Euptychiina

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.[1][2]

Euptychiina
Arizona pine satyr (Paramacera allyni)
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Lepidoptera
Family:	Nymphalidae
Tribe:	Satyrini
Subtribe:	Euptychiina L. D. Miller
Genera
About 50

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.[2] However, with the exception of pioneering work by W. Forster[3] and L. D. Miller,[4] 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,[5][6] but the group is estimated to contain over 500 species in 70 genera.[6] The current classification of Euptychiina is based on the Lamas checklist,[7] 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.[1][8]

Phylogeny showing relationships of major euptychiine genera[8]

History of classification

A. Butler[9] 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).[10] 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.[11] 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.[3][7] 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.[6][12] Although the subtribal name was first introduced by L. D. Miller when he treated Euptychiina as a tribal level taxon "Euptychiini",[4] 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.[13] 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.[14][15][16][17][18][19] Larva of euptychiine species often lack body scoli and possessing short head scoli and caudal filaments, but there exist some variation.[19] Hostplant records are known for approximately 100 species, those records are mainly grass and bamboo species,[20] although the genus Euptychia is known to feed on mosses and lycopsids.[21][22][23]

Euptychia westwoodi mating on its host plant (Selaginella species) in western Ecuador.

Early stage biology (egg, larva, pupa and adult) of Splendeuptychia quadrina

Generic classification and species accounts (as of February 2019)

genera	species and/or subspecies		genera	species and/or subspecies
Amiga Nakahara, Willmott & Espeland, 2019 Type species - Papilio arnaca Fabricius, 1776	A. arnaca arnaca (Fabricius, 1776) A. arnaca adela Nakahara & Espeland, 2019 A. arnaca sericeella (Bates, 1865) A. arnaca indianacristoi Nakahara & Marín, 2019		Orotaygetis Nakahara & Zacca, 2018 Type species - Orotaygetis surui Nakahara, Zacca & Lamas, 2018	O. surui Nakahara, Zacca & Lamas, 2018
Amphidecta Butler, 1867 Type species - Euptychia pignerator Butler, 1867	A. calliomma (C. Felder & R. Felder, 1862) A. pignerator pignerator (Butler, 1867) A. pignerator simplicia Weymer, 1910 A. reynoldsi Sharpe, 1890		Paleonympha Butler, 1871 Type species - Paleonympha opalina Butler, 1871	P. opalina Butler, 1871
Archeuptychia Forster, 1964 Type species - Papilio cluena Drury, 1782	A. cluena (Drury, 1782)		Paramacera Butler, 1868 Type species - Neonympha xicaque Reakirt, [1867]	P. allyni Miller, 1972 P. chinanteca Miller, 1972 P. copiosa Miller, 1972 P. xicaque xicaque (Reakirt, [1867]) P. xicaque rubrosuffusa Miller, 1972
Atlanteuptychia Freitas, Barbosa & Mielke, 2013 Type species - Euptychia ernestina Weymer, 1911	A. ernestina (Weymer, 1911)		Parataygetis Forster, 1964 Type species - Euptychia albinota Butler, 1867	P. albinotata (Butler, 1867) P. lineata (Godman & Salvin, 1880)
