# Satyrini

The Satyrini is one of the tribes of the subfamily Satyrinae. It includes about 2200 species and is therefore the largest tribe in the subfamily which comprises 2500 species.

## Distribution

Satyrini butterflies have a worldwide distribution, but the distribution pattern differs between subtribes. Some subtribes are almost restricted to a single biogeographic region, such as the Pronophilina, which is found only in Andean cloud forests from Venezuela to Bolivia.

## Biology

The larval food plants of many species in this tribe are grasses, i.e. Poaceae. It is considered that the Satyrini diversified at about the same time as the grasses did, and that the radiation of the tribe is therefore closely related to the evolution of the grasses.
In contrast, the tribe has a few genera which show uncommon feeding preferences. Three genera, Euptychia, Ragadia and Acrophtalmia, feed on Lycopsida, and moreover, some species of Euptychia have been reported to feed on mosses of Neckeraceae. This is interesting cases as these genera are not considered to be very closely related to each other.

## Taxonomy

The taxonomy of the subfamily Satyrinae has recently undergone a significant revision. Peña et al. (2006) showed that most of the traditionally well known group are paraphyletic or polyphyletic and developed a new taxonomy, based on molecular phylogenetic. The classification scheme transferred many genera traditionally placed in the Elymniini to the Satyrini, and later studies have largely followed this classification. Thus, the Satyrini is now considered to include the genera previously placed in the Elymniini, such as Lethe, Pararge and Mycalesis. However, the phylogenetic statuses of some subtribes within the tribe are still unclear and the revision of the classification is likely to continue.
According to Wahlberg (2019), the Satyrini includes the following subtribes and genera:

### Parargina Moore, 1893

   * Chonala Moore, 1893
   * Kirinia Westwood, 1841
   * Lasiommata Tutt, 1896
   * Lopinga Moore, 1893
   * Orinoma Gray, 1846
   * Pararge Hübner, 1819
   * Rhaphicera Butler, 1867
   * Tatinga Moore, 1893
   * Nosea Koiwaya 1993

### Lethina Reuter, 1896

   * Aphysoneura Karsch, 1894
   * Lethe Hübner, 1819
   * Neope Moore, 1866
   * Ninguta Moore, 1892
   * Ptychandra Felder & Felder, 1861
   * Mandarinia Leech, 1892

### Mycalesina Reuter, 1896

   * Lohora Moore, 1880
   * Brakefieldia Aduse-Poku, Lees & Wahlberg, 2016
   * Telinga Moore, 1880
   * Heteropsis Westwood, 1850
   * Culapa Moore, 1879
   * Mydosama Moore, 1880
   * Mycalesis Hübner, 1818
   * Devyatkinia Monastyrskii & Uémura, 2016
   * Hallelesis Condamin, 1961
   * Bicyclus Kirby, 1871

### Eritina Miller, 1968

   * Coelites Westwood, 1850
   * Erites Westwood, 1851
   * Orsotriaena Wallengren, 1858
   * Zipaetis Hewitson, 1863

### Ragadiina Herrich-Schäffer, 1864

   * Acrophtalmia Felder & Felder, 1861
   * Acropolis Hemming, 1934
   * Ragadia Westwood, 1851

### Coenonymphina Tutt, 1896

   * Coenonympha Hübner, 1819
   * Sinonympha Lee, 1994
   * Oressinoma Doubleday, 1849
   * Altiapa Parsons, 1986
   * Argynnina Butler, 1867
   * Argyronympha Mathew, 1886
   * Argyrophenga Doubleday, 1845
   * Dodonidia Butler, 1884
   * Erebiola Fereday, 1879
   * Erycinidia Rothschild & Jordan, 1905
   * Geitoneura Butler, 1867
   * Harsiesis Fruhstorfer, 1911
   * Heteronympha Wallengren, 1858
   * Hyalodia Jordan, 1924
   * Hypocysta Westwood, 1851
   * Lamprolenis Godman & Salvin, 1881
   * Nesoxenica Waterhouse & Lyell, 1914
   * Oreixenica Waterhouse & Lyell, 1914
   * Paratisiphone Watkins, 1928
   * Percnodaimon Butler, 1876
   * Platypthima Rothschild & Jordan, 1905
   * Tisiphone Hübner, 1819

