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.

Satyrini
Satyrus actaea, Languedoc-Roussillon, France
Lethe appalachia (syn. Satyrodes appalachia), Virginia, United States
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Lepidoptera
Family:	Nymphalidae
Subfamily:	Satyrinae
Tribe:	Satyrini Boisduval, 1833
Subtribes
Parargina Tutt, 1896 Lethina Reuter, 1896 Mycalesina Reuter, 1896 Eritina Miller, 1968 Ragadiina Herrich-Schäffer, 1864 Coenonymphina Tutt, 1896 Euptychiina Reuter, 1896 Erebiina Tutt, 1896 Maniolina Grote, 1897 Melanargiina Wheeler, 1903 Pronophilina Reuter, 1896 Satyrina Boisduval, 1833 Ypthimina Reuter, 1896 some genera incertae sedis
Diversity
approximately 220 genera, 2200 species

Toothed satyr
Praepronophila petronius, Colombia

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

See also: Elymniini

Pararge aegeria ♂, Portugal

Lethe europa pavida, Taiwan

Mycalesis francisca formosana, Taiwan

Coelites epiminthia, Malaysia

Ragadia makuta, Malaysia

Coenonympha oedippus, Japan

Euptychia westwoodi, Panama

Erebia ligea takanonis, Japan

Maniola telmessia, Turkey

Melanargia galathea, UK

Lasiophila sp., Peru

Ypthima baldus, India

Calisto herophile, Cuba

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.

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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
Tribe	Lethini		Subtribe	Lethiti
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
Tribe	Mycalesini		Subtribe	Mycalesiti
Genera	Bicyclus Hallelesis Henotesia Mycalesis Orsotriaena		Genera	Bicyclus Hallelesis Henotesia Mycalesis Orsotriaena
Tribe	Zetherini		Subtribe	Zetheriti
Genus	Zethera			Zethera
			Tribe		Zetherini
			Genera		Neorina Penthema Ethope Zethera
Subfamily	Biinae		Tribe	Biini	Melanitini
Tribe	Melanitini		Subtribe	Melanititi	Aeropetes Paralethe Manataria Gnophodes Melanitis
Genera	Gnophodes Melanitis		Genera	Gnophodes Melanitis
Tribe	tribe uncertain		Tribe	tribe uncertain
Genus	Manataria		Genera	Manataria
Subfamily	Haeterinae		Tribe	Haeterini	Haeterini
Tribe	Haeterini		Genera	Cithaerias Haetera Pierella Pseudohaetera	Cithaerias Haetera Pierella Pseudohaetera
Genera	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
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
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

References

Sources

• Chen, Lu; Wahlberg, Niklas; Liao, Cheng-Qing; Wang, Chen-Bin; Ma, Fang-Zhou; Huang, Guo-Hua (2020). "Fourteen complete mitochondrial genomes of butterflies from the genus Lethe (Lepidoptera, Nymphalidae, Satyrinae) with mitogenome-based phylogenetic analysis". Genomics. 112 (6): 4435–4441. doi:10.1016/j.ygeno.2020.07.042. PMID 32745503. S2CID 220966325.
• Freitas, André V.L.; Mota, Luísa L.; Barbosa, Eduardo P.; Carreira, Junia Y.O. (2019). "Immature stages of the Selaginella-feeding Euptychia mollina (Nymphalidae: Satyrinae) from the Brazilian Amazon". Zoologia. 36: 1–7. doi:10.3897/zoologia.36.e32053. S2CID 195398071.
• Harvey, D.J. (1991). "Higher classification of the Nymphalidae, Appendix B.". In Nijhout, H.F. (ed.). The Development and Evolution of Butterfly Wing Patterns. Smithsonian Institution Press. pp. 255–273.
• Lamas, G. (2004). Checklist: Part 4A. Hesperioidea-Papilionoidea. Atlas of neotropical Lepidoptera. Association for Tropical Lepidoptera/Scientific Publishers. ISBN 0945417284. OCLC 58754755.
• Marín, M.A.; Peña, C.; Freitas, A.V.L.; Wahlberg, N.; Uribe, S.I. (2011). "From the Phylogeny of the Satyrinae Butterflies to the Systematics of Euptychiina (Lepidoptera: Nymphalidae): History, Progress and Prospects". Neotropical Entomology. 40 (1): 1–13. doi:10.1590/S1519-566X2011000100001. PMID 21437476.
• Miller, Lee D. (1968). "The higher classification, phylogeny and zoogeography of the Satyridae (Lepidoptera)". Memoirs of the American Entomological Society. 24: 1–174.
• Peña, Carlos; Wahlberg, Niklas; Weingartner, Elisabet; Kodandaramaiah, Ullasa; Nylin, Sören; Freitas, André V.L.; Brower, Andrew V.Z. (2006). "Higher level phylogeny of Satyrinae butterflies (Lepidoptera: Nymphalidae) based on DNA sequence data" (PDF). Molecular Phylogenetics and Evolution. 40 (1): 29–49. Bibcode:2006MolPE..40...29P. doi:10.1016/j.ympev.2006.02.007. PMID 16563805.
• Peña, Carlos; Wahlberg, Niklas (2008). "Prehistorical climate change increased diversification of a group of butterflies" (PDF). Biology Letters. 4 (3): 274–278. doi:10.1098/rsbl.2008.0062. PMC 2610051. PMID 18364308.
  • Peña, Carlos; Wahlberg, Niklas (2008). "ELECTRONIC SUPPLEMENTARY MATERIAL" (PDF).
• PEÑA, CARLOS; NYLIN, SÖREN; WAHLBERG, NIKLAS (2011). "The radiation of Satyrini butterflies (Nymphalidae: Satyrinae): a challenge for phylogenetic methods". Zoological Journal of the Linnean Society. 161 (1): 64–87. doi:10.1111/j.1096-3642.2009.00627.x.
• Wahlberg, Niklas. "The Nymphalidae Systematics Group". Retrieved 2021-10-21.
  • Wahlberg, Niklas (2019). "The higher classification of Nymphalidae". NYMPHALIDAE.net. Retrieved 2021-10-21.

External links

• Satyrinae at NYMPHALIDAE.net (There is no individual page for Satyrini as it is split into pages for each subtribes.)
• Satyrini Boisduval 1833 at TREE OF LIFE web project
• ジャノメチョウ族（Satyrini）のページ at Pteron World (in Japanese)