Caeruleuptychia Forster, 1964 Type species - Euptychia caerulea Butler, 1867	C. aegrota (Butler, 1867) C. aetherialis (Butler, 1867) C. brixius (Godart, [1824]) C. caerulea (Butler, 1869) C. coelestis (Butler, 1867) C. coelica (Hewitson, 1869) C. cyanites (Butler, 1871) C. divina (Weymer, 1911) 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. scopulata (Godman, 1905) C. scripta Nakahara, Zacca & Huertas, 2017 C. tenera (Weymer, 1911) C. trembathi Willmott, Nakahara, Hall & Neild, 2017 C. twalela (Brévignon, 2005) C. umbrosa (Butler, 1870) C. urania (Butler, 1867) C. ziza (Butler, 1869)		Pareuptychia Forster, 1964 Type species - Pareuptychia hesionides Forster, 1964	P. binocula (Butler, 1869) P. difficilis (Forster, 1964) P. hervei Brévignon, 2005 P. hesioniades hesioniades Forster, 1964 P. hesioniades deviae Brévignon, 2005 P. lydia (Cramer, 1777) P. metaleuca (Boisduval, 1870) P. milleri Nakahara, Marín & Neild, 2016 P. ocirrhoe ocirrhoe (Fabricius, 1776) P. ocirrhoe interjecta (d'Almeida, 1952) P. summandosa (Gosse, 1880)
Capronnieria Forster, 1964\ Type species - Satyrus galesus Godart, [1824]	C. galesus (Godart, [1824])		Paryphthimoides Forster, 1964 Type species - Neonympha polytis Prittwitz, 1865	P. poltys poltys (Prittwitz, 1865) P. poltys numilia (C. Felder & R. Felder, 1867) P. vestigiata (Butler, 1867) P. undulata (Butler, 1867) P. grimon (Godart, [1824]) P. argulus (Godart, [1824]) P. difficilis (Forster, 1964) P. numeria (C. Felder & R. Felder, 1867) P. melobosis (Capronnier, 1874) P. sylvina (C. Felder & R. Felder, 1867)
Cepheuptychia Forster, 1964 Type species - Papilio cephus Fabricius, 1775	C. angelica (Butler, 1874) C. cephus (Fabricius, 1775) C. glaucina (Bates, 1864) C. romani (Aurivillius, 1929)		Pharneuptychia Forster, 1964 Type species - Satyrus phares Godart, [1824]	P. haywardiana Lamas, 2010 P. phares (Godart, [1824])* P. pharnabazos (Bryk, 1953) P. pharnaces (Weymer, 1911) P. romanina (Bryk, 1953) P. innocentia (C. Felder & R. Felder, 1867) P. estoraquensis Henao & Meneses, 2017
Chloreuptychia Forster, 1964 Type species - Papilio chloris Cramer, 1778	C. agatha (Butler, 1867) C. amenthesysta Brévignon & Benmesbah, 2011 C. callichloris (Butler, 1867) C. catharina (Staudinger, [1886]) C. chlorimene (Hübner, [1819]) C. herseis (Godart, [1824]) C. hewitsonii (Butler, 1867) C. marica (Weymer, 1911) C. tolumnia (Cramer, 1777)		Pindis R. Felder, 1869 Type species - Pindis squamistriga R. Felder, 1869	P. squamistriga R. Felder, 1869
Cissia Doubleday, 1848 Type species - Papilio penelope Fabricius, 1775	C. eous (Butler, 1867) C. penelope (Fabricius, 1775) C. phronius (Godart, [1824]) C. pompilia (C. Felder & R. Felder, 1867) C. proba (Butler, 1867) C. rubricata rubricata (Edwards, 1871) C. rubricata anabelae (Miller, 1976) C. rubricata cheneyorum (Chermock, 1949) C. rubricata pseudocleophes (Miller, 1976) C. rubricata smithorum (Wind, 1946) C. touloulou (Benmesbah, 2015) C. labe (Butler, 1870) C. lesbia (Staudinger, [1886]) C. palladia (Butler, 1867) C. joycea (Singer, DeVries & Ehrlich, 1983) C. pseudoconfusa (Singer, DeVries & Ehrlich, 1983) C. similis (Butler, 1867) C. themis (Butler, 1870) C. myncea (Cramer, 1780) C. confusa (Staudinger, 1887) C. maripa (Brévignon, 2005) C. cleophes (Godman & Salvin, 1889) C. terrestris (Butler, 1867) C. terrestris terrestris (Butler, 1867)		Posttaygetis Forster, 1964 Type species - Papilio penelea Cramer, 1777	P. penelea (Cramer, 1777)