### Euptychiina Reuter, 1896

   * Euptychia Hübner, 1818
   * Palaeonympha Butler, 1871
   * Amphidecta Butler, 1867
   * Archeuptychia Forster, 1964
   * Amiga Nakahara, Willmott & Espeland, 2019
   * Caeruleuptychia Forster, 1964
   * Capronnieria Forster, 1964
   * Cepheuptychia Forster, 1964
   * Chloreuptychia Forster, 1964
   * Cissia Doubleday, 1848
   * Coeruleotaygetis Forster, 1964
   * Cyllopsis Felder, 1869
   * Erichthodes Forster, 1964
   * Euptychoides Forster, 1964
   * Forsterinaria Gray, 1973
   * Graphita Nakahara, Marín & Barbosa 2016
   * Godartiana Forster, 1964
   * Harjesia Forster, 1964
   * Hermeuptychia Forster, 1964
   * Huberonympha Viloria & Costa, 2016
   * Inbio Nakahara & Espeland, 2015
   * Magneuptychia Forster, 1964
   * Megeuptychia Forster, 1964
   * Megisto Hübner, 1819
   * Moneuptychia Forsterm, 1964
   * Neonympha Hübner 1818
   * Nhambikuara Freitas, Barbosa & Zacca, 2018
   * Optimandes Marín, Nakahara & Willmott, 2019
   * Orotaygetis Nakahara & Zacca, 2018
   * Paramacera Butler, 1868
   * Parataygetis Forster, 1964
   * Pareuptychia Forster, 1964
   * Paryphthimoides Forster, 1964
   * Pharneuptychia Forster, 1964
   * Pindis Felder, 1869
   * Posttaygetis Forster, 1964
   * Prenda Freitas & Mielke, 2011
   * Pseudeuptychia Forster, 1964
   * Pseudodebis Forster, 1964
   * Rareuptychia Forster, 1964
   * Satyrotaygetis Forster, 1964
   * Sepona Freitas & Barbosa, 2016
   * Splendeuptychia Forster, 1964
   * Stegosatyrus Zacca, Mielke & Pyrcz, 2013
   * Stevenaria Viloria, Costa, Neild & Nakahara, 2016
   * Taydebis Freitas, 2003
   * Taygetina Forster, 1964
   * Taygetis Hübner, 1819
   * Taygetomorpha Miller, 2004
   * Yphthimoides Forster, 1964
   * Zischkaia Forster, 1964

### Erebiina Tutt, 1896

   * Erebia Elwes, 1899

### Maniolina Grote, 1897

   * Aphantopus Wallengren, 1853
   * Cercyonis Scudder, 1875
   * Hyponephele Muschamp, 1915
   * Maniola Schrank, 1801
   * Pyronia Hübner, 1819

