The butterfly subtribe Euptychiina (Lepidoptera: Nymphalidae) is a diverse group within the tribe Satyrini, occurring throughout Central and South America, in addition to a few species known from North America. Euptychiina is a predominantly lowland group, with the exception of one Asian taxon Palaeonympha opalina Butler, 1871 and the Andean genus Forsterinaria Gray, 1973. The taxon was erected by Lee Denmar Miller.
Euptychiina Despite its members being common, this subtribe has been a challenging subject for taxonomic and phylogenetic studies for many years because of their dull coloration, intraspecific variation, lack of clear morphological characters, and morphological homogeneity. However, with the exception of pioneering work by W. Forster and L. D. Miller, the group received little attention from butterfly researchers until recently due to their typically dull brownish coloration. Currently, 50 genera and over 400 described species are recognized within this subtribe, but the group is estimated to contain over 500 species in 70 genera. The current classification of Euptychiina is based on the Lamas checklist, who retained and reorganized many of the genera erected by Forster. Forster described 33 euptychiine genera that are now widely accepted, but since he erected these genera without testing monophyly and synapomorphies, many of his genera have been recovered as polyphyletic or paraphyletic in recent molecular phylogenetic studies.
History of classification A. Butler was probably the first to propose a systematic classification for many euptychiine taxa, excluding species now in the "Taygetis clade". In his monograph of Euptychia (then used as a catch-all genus to include most euptychiine species), Butler divided the genus into seven groups (Division I to VII). Subsequently, Butler proposed an updated classification for the group and recognized 27 "species groups" within Euptychia sensu Butler (1867). G. Weymer recognized 29 "species group" within Euptychia sensu Butler, in addition to treating Taygetis and Amphidecta in his monograph of Satyridae in the "Macrolepidoptera of the American Faunistic region" by A. Seitz.Weymer, G. 1910-1911. 4. Familie: Satyridae. In: Seitz, A. (Ed.), Die Gross-Schmetterlinge der Erde. Stuttgart, A. Kernen. 5: 185-225. Based on Weymer (1911)'s classification, Forster (1964) introduced 33 euptychiine genera and his classification is largely retained in Lamas (2004), a work considered as a vital foundation regarding Neotropical butterfly classification. The recent years have seen an explosion of interest in euptychiine systematics, resulting in many changes in generic classification of the group as well as improvement in our understanding of its species diversity. Although the subtribal name was first introduced by L. D. Miller when he treated Euptychiina as a tribal level taxon "Euptychiini", the genus Euptychia Hübner, 1818 was historically used to place many euptychiine species now no longer classified in that genus, perhaps explaining why the generic name Euptychia was used in a much broader sense to include many other euptychiine species. Consequently, Forster included this name "Euptychia" as part of new generic names he described, a trend also followed when the new generic name Atlanteuptychia was introduced for Euptychia ernestina Weymer, 1911. However, other recently described euptychiine genera do not follow this trend (references above).
Biology There exist detailed early stage biology (i.e. complete life cycle documented) information for 30 euptychiine species.Freitas, A. V. L., Carreira, J. Y. O., Santos, J. P., Barbosa, E. P. 2016. Immature stages and natural history of two species of Forsterinaria from southeastern Brazil (Lepidoptera: Nymphalidae). Tropical Lepidoptera Research, 26(1): 13-18.Freitas, A. V. L., Barbosa, E. P., Marin, M. A. 2016. Immature stages and natural history of the neotropical satyrine Pareuptychia ocirrhoe interjecta (Nymphalidae: Satyrinae). Journal of the Lepidopterists' Society, 70(4): 271-276.Freitas, A. V. L. 2017. Immature stages of the Neotropical satyrine Taygetis acuta (Nymphalidae: Euptychiina). Tropical Lepidoptera Research, 27: 1-5.Freitas, A. V. L., Rosa, A. H. B., Carreira, J. Y. O., Gueratto, P. Y., Santos J. P., Tacioli A. 2018. Immature stages of two Moneuptychia from southeastern Brazil (Nymphalidae: Euptychiina). Tropical Lepidoptera Research 28: 100-105.See, J., Nakahara, S., Gallice, G. 2018 Immature stages of Splendeuptychia quadrina (Butler, 1869) (Lepidoptera: Nymphalidae: Satyrinae). Tropical Lepidoptera Research, 28 (2): 49–53. Larva of euptychiine species often lack body scoli and possessing short head scoli and caudal filaments, but there exist some variation. Hostplant records are known for approximately 100 species, those records are mainly grass and bamboo species,Beccaloni, G. W., Viloria, A. L., Hall, S. K., Robinson, G. S. 2008. Catalogue of the Hostplants of the Neotropical Butterflies / Catálogo de las Plantas Huésped de las Mariposas Neotropicales. m3m-Monografías Tercer Milenio, Volume 8. Zaragoza, Spain: Sociedad Entomológica Aragonesa (SEA)/Red Iberoamericana de Biogeografía y Entomología Sistemática (RIBES)/Ciencia y Tecnología para el Desarrollo (CYTED)/Natural History Museum, London, U. K. (NHM)/Instituto Venezolano de Investigaciones Científicas, Venezuela (IVIC). 1-536 pp. although the genus Euptychia is known to feed on mosses and lycopsids.Singer, M. C., Ehrlich, P. R., Gilbert, L. E. 1971. Butterfly feeding on lycopsid. Science, 172: 1341-1342.DeVries, P. J. 1986. Hostplant records and natural history notes on Costa Rican butterflies (Papilionidae, Pieridae & Nymphalidae). Journal of Research on the Lepidoptera, 24: 290‑333.Brévignon, C. 2008. Inventaire des Satyrinae de Guyane Française (Lepidoptera: Nymphalidae), pp. 62-94. In: Lacomme, D., Manil, L. (Eds.),Lépidoptères de Guyane, Tome 3, Rhopalocères 2. Paris, Lepidopteristes de France.