Cyllopsis R. Felder, 1869 Type species - Neonympha hedemanni R. Felder, 1869	C. 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 gemma (Hübner, 1808) C. gemma freemani (Stallings & Turner, 1947) C. uatemalena Miller, 1974 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. 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 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 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, 1974		Prenda Freitas & Mielke, 2011 Type species - Prenda clarissa Freitas & Mielke, 2011	P. clarissa Freitas & Mielke, 2011
Erichthodes Forster, 1964 Type species - Neonympha antonina C. Felder & R. Felder, 1867	E. antonina (C. Felder & R. Felder, 1867) E. arius (Weymer, 1911) E. eremita Lamas, Willmott & Radford, 2018 E. jovita C. Felder & R. Felder, 1867 E. julia (Weymer, 1911) E. narapa (Schaus, 1902)		Pseudeuptychia Forster, 1964 Type species - Euptychia languida Butler, 1867	P. cuzquenya Nakahara & Lamas, 2018 P. hemileuca (Staudinger, [1886]) P. languida languida (Butler, 1867) P. languida austrina Nakahara & Lamas, 2018
Euptychia Hübner, 1818 Type species - Euptychia mollina Hübner, 1818	E. alacristata Neild, Nakahara & Fratello, 2014 E. aquila Fratello, Nakahara & Brévigon, 2015 E. atlantica Nakahara & Freitas, 2017 E. attenboroughi Neild, Nakahara, Fratello & Le Crom, 2015 E. audacia Brévignon, Fratello & Nakahara, 2015 E. boulleti (Le Cerf, 1919) E. cesarense cesarense Pulido, Andrade, Peña & Lamas, 2011 E. cesarense obtusa Nakahara, 2015 E. cesarense viloriai Andrade, Pulido, Peña & Lamas, 2011 E. enyita Nakahara, Lamas & Willmott, 2015 E. enyo Butler, 1867 E. favonius Nakahara, Vega & Willmott, 2016 E. fernandae Nakahara & Willmott, 2015 E. fetna Butler, 1870 E. granatina Nakahara, Le Crom & Hall, 2015 E. hannemanni Forster, 1964 E. insolata Butler & H. Druce, 1872 E. jesia Butler, 1869 E. juanjoi Le Crom, Nakahara & Lamas, 2015 E. lacandona Warren & Nakahara, 2015 E. marceli Brévignon, 2005 E. meta Weymer, 1911 E. mollina mollina Hübner, 1818 E. mollina suzannae Brévignon, 2005 E. mollis Staudinger, 1876 E. neblina Warren & Nakahara, 2015 E. neildi Brévignon, 2005 E. padroni Nakahara, Lamas & Willmott, 2015 E. pegasus Nakahara & Hall, 2015 E. picea Butler, 1867 E. pillaca Nakahara & Willmott, 2015 E. roraima Nakahara, Fratello & Harvey, 2014 E. rubrofasciata Miller & Miller, 1988 E. rufocincta Weymer, 1911 E. sophiae Zacca, Nakahara, Dolibaina & Dias, 2015 E. truncata Nakahara & Hall, 2015 E. westwoodi westwoodi Butler, 1867 E. westwoodi muli Brévignon, 2005		Pseudodebis Forster, 1964 Type species - Papilio valentina Cramer, 1779	P. celia (Cramer, 1779) P. dubiosa Forster, 1964 P. euptychidia (Butler, 1868) P. marpessa (Hewitson, 1862) P. puritana (Weeks, 1902) P. valentina (Cramer, 1779) P. vrazi (Kheil, 1896) P. zimri (Butler, 1869)