### Melanargiina Wheeler, 1903

   * Melanargia Meigen, 1828

### Pronophilina Reuter, 1896

   * Altopedaliodes Forster, 1964
   * Antopedaliodes Forster, 1964
   * Apexacuta Pyrcz, 2004
   * Arhuaco Adams & Bernard, 1977
   * Cheimas Thieme, 1907
   * Corades Doubleday, 1849
   * Corderopedaliodes Forster, 1964
   * Daedalma Hewitson, 1858
   * Dangond Adams & Bernard, 1979
   * Drucina Butler, 1872
   * Druphila Pyrcz, 2004
   * Eretris Thieme, 1905
   * Eteona Doubleday, 1848
   * Foetterleia Viloria, 2004
   * Junea Hemming, 1964
   * Lasiophila Felder & Felder, 1859
   * Lymanopoda Westwood, 1851
   * Mygona Thieme, 1907
   * Neopedaliodes Viloria, Miller & Miller 2004
   * Oxeoschistus Butler, 1867
   * Panyapedaliodes Forster, 1964
   * Paramo Adams & Bernard, 1977
   * Parapedaliodes Forster, 1964
   * Pedaliodes Butler, 1867
   * Pherepedaliodes Forster, 1964
   * Physcopedaliodes Forster, 1964
   * Praepedaliodes Forster, 1964
   * Praepronophila Forster, 1964
   * Proboscis Thieme, 1907
   * Pronophila Doubleday, 1849
   * Protopedaliodes Viloria & Pyrcz, 1994
   * Pseudomaniola Röber, 1889
   * Punapedaliodes Forster, 1964
   * Steremnia Thieme, 1905
   * Steroma Westwood, 1850
   * Steromapedaliodes Forster, 1964
   * Thiemeia Weymer, 1912
   * Diaphanos Adams & Bernard, 1981
   * Ianussiusa Pyrcz & Viloria, 2004
   * Idioneurula Strand, 1932
   * Manerebia Staudinger, 1897
   * Neomaniola Hayward, 1949
   * Argyrophorus Blanchard, 1852
   * Auca Hayward, 1953
   * Chillanella Herrera, 1966
   * Cosmosatyrus Felder & Felder, 1867
   * Elina Blanchard, 1852
   * Faunula Felder & Felder, 1867
   * Haywardella Herrera, 1966
   * Homoeonympha Felder & Felder, 1867
   * Nelia Hayward, 1953
   * Neomaenas Wallengren, 1858
   * Neosatyrus Wallengren, 1858
   * Pampasatyrus Hayward, 1953
   * Quilaphoestosus Herrera, 1966
   * Spinantenna Hayward, 1953
   * Tetraphlebia Felder & Felder, 1867
   * Gyrocheilus Butler, 1867

### Satyrina Boisduval, 1833

   * Argestina Riley, 1923
   * Davidina Oberthür, 1879
   * Berberia de Lesse, 1951
   * Hipparchia Fabricius, 1807
   * Kanetisa Moore, 1893
   * Aulocera Butler, 1867
   * Karanasa Moore, 1893
   * Oeneis Hübner, 1819
   * Paroeneis Moore, 1893
   * Arethusana de Lesse, 1951
   * Brintesia Fruhstorfer, 1911
   * Minois Hübner, 1819
   * Chazara Moore, 1893
   * Pseudochazara de Lesse, 1951
   * Satyrus Latreille, 1810

### Ypthimina Reuter, 1896

   * Cassionympha van Son, 1955
   * Coenyra Hewitson, 1865
   * Coenyropsis van Son, 1958
   * Mashuna van Son, 1955
   * Mashunoides Mendes & Bivar de Sousa, 2009
   * Melampias Hübner, 1819
   * Neita van Son, 1955
   * Neocoenyra Butler, 1886
   * Pseudonympha Wallengren, 1857
   * Paternympha Henning & Henning, 1997
   * Physcaeneura Wallengren, 1857
   * Strabena Mabille, 1877
   * Stygionympha van Son, 1955
   * Ypthima Hübner, 1818
   * Austroypthima Holloway, 1974
   * Ypthimomorpha van Son, 1955