Generic classification and species
The following genera and species belong to subtribe Euptychina. +generaspecies and/or subspeciesimages of example speciesAmiga Nakahara, Willmott & Espeland, 2019
Type species — Papilio arnaca Fabricius, 1776 A. arnaca (Fabricius, 1776) A. arnaca arnaca (Fabricius, 1776) A. arnaca adela Nakahara & Espeland, 2019 A. arnaca indianacristoi Nakahara & Marín, 2019 A. sericeella (Bates, 1865)Amphidecta Butler, 1867
Type species — Euptychia pignerator Butler, 1867A. calliomma (C. Felder & R. Felder, 1862) A. clio (Weymer, 1911) A. pignerator (Butler, 1867) A. pignerator pignerator (Butler, 1867) A. pignerator simplicia Weymer, 1910 A. reynoldsi Sharpe, 1890Archeuptychia Forster, 1964
Type species — Papilio cluena Drury, 1782 A. cluena (Drury, 1782)Argentaria Huertas & Willmott, 2023
Type species — Euptychia itonis Hewitson, 1862 A. argyropsacas (Bryk, 1953) A. clementia (Butler, 1877) A. clorimena (Stoll, 1790) A. cosmophila (Hübner, 1823) A. hygina (Butler, 1877) A. itonis (Hewitson, 1862) A. jadea (Brévignon & Benmesbah, 2011) A. kendalli (L.D. Miller, 1976) A. libitina (Butler, 1870) A. pagyris (Godart, ) A. quadrina (Butler, 1869) A. salvini (Butler, 1867) A. telesphora (Butler, 1877) A. zischkai (Forster, 1964)Atlanteuptychia Freitas, Barbosa & Mielke, 2013
Type species — Euptychia ernestina Weymer, 1911 A. ernestina (Weymer, 1911)Caeruleuptychia Forster, 1964
Type species — Euptychia caerulea Butler, 1867C. aegrota (Butler, 1867) C. aetherialis (Butler, 1867) C. caerulea (Butler, 1869) C. coelestis (Butler, 1867) C. coelica (Hewitson, 1869) C. cyanites (Butler, 1871) C. divina (Weymer, 1911) C. francisca (Butler, 1870) C. glauca (Weymer, 1911) C. helena (Anken, 1994) C. helios (Weymer, 1911) C. lobelia (Butler, 1870) C. mare (Butler, 1869) C. maryzenderae Lamas & Nakahara, 2017 C. penicillata (Godman, 1905) C. pilata (Butler, 1867) C. romani (Aurivillius, 1929) C. saul Brévignon & Benmesbah, 2011 C. scopulata (Godman, 1905) C. scripta Nakahara, Zacca & Huertas, 2017 C. tenera (Weymer, 1911) C. thaliana Nakahara & Piovesan, 2022 C. trembathi Willmott, Nakahara, Hall & Neild, 2017 C. twalela (Brévignon, 2005) C. umbrosa (Butler, 1870) C. urania (Butler, 1867) C. ziza (Butler, 1869)Capronnieria Forster, 1964
Type species — Satyrus galesus Godart, C. galesus (Godart, ) C. narapa (Schaus, 1902)Carminda Ebert & Dias, 1998
Type species — Satyrus paeon Godart C. griseldis (Weymer, 1911) C. paeon (Godart, ) C. surpresa Barbosa, Aguiar, Rosa, Zacca & Freitas, 2020 C. umuarama (Ebert & Dias, 1997)Cepheuptychia Forster, 1964
Type species — Papilio cephus Fabricius, 1775C. angelica (Butler, 1874) C. cephus (Fabricius, 1775) C. glaucina (Bates, 1864)Chloreuptychia Forster, 1964
Type species — Papilio chloris Cramer, 1778C. agatha (Butler, 1867) C. amenthesysta Brévignon & Benmesbah, 2011 C. chlorimene (Hübner, ) C. gordonsmalli Thurman, MacDonald & Nakahara, 2021Cisandina Nakahara & Espeland, 2022