Euptychoides Forster, 1964 Type species - Euptychia saturnus Butler, 1867	E. albofasciata (Hewitson, 1869) E. castrensis (Schaus, 1902) E. eugenia (C. Felder & R. Felder, 1867) E. eugenia eugenia (C. Felder & R. Felder, 1867) E. eugenia transversa (Weymer, 1911) E. fida (Weymer, 1911) E. hotchikissi (Dyar, 1913) E. laccine (C. Felder & R. Felder, 1867) E. nossis (Hewitson, 1862) E. pseudosaturnus Forster, 1964 E. sanmarcos Nakahara & Lamas, 2018 E. saturnus (Butler, 1867)		Rareuptychia Forster, 1964 Type species - Euptychia clio Weymer, 1911	R. clio (Weymer, 1911)
Forsterinaria Gray, 1973 Type species - Satyrus necys Godart, [1824]	F. anophthalma (C. Felder & R. Felder, 1867) F. antje Peña & Lamas, 2005 F. boliviana (Godman, 1905) F. coipa Peña & Lamas, 2005 F. difficilis (Forster, 1964) F. emo Zubek, Pyrcz & Boyer, 2014 F. enjuerma Peña & Lamas, 2005 F. falcata Peña & Lamas, 2005 F. guaniloi Peña & Lamas, 2005 F. hannieri Zubek & Pyrcz, 2011 F. inornata inornata (C. Felder & R. Felder, 1867) F. inornata magdalena (Hayward, 1957) F. itatiaia Peña & Lamas, 2005 F. necys (Godart, [1824]) F. neonympha (C. Felder & R. Felder, 1867) F. pallida Peña & Lamas, 2005 F. pichita Peña & Lamas, 2005 F. pilosa Peña & Lamas, 2005 F. pronophila (Butler, 1867) F. proxima (Hayward, 1957) F. pseudinornata (Forster, 1964) F. punctata Peña & Lamas, 2005 F. pseudonecys (Strand, 1916) F. pyrczi Peña & Lamas, 2005 F. quantius (Godart, [1824]) F. rotunda Peña & Lamas, 2005 F. rustica rustica (Butler, 1868) F. rustica villarresi (Dognin, 1887) F. stella (Hayward, 1957)		Satyrotaygetis Forster, 1964 Type species - Taygetis satyrina Bates, 1865	S. satyrina (Bates, 1865)
Godartiana Forster, 1964 Type species - Satyrus byses Godart, [1824]	G. amadoi Paluch, Zacca & Freitas, 2016 G. armilla (Butler, 1867) G. astronesthes Lamas & Nakahara, 2018 G. byses (Godart, [1824]) G. luederwaldti (Spitz, 1931) G. muscosa (Butler, 1870)		Sepona Freitas & Barbosa, 2016 Type species - Euptychia punctata Weymer, 1911	S. punctata (Weymer, 1911)
Graphita Nakahara, Marín & Barbosa, 2016 Type species - Neonympha griphe C. Felder & R. Felder, 1867	G. griphe (C. Felder & R. Felder, 1867)		Splendeuptychia Forster, 1964 Type species - Euptychia ashna Hewitson, 1869	S. ackeryi Huertas, Ríos & Le Crom, 2009 S. ambra (Weymer, [1911]) S. ashna (Hewitson, 1869) S. aurigera (Weymer, [1911]) S. boliviensis Forster, 1964 S. clementia (Butler, 1877) S. clorimena (Stoll, 1790) S. cosmophila (Hübner, 1823) S. doxes (Godart, [1824]) S. furina (Hewitson, 1862) S. hygina (Butler, 1877) S. itonis (Hewitson, 1862) S. jadea Brévignon & Benmesbah, 2011 S. junonia (Butler, 1867) S. kendalli Miller, 1976 S. latia (Butler, 1867) S. libitina (Butler, 1870) S. mercedes Huertas, 2011 S. pagyris (Godart, [1824]) S. purusana (Aurivillius, 1929) S. quadrina (Butler, 1869) S. salvini (Butler, 1867) S. telesphora (Butler, 1877) S. toynei Willmott & Hall, 1995 S. triangula (Aurivillius, 1929) S. zischkai Forster, 1964