### subtribe uncertain

   * Calisto Hübner, 1823
   * Paralasa Moore, 1893
   * Callerebia Butler, 1867
   * Proterebia Roos & Amschied, 1980
   * Loxerebia Watkins, 1925
   * Dyndirus Capronnier, 1874
Table for representative higher level classifications of satyrines in Peña et al. (2006) (excerpts with some modifications) :
| | Miller (1968) | | Harvey (1991) | Peña et al. (2006) |
| Family | Satyridae | Subfamily | Satyrinae | Satyrinae |
| Subfamily | Elymniinae | Tribe | Elymniini | Elymniini |
| Tribe | Elymniini | Subtribe | Elymniiti | Elymnias |
| Genera | Elymnias Elymniopsis | Genera | Elymnias Elymniopsis | Elymnias |
| Tribe | Lethini | Subtribe | Lethiti | Elymnias |
| Genera | Aeropetes Paralethe Enodia Lethe Neope Satyrodes Kirinia Lasiommata Lopinga Pararge Ethope Neorina | Genera | Aeropetes Paralethe Enodia Lethe Neope Satyrodes Kirinia Lasiommata Lopinga Pararge Ethope Neorina | Elymnias |
| Tribe | Mycalesini | Subtribe | Mycalesiti | Elymnias |
| Genera | Bicyclus Hallelesis Henotesia Mycalesis Orsotriaena | Genera | Bicyclus Hallelesis Henotesia Mycalesis Orsotriaena | Elymnias |
| Tribe | Zetherini | Subtribe | Zetheriti | Elymnias |
| Genus | Zethera | | Zethera | Elymnias |
| Genus | Zethera | Tribe | Zethera | Zetherini |
| Genus | Zethera | Genera | Zethera | Neorina Penthema Ethope Zethera |
| Subfamily | Biinae | Tribe | Biini | Melanitini |
| Tribe | Melanitini | Subtribe | Melanititi | Aeropetes Paralethe Manataria Gnophodes Melanitis |
| Genera | Gnophodes Melanitis | Genera | Gnophodes Melanitis | Aeropetes Paralethe Manataria Gnophodes Melanitis |
| Tribe | tribe uncertain | Tribe | tribe uncertain | Aeropetes Paralethe Manataria Gnophodes Melanitis |
| Genus | Manataria | Genera | Manataria | Aeropetes Paralethe Manataria Gnophodes Melanitis |
| Subfamily | Haeterinae | Tribe | Haeterini | Haeterini |
| Tribe | Haeterini | Genera | Cithaerias Haetera Pierella Pseudohaetera | Cithaerias Haetera Pierella Pseudohaetera |
| Genera | Cithaerias Haetera Pierella Pseudohaetera | Genera | Cithaerias Haetera Pierella Pseudohaetera | Cithaerias Haetera Pierella Pseudohaetera |
| Subfamily | Satyrinae | Tribe | Satyrini | Satyrini |
| | | Subtribe | | Parargina |
| | | Genera | | Kirinia Lopinga Lasiommata Pararge |
| | | Subtribe | | Lethina |
| | | Genera | | Lethe Enodia Satyrodes Neope |
| | | Subtribe | | Mycalesina |
| | | Genera | | Bicyclus Hallelesis Henotesia Mycalesis |
| Tribe | Coenonymphini | Subtribe | Coenonymphiti | Coenonymphina |
| Genera | Coenonympha Aphantopus | Genera | Coenonympha Aphantopus | Oreixenica Tisiphone Nesoxenica Hypocysta Lamprolenis Dodonidia Argyrophenga Erebiola Percnodaimon Heteronympha Geitoneura Oressinoma Coenonympha Orsotriaena Zipaetis Argyronympha |
| Tribe | Hypocystini | Subtribe | Hypocystiti | Oreixenica Tisiphone Nesoxenica Hypocysta Lamprolenis Dodonidia Argyrophenga Erebiola Percnodaimon Heteronympha Geitoneura Oressinoma Coenonympha Orsotriaena Zipaetis Argyronympha |
| Genera | Argyrophenga Dodonidia Erebiola Geitoneura Heteronympha Hypocysta Lamprolenis Nesoxenica Oreixenica Percnodaimon Tisiphone Zipaetis | Genera | Argyrophenga Dodonidia Erebiola Geitoneura Heteronympha Hypocysta Lamprolenis Nesoxenica Oreixenica Percnodaimon Tisiphone Zipaetis | Oreixenica Tisiphone Nesoxenica Hypocysta Lamprolenis Dodonidia Argyrophenga Erebiola Percnodaimon Heteronympha Geitoneura Oressinoma Coenonympha Orsotriaena Zipaetis Argyronympha |