Type species — C. castanya Lamas & Nakahara, 2022 C. esmeralda Nakahara & Barbosa, 2022 C. fida (Weymer, 1911) C. fida fida (Weymer, 1911) C. fida directa Nakahara & Willmott, 2022 C. lea (Cramer, 1777) C. philippa (Butler, 1867) C. sanmarcos (Nakahara & Lamas, 2018) C. trinitensis (Brévignon & Benmesbah, 2012)Cissia Doubleday, 1848
Type species — Papilio penelope Fabricius, 1775C. anabelae (L.D. Miller, 1976) C. cheneyorum (Chermock, 1949) C. cleophes (Godman & Salvin, 1889) C. eous (Butler, 1867) C. penelope (Fabricius, 1775) C. phronius (Godart, ) C. pompilia (C. Felder & R. Felder, 1867) C. proba (Butler, 1867) C. pseudocleophes (Miller, 1976) C. rubricata (Edwards, 1871) C. rubricata rubricata (Edwards, 1871) C. rubricata smithorum (Wind, 1946) C. wahala Grishin, 2022Colombeia Viloria, Andrade & Le Crom, 2019
Type species —C. hotchkissi (Dyar, 1913) C. mycalesis (Röber, 1927) C. nossis (Hewitson, 1862)Cristalinaia Freitas, Barbosa & Zacca, 2019
Type species — Cristalinaia vitoria Mota, Zacca & Freitas, 2019C. vitoria Mota, Zacca & Freitas, 2019Cyllopsis R. Felder, 1869 Type species — Neonympha hedemanni R. Felder, 1869C. argentella (Butler & H. Druce, 1872) C. caballeroi (Beutelspacher, 1982) C. clinas (Godman & Salvin, 1889) C. diazi Miller, 1974 C. dospassosi Miller, 1974 C. emilia Chacón & Nishida, 2002 C. gemma (Hübner, 1808) C. gemma gemma (Hübner, 1808) C. gemma freemani (Stallings & Turner, 1947) C. guatemalena Miller, 1974 C. hedemanni R. Felder, 1869 C. hedemanni hedemanni R. Felder, 1869 C. hedemanni tamaulipensis Miller, 1974 C. hedemanni vetones (Godman & Salvin, 1878) C. hilaria (Godman, 1901) C. jacquelineae Miller, 1974 C. nayarit (Chermock, 1947) C. nelsoni (Godman & Salvin, 1881) C. pallens Miller, 1974 C. parvimaculata Miller, 1974 C. pephredo (Godman, 1901) C. perplexa Miller, 1974 C. perpepida (Dyar, 1912) C. pertepida perpepida (Dyar, 1912) C. pertepida avicula (Nabokov, 1942) C. pertepida intermedia Miller, 1974 C. pertepida maniola (Nabokov, 1942) C. philodice (Godman & Salvin, 1878) C. pseudopephredo (Chermock, 1947) C. pyracmon (Butler, 1867) C. pyracmon pyracmon (Butler, 1867) C. pyracmon henshawi (Edwards, 1876) C. rogersi (Godman & Salvin, 1878) C. schausi (Miller, 1974) C. steinhauserorum Miller, 1974 C. suivalenoides Miller, 1974 C. suivalens (Dyar, 1914) C. suivalens suivalens (Dyar, 1914) C. suivalens escalantei Miller, 1974 C. tomemmeli Warren & Nakahara, 2018 C. wellingi Miller, 1978 C. whiteorum Miller & de la Maza, 1984 C. windi Miller, 1974Deltaya Willmott, Nakahara & Espeland, 2023
Type species — Papilio ocypete Fabricius, D. andrei (Zacca, Casagrande & Mielke, 2017) D. louisammour (Benmesbah & Zacca, 2018) D. ocypete (Fabricius, 1776) D. opima (Weymer, 1911) D. pallema (Schaus, 1902) D. probata (Weymer, 1911)Emeryus Zacca, Casagrande & Mielke, 2020
Type species — Papilio ocypete Fabricius, E. argulus (Godart, ) E. argulus argulus (Godart, ) E. argulus magnum Zacca, Casagrande & Mielke, 2020