Harjesia Forster, 1964 Type species - Taygetis blanda Möschler, 1877	H. argentata Nakahara, Zacca & Lamas, 2018 H. blanda (Möschler, 1877) H. obscura (Butler, 1867)		Stegosatyrus Zacca, Mielke & Pyrcz, 2013 Type species - Satyrus periphas Godart, [1824]	S. hemiclara Pyrcz, Boyer & Zacca, 2013 S. imbrialis (Weeks, 1901) S. ocelloides (Schaus, 1902) S. periphas (Godart, [1824])
Hermeuptychia Forster, 1964 Type species - Papilio hermes Fabricius, 1775	H. atalanta (Butler, 1867) H. clara Nakahara, Tan, Lamas & Willmott, 2016 H. cucullina (Weymer, 1911) H. fallax fallax (C. Felder & R. Felder, 1867) H. fallax marinha Anken, 1994 H. gisella (Hayward, 1957) H. harmonia (Butler, 1867) H. hermes (Fabricius, 1775) H. hermybius Grishin, 2014 H. intricata Grishin, 2014 H. maimoune (Butler, 1870) H. pimpla (C. Felder & R. Felder, 1867) H. sosybius (Fabricius, 1793)		Stevenaria Viloria, Costa, Neild & Nakahara, 2016 Type species - Stevenaria yutajensis Viloria & Costa, 2016	S. divergens (Butler, 1867) S. nakaharai Viloria, Costa, Fratello & Neild, 2016 S. yutajensis Viloria & Costa, 2016
Huberonympha Viloria & Costa, 2016 Type species - Huberonympha neildi Viloria, Costa, Fratello & Nakahara, 2016	H. neildi Viloria, Costa, Fratello & Nakahara, 2016		Taydebis Freitas, 2003 Type species - Euptychia peculiaris Butler, 1874	peculiaris Butler, 1874
Inbio Nakahara & Espeland, 2015 Type species - Neonympha hilara C. Felder & R. Felder, 1867	I. hilara (C. Felder & R. Felder, 1867)		Taygetina Forster, 1964 Type species - Taygetis banghaasi Weymer, 1910	banghaasi (Weymer, 1910) kerea (Butler, 1869) oreba (Butler, 1870) peribaea (Godman & Salvin, 1880) weymeri (Draudt, 1912)
Magneuptychia Forster, 1964 Type species - Papilio libye Linneaus, 1767	M. agnata (Schaus, 1913) M. alcinoe (C. Felder & R. Felder, 1867) M. analis (Godman, 1905) M. andrei Zacca, Casagrande & Mielke, 2017 M. drymo (Schaus, 1913) M. flavofascia Zacca & Siewert, 2014 M. francisca (Butler, 1870) M. fugitiva Lamas, [1997] M. gera gera (Hewitson, 1850) M. gera nobilis (Weymer, 1911) M. gera nortia (Hewitson, 1862) M. gomezi (Singer, DeVries & Ehrlich, 1983) M. harpyia harpyia (C. Felder & R. Felder, 1867) M. harpyia batesii (Butler, 1867) M. inani (Staudinger, [1886]) M. iris (C. Felder & R. Felder, 1867) M. kamel Benmesbah & Zacca, 2018 M. lea lea (Cramer, 1777) M. lea philippa (Butler, 1867) M. libye (Linnaeus, 1767) M. lethra (Möschler, 1883) M. louisammour Benmesbah & Zacca, 2018 M. metagera (Butler, 1867) M. mimas (Godman, 1905) M. moderata (Weymer, 1911) M. modesta (Butler, 1867) M. murrayae Brévignon, 2005 M. mycalesis (Röber, 1927) M. nebulosa (Butler, 1867) M. ocnus (Butler, 1867) M. ocypete (Fabricius, 1776) M. opima (Weymer, 1911) M. pallema (Schaus, 1902) M. pax Huertas, Lamas, Fagua & Willmott, 2016 M. probata (Weymer, 1911) M. saul Brévignon & Benmesbah, 2011 M. segesta (Weymer, 1911) M. sheba Brévignon & Benmesbah, 2012 M. tiessa (Hewitson, 1869) M. tricolor tricolor (Hewitson, 1850) M. tricolor fulgora (Butler, 1869) M. trinitensis Brévignon & Benmesbah, 2011		Taygetis Hübner, [1819] Type species - Papilio mermera Cramer, 1776	acuta Weymer, 1910 angulosa Weymer, 1907 asterie Weymer, 1910 chiquitana Forster, 1964 chrysogone Doubleday, [1849] cleopatra C. Felder & R. Felder, 1867 drogoni Siewert, Zacca, Dias & Freitas, 2013 echo (Cramer, 1775) elegia