Satyrini

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

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.^[2]

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.^[2]

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.^[2]^[3]

In contrast, the tribe has a few genera which show uncommon feeding preferences.^[3] 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.^[4]

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,^[5] and later studies have largely followed this classification.^[6]^[7] Thus, the Satyrini is now considered to include the genera previously placed in the Elymniini, such as Lethe, Pararge and Mycalesis.^[1] However, the phylogenetic statuses of some subtribes within the tribe are still unclear and the revision of the classification is likely to continue.^[7]

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^[a])
	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	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
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

Notes

^ Lamas (2004) was excluded because it only listed genera distributed in the Neotropics. Also, note that the table does not include all subtribes of this tribe (e.g. there are not subtribes Eritina, Ragadiina and tribe Dirini), and some genera have been further reclassified (e.g. Aeropetes and Paralethe from Melanitini to Dirini, Orsotriaena and Zipaetis from Coenonymphina to Eritina) or synonymized (e.g. Satyrodes was synonymized with Lethe), since Peña et al. (2006).^[1]

References

Citations

^ ^a ^b ^c Wahlberg 2019.
^ ^a ^b ^c ^d PEÑA, NYLIN & WAHLBERG 2011.
^ ^a ^b Peña & Wahlberg 2008.
^ Freitas et al. 2019.
^ Peña et al. 2006.
^ Chen et al. 2020.
^ ^a ^b Marín et al. 2011.

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)

[8] Lamas (2004) was excluded because it only listed genera distributed in the Neotropics. Also, note that the table does not include all subtribes of this tribe (e.g. there are not subtribes Eritina, Ragadiina and tribe Dirini), and some genera have been further reclassified (e.g. Aeropetes and Paralethe from Melanitini to Dirini, Orsotriaena and Zipaetis from Coenonymphina to Eritina) or synonymized (e.g. Satyrodes was synonymized with Lethe), since Peña et al. (2006).^[1]

[FOOTNOTEWahlberg2019-1] Wahlberg 2019.

[FOOTNOTEPEÑANYLINWAHLBERG2011-2] PEÑA, NYLIN & WAHLBERG 2011.

[FOOTNOTEPeñaWahlberg2008-3] Peña & Wahlberg 2008.

[FOOTNOTEFreitasMotaBarbosaCarreira2019-4] Freitas et al. 2019.

[FOOTNOTEPeñaWahlbergWeingartnerKodandaramaiah2006-5] Peña et al. 2006.

[FOOTNOTEChenWahlbergLiaoWang2020-6] Chen et al. 2020.

[FOOTNOTEMarínPeñaFreitasWahlberg2011-7] Marín et al. 2011.

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