| Tribe | Euptychiini | Subtribe | Euptychiiti | Euptychiina |
| Genera | Caeruleuptychia Cepheuptychia Chloreuptychia Cissia Cyllopsis Erichthodes Euptychia Euptychoides Forsterinaria Godartiana Harjesia Hermeuptychia Moneuptychia Neonympha Oressinoma Paramacera Parataygetis Pareuptychia Paryphthimoides Pharneuptychia Pindis Posttaygetis Oressinoma Rareuptychia Splendeuptychia Taygetis Yphthimoides | Genera | Caeruleuptychia Cepheuptychia Chloreuptychia Cissia Cyllopsis Erichthodes Euptychia Euptychoides Forsterinaria Godartiana Harjesia Hermeuptychia Moneuptychia Neonympha Oressinoma Paramacera Parataygetis Pareuptychia Paryphthimoides Pharneuptychia Pindis Posttaygetis Oressinoma Rareuptychia Splendeuptychia Taygetis Yphthimoides | Euptychia Cyllopsis Paramacera Palaeonympha Pharneuptychia Euptychoides Yphthimoides Moneuptychia Paryphthimoides Amphidecta Rareuptychia Godartiana Hermeuptychia Splendeuptychia Pindis Cepheuptychia Cissia Caeruleuptychia Magneuptychia Chloreuptychia Neonympha Erichthodes Pareuptychia Taygetis Harjesia Parataygetis Posttaygetis Forsterinaria |
| Tribe | tribe uncertain | Subtribe | subtribe uncertain | subtribe uncertain |
| Genus | Palaeonympha | Genera | Palaeonympha | Cercyonis Hyponephele Neocoenyra |
| Tribe | Ypthimini | Subtribe | Ypthimiti | Ypthimina |
| Genera | Neocoenyra Ypthima Ypthimomorpha | Genera | Neocoenyra Ypthima Ypthimomorpha | Paralasa Ypthima Ypthimomorpha |
| Tribe | Melanargiini | Subtribe | Melanargiiti | Melanargiina |
| Genus | Melanargia | Genus | Melanargia | Melanargia |
| Tribe | Maniolini | Subtribe | Manioliti | Maniolina |
| Genera | Cercyonis Hyponephele Maniola Pyronia | Genera | Cercyonis Hyponephele Maniola Pyronia | Pyronia Maniola Aphantopus |
| Tribe | Pronophilini | Subtribe | Pronophiliti | Pronophilina |
| Genera | Amphidecta Corades Daedalma Eteona Junea Lasiophila Lymanopoda Oxeoschistus Panyapedaliodes Parapedaliodes Pedaliodes Praepedaliodes Proboscis Pronophila Pseudomaniola Punapedaliodes Steremnia Steroma Idioneurula Manerebia Argyrophorus Quilaphoetosus Auca Chillanella Cosmosatyrus Elina Etcheverrius Nelia Pampasatyrus | Genera | Amphidecta Corades Daedalma Eteona Junea Lasiophila Lymanopoda Oxeoschistus Panyapedaliodes Parapedaliodes Pedaliodes Praepedaliodes Proboscis Pronophila Pseudomaniola Punapedaliodes Steremnia Steroma Idioneurula Manerebia Argyrophorus Quilaphoetosus Auca Chillanella Cosmosatyrus Elina Etcheverrius Nelia Pampasatyrus | Nelia Steremnia Steroma Manerebia Idioneurula Tamania Ianussiusa Lymanopoda Argyrophorus Etcheverrius Pampasatyrus Elina Quilaphoetosus Cosmosatyrus Chillanella Auca Panyapedaliodes Pedaliodes Punapedaliodes Praepedaliodes Corades Junea Pronophila Eteona Foetterleia Daedalma Oxeoschistus Proboscis Lasiophila Apexacuta Pseudomaniola |
| Tribe | Erebiini | Subtribe | Erebiiti | Erebiina |
| Genus | Erebia | Genus | Erebia | Erebia |
| Tribe | Satyrini | Subtribe | Satyriti | Satyrina |
| Genera | Arethusana Berberia Brintesia Chazara Hipparchia Karanasa Neominois Oeneis Paralasa Pseudochazara Satyrus | Genera | Arethusana Berberia Brintesia Chazara Hipparchia Karanasa Neominois Oeneis Paralasa Pseudochazara Satyrus | Berberia Hipparchia Chazara Pseudochazara Satyrus Arethusana Brintesia Karanasa Neominois Oeneis |
| | | Tribe | tribe uncertain | |
| | | Genera | Penthema | |