Weymer, 1910 fulginia d'Almeida, 1922 inambari Miller & Lamas, 1999 inconspicua Draudt, 1931 kharisma Brévigon & Benmesbah, 2011 laches (Fabricius, 1793) larua C. Felder & R. Felder, 1867 leuctra Butler, 1870 mermeria (Cramer, 1776) oyapock Brévignon, 2007 rectifascia Weymer, 1907 rufomarginata rufomarginata Staudinger, 1888 rufomarginata tapulunmi Brévignon, 2005 servius Weymer, 1919 sosis Hopffer, 1874 sylvia Bates, 1866 thamyra (Cramer, 1779) tripunctata Weymer, 1907 uncinata Weymer, 1907 uzza Butler, 1869 virgilia (Cramer, 1776) zippora Butler, 1869 ypthima Hübner, [1821]
Megeuptychia Forster, 1964 Type species - Papilio antonoe Cramer, 1775	M. antonoe (Cramer, 1775) M. monopunctata Willmott & Hall, 1995 M. souadae Benmesbah, 2015		Yphthimoides Forster, 1964 Type species - Neonympha yphthima C. Felder & R. Felder, 1867	acmenis (Hübner, 1823) affinis (Butler, 1867) angularis (Butler, 1867) argyrospila (Butler, 1867) bella Barbosa & Freitas, 2015 blanquita Barbosa, Marín & Freitas, 2017 borasta (Schaus, 1902) celmis (Godart, [1824]) cipoensis Freitas, 2004 eriphule (Butler, 1867) gabriela Barbosa, Freitas & Paluch, 2015 iserhardi Freitas & Barbosa, 2015 leguialimai (Dyar, 1913) maepius maepius (Godart, [1824]) maepius guenzeli Anken, 1994 maepius perfuscata (Butler, 1868) manasses (C. Felder & R. Felder, 1867) mimula (Hayward, 1954) mythra (Weymer, 1911) nareta Barbosa & Freitas, 2017 neomaenas (Hayward, 1967) ochracea (Butler, 1867) ordinaria Freitas, Kaminski & Mielke, 2012 pacta (Weymer, 1911) patricia (Hayward, 1957) renata (Stoll, 1780) straminea (Butler, 1867) viviana (Romieux, 1927) yphthima (C. Felder & R. Felder, 1867)
Megisto Hübner, [1819] Type species - Papilio cymela Cramer, 1777	M. cymela (Cramer, 1777)		Zischkaia Forster, 1964 Type species - Euptychia amalda Weymer, 1911	amalda (Weymer, 1911) pacarus (Godart, [1824]) saundersii (Butler, 1867)
Moneuptychia Forster, 1964 Type species - Euptychia soter Butler, 1877	M. giffordi Freitas, Emery & Mielke, 2010 M. griseldis (Weymer, 1911) M. itapeva Freitas, 2007 M. melchiades (Butler, 1877) M. montana Freitas, 2015 M. paeon (Godart, [1824]) M. pervagata Freitas, Siewert & Mielke, 2015 M. soter (Butler, 1877) M. umuarama (Ebert & Dias, 1997) M. vitellina Freitas & Barbosa, 2015 M. wahlbergi Freitas, Barbosa, Siewert & Mielke, 2015		Incertae sedis	insignis (Butler, 1867) ordinata (Weymer, 1911) pellonia (Godman, 1901)
Neonympha Hübner, 1818 Type species - Papilio areolatus Smith, 1797	N. areolatus (Smith, 1797) N. helicta helicta (Hübner, [1808]) N. helicta dadeensis Gatrelle, 1999 N. helicta septentrionalis Davis, 1924 N. mitchellii mitchellii French, 1889 N. mitchellii francisci Parshall & Kral, 1989
Nhambikuara Freitas, Barbosa & Zacca, 2018 Type species - Nhambikuara cerradensis Freitas, Barbosa & Zacca, 2018	N. cerradensis Freitas, Barbosa & Zacca, 2018 N. mima (Butler, 1867)

References

1. 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. doi:10.1111/j.1463-6409.2010.00421.x.
2. Peña, C.; Lamas, G. (2005). "Revision of the butterfly genus Forsterinaria Gray, 1973 (Lepidoptera: Nymphalidae, Satyrinae)". Revista Peruana de Biología. 12 (1): 5–48.
