revista Biociências, Taubaté, 17 (1), 27–32.
    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. 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, 487–524.
    Stichel, H. (1904) Lepidoptera Rhopalocera, Fam. Nymphalidae, Subfam. Brassolinae. Genera Insectorum, 20, 1–48.
    Stichel, H. (1907) Kommentar zum Artikel: Röber, Neue Brassoliden im Jahrgang XXI, pp. 18–21 und 27–28 dieser Zeitschrift. Societas entomologica, 22 (13), 98–101.
    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. (1916) Lepidoptera Niepeltiana. 2 Teil. Wilhelm Niepelt, Zirlau, 26 pp., pls. 13–17.
    Vane-Wright, R.I. (1972) Scent organs of male bufferflies. Report of the British Museum (Natural History), 1969/71, 31–35, 4 pls.
    Vargas-Fernández, I., Llorente-Bousquets, J. & Luis-Martínez, A. (2016) Adiciones a la serie Papilionoidea de México. Distribución geográfica e ilustración. Universidad Nacional Autónoma de México, México City, 120 pp.
    Vargas-Fernández, I., Llorente-Bousquets, J.E., Luis-Martínez, A. & Pozo, M. del C. (2008) Nymphalidae de México II (Libytheinae, Morphinae y Charaxinae): Distribución geográfica e ilustración. Universidad Nacional autónoma de México, México City, 226 pp.
    Waterhouse, C.O. (1881) Aid to the identification of insects. Vol. 1. E. W. Janson, London, pls. 9–83­­.
    Winter, S., Friedman, A.L.L., Astrin, J.J., Gottsberger, B. & Letsch, H. (2017) Timing and host plant associations in the evolution of the weevil tribe Apionini (Apioninae, Brentidae, Curculionoidea, Coleoptera) indicate an ancient co-diversification pattern of beetles and flowering plants. Molecular Phylogenetics and Evolution, 107, 179–190. https://doi.org/10.1016/j.ympev.2016.10.015
    Young, A.M. (1977) Notes on the defoliation of Coconut Palm (Cocos nucifera) by the butterfly Opsiphanes quiteria quirinus in northeastern Costa Rica (Lepidoptera: Brassolidae). Deutsche Entomologische Zeitschrift, 24 (4/5), 353–365. https://doi.org/10.1002/mmnd.19770240408
    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, 348–355. https://doi.org/10.1111/1755–0998.13096

  3.