3. Forster, W. (1964). "Beiträge zur Kenntnis der Insektenfauna Boliviens XIX. Lepidoptera III. Satyridae". Veröffentlichungen der Zoologischen Staatssammlung München. 8: 51–188.
4. Miller, L. D. (1968). "The higher classification, phylogeny and zoogeography of the Satyridae (Lepidoptera)". Memoirs of the American Entomological Society. 24: [6] + iii + 174 pp.
5. Nakahara, Shinichi; Lamas, Gerardo; Tyler, Stephanie; Marín, Mario Alejandro; Huertas, Blanca; Willmott, Keith R.; Mielke, Olaf H. H.; Espeland, Marianne (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. doi:10.3897/zookeys.821.31782. PMC 6367311. PMID 30740021.
6. Nakahara, S.; Willmott, K. R.; Mielke, O. H. H.; Schwartz, J.; Zacca, T.; Espeland, M.; Lamas, G. (2018). "Seven new taxa from the butterfly subtribe Euptychiina (Lepidoptera: Nymphalidae: Satyrinae) with revisional notes on Harjesia Forster, 1964 and Pseudeuptychia Forster, 1964". Insecta Mundi (639): 1–38.
7. Lamas, G. (2004). "Checklist: Part 4A. Hesperioidea – Papilionoidea". In Heppner, J. B. (ed.). Atlas of Neotropical Lepidoptera. Volume 5A. Gainesville: Scientific Publishers. pp. 430 pp.
8. Espeland, Marianne; Breinholt, Jesse W.; Barbosa, Eduardo P.; Casagrande, Mirna M.; Huertas, Blanca; Lamas, Gerardo; Marín, Mario A.; Mielke, Olaf H.H.; Miller, Jacqueline Y.; Nakahara, Shinichi; Tan, Denise; Warren, Andrew D.; Zacca, Thamara; Kawahara, Akito Y.; Freitas, André V.L.; Willmott, Keith 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. doi:10.1016/j.ympev.2018.10.039. PMID 30423438.
9. Butler, A. G. (1867). "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". Proceedings of the Zoological Society of London. 1866 (3): 484–504.
10. Butler, A. G. (1877). "On new species of the genus Euptychia, with a tabular view of those hitherto recorded". Zoological Journal of the Linnean Society. 13 (67): 116–128. doi:10.1111/j.1096-3642.1877.tb02375.x.
11. Weymer, G. 1910-1911. 4. Familie: Satyridae. In: Seitz, A. (Ed.), Die Gross-Schmetterlinge der Erde. Stuttgart, A. Kernen. 5: 185-225.
12. Nakahara, S.; Barbosa, E. P.; Freitas, A V L. (2017). "A Potentially Endangered New Species of Euptychia Hübner, 1818 (Lepidoptera: Nymphalidae: Satyrinae) from the Atlantic Coastal Forest of Brazil". Neotropical Entomology. 46 (3): 302–309. doi:10.1007/s13744-016-0466-y. PMID 28063143.
13. Freitas, André V. L.; Barbosa, Eduardo P.; Santos, Jessie P.; Mielke, Olaf H. H. (2013). "A new genus, Atlanteuptychia gen. Nov., for Euptychia ernestina (Lepidoptera: Nymphalidae: Satyrinae)". Zoologia (Curitiba). 30 (6): 661–668. doi:10.1590/S1984-46702013005000005.
14. "Systematics of Euptychiina".
15. 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.
16. 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.
17. Freitas, A. V. L. 2017. Immature stages of the Neotropical satyrine Taygetis acuta (Nymphalidae: Euptychiina). Tropical Lepidoptera Research, 27: 1-5.
18. 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.
19. See, J., Nakahara, S., Gallice, G. 2018 Immature stages of Splendeuptychia quadrina (Butler, 1869) (Lepidoptera: Nymphalidae: Satyrinae). Tropical Lepidoptera Research, 28 (2): 49–53.
20. 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.
21. Singer, M. C., Ehrlich, P. R., Gilbert, L. E. 1971. Butterfly feeding on lycopsid. Science, 172: 1341-1342.
22. 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.
23. 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.