Carlos Peña & Niklas Wahlberg

Eretris sp.

Satyrinae is the biggest subfamily in the Nymphalidae comprising 2848 species of worldwide distribution. According to the latest version of the Classification of Nymphalidae, these are placed in 261 genera and 9 tribes. As currently delimited, the subfamily includes the tribes Morphini, Brassolini and Amathusiini, which have traditionally been placed in their own subfamily. However, almost all molecular studies show that the three tribes are within Satyrinae. The taxonomy of the subfamily is quite chaotic at the moment and needs to be revised, though the pioneering work of Miller in 1968 has helped much in creating some sort of order. The phylogenetic relationships of the subfamily are slowly being resolved.

Satyrinae butterflies, or satyrines, are worldwide distributed, including all continents and even remote isolated places like the Greater Antilles, New Caledonia, New Zealand and Fiji islands.

As larvae, virtually all satyrines feed on monocot plants, with the striking exception of two genera that do not behave as proper butterflies since they do not feed on angiosperms, feeding instead on lower plants from the Lycopodiophyta division. These are the only butterflies exhibiting this behavior, one group is the Neotropical genus Euptychia and the other is the Oriental genus Ragadia. Even more bizarre is the fact that some Euptychia species feed on mosses¹, departing from every established rule in the "how to be a good butterfly" manual.

Despite being such a big group of butterflies, there is little general knowledge on the Satyrinae and most of what we know is due to anecdotal data available from widely sparse scientific literature. This is due to the lack of interest in the study of satyrines, probably because satyrines exhibit mainly brownish coloration, being regarded as very dull butterflies when compared with other Nymphalidae. However, some satyrines have not been totally neglected. The Palearctic Pararge is used as model organism for ecological studies and biogeography, members of Erebia have been the subject of phylogeographic studies and the African Bicyclus anynana is heavily used as model organism in evo-devo studies, particularly in the area of interactions between evolutionary and developmental processes.

The tribes of Satyrinae


Key publications for Satyrinae systematics:

  1. Bozano, G. C. 2002. Satyrinae part III. Pages 1-71 in Guide to the Butterflies of the Palearctic Region (G. C. Bozano, ed.) Omnes Artes, Milano.
  2. Brooks, C. J. 1950. A revision of the genus Tenaris Hübner (Lepidoptera: Amathusiidae). Trans. R. Ent. Soc. Lond. 101: 179-238.
  3. Della Bruno, C., E. Gallo, M. Lucarelli, and V. Sbordoni. 2002. Satyrinae part II. Pages 1-59 in Guide to the Butterflies of the Palearctic Region (G. C. Bozano, ed.) Omnes Artes, Milano.
  4. DeVries, P. J., I. J. Kitching, and R. I. Vane-Wright. 1985. The systematic position of Antirrhea and Caerois, with comments on the classification of the Nymphalidae (Lepidoptera). Systematic Entomology 10:11-32.
  5. Freitas, A. V. L. 2002. Immature stages of Eteona tisiphone (Nymphalidae: Satyrinae). Journal of the Lepidopterists' Society 56:286-288.
  6. Freitas, A. V. L. 2003. Description of a new genus for "Eupthychia" peculiaris (Nymphalidae: Satyrinae): immature stages and systematic position. Journal of the Lepidopterists' Society 57:100-106.
  7. Freitas, A. V. L. 2004. Immature stages of Amphidecta reynoldsi (Nymphalidae: Satyrinae). Journal of the Lepidopterists' Society 58:53-55.
  8. Freitas, A. V. L. 2004. A new species of Yphthimoides (Nymphalidae, Satyrinae) from southeastern Brazil. Journal of the Lepidopterists' Society 58:7-12.
  9. Freitas, A. V. L., W. W. Benson, O. J. Marini-Filho, and R. M. de Carvalho. 1995. Territoriality by the dawn's early light: the Neotropical owl butterfly Caligo idomenaeus (Nymphalidae: Brassolinae). Journal of Research on the Lepidoptera 34:14-20.
  10. Freitas, A. V. L., D. Murray, and K. S. J. Brown. 2002. Immatures, natural history and the systematic position of Bia actorion (Nymphalidae). Journal of the Lepidopterists' Society 56:117-122.
  11. Freitas, A. V. L., and C. Peña. 2006. Description of Genus Guaianaza for "Euptychia" pronophila (Lepidoptera: Nymphalidae: Satyrinae) with a description of the immature stages. Zootaxa 1163:49-59.
  12. Jordan, K. 1924. On Hypocysta and some allied genera of Satyrinae (Lepidoptera: Rhopalocera) from New Guinea and the Solomon Islands. Novitates Zoologica 31: 279-297.
  13. Martin, J.-F., A. Gilles, and H. Descimon. 2000. Molecular phylogeny and evolutionary patterns of the European satyrids (Lepidoptera: Satyridae) as revealed by mitochondrial gene sequences. Molecular Phylogenetics and Evolution 15:70-82.
  14. Matos-Maravi, P. F., Peña, C., Willmott, K. R., Freitas, A. V. L. & Wahlberg, N. 2013: Systematics and evolutionary history of butterflies in the "Taygetis clade" (Nymphalidae: Satyrinae: Euptychiina): towards a better understanding of Neotropical biogeography. Molecular Phylogenetics and Evolution 66: 54-68. doi:10.1016/j.ympev.2012.09.005
  15. Miller, L. D. 1968. The higher classification, phylogeny, and zoogeography of the Satyridae. Memmoirs of the American Entomological Society, Philadelphia.
  16. Murray, D., and D. Pashley Prowell. 2005. Molecular phylogenetics and evolutionary history of the neotropical Satyrine subtribe Euptychiina (Nymphalidae: Satyrinae). Molecular Phylogenetics and Evolution 34:67-80.
  17. Parsons, M. J. 1999. The Butterflies of Papua New Guinea: Their Systematics and Biology. Academic Press, London.
  18. Peña, C., and G. Lamas. 2005. Revision of the butterfly genus Forsterinaria Gray, 1973 (Lepidoptera: Nymphalidae, Satyrinae). Revista peruana biologia 12:5-48.
  19. Peña, C., N. Wahlberg, E. Weingartner, U. Kodandaramaiah, S. Nylin, A. V. L. Freitas, and A. V. Z. Brower. 2006. Higher level phylogeny of Satyrinae butterflies (Lepidoptera: Nymphalidae) based on DNA sequence data. Molecular Phylogenetics and Evolution 40:29-49.
  20. Peña, C., Nylin, S., Freitas, A. V. L. & Wahlberg, N. 2010: Biogeographic history of the butterfly subtribe Euptychiina (Lepidoptera, Nymphalidae, Satyrinae). Zoologica Scripta 39: 243-258. doi:10.1111/j.1463-6409.2010.00421.x pdf
  21. Peña, C., Nylin, S. & Wahlberg, N. 2011: The radiation of Satyrini butterflies (Nymphalidae: Satyrinae): a challenge for phylogenetic methods. Zoological Journal of the Linnean Society 161: 64-87. doi:10.1111/j.1096-3642.2009.00627.x
  22. Penz, C. M., and P. J. DeVries. 2002. Phylogenetic analysis of Morpho butterflies (Nymphalidae, Morphinae): implications for classification and natural history. American Museum Novitates 3374: 1-33.
  23. Penz, C., DeVries, P. J. & Wahlberg, N. 2012: Diversification of Morpho butterflies (Lepidoptera, Nymphalidae): a re-evaluation of morphological characters and new insight from DNA sequence. Systematic Entomology 37: 670-685. doi:10.1111/j.1365-3113.2012.00636.x
  24. ¹Singer C. and Mallet J. 1986. Moss-Feeding by a Satyrine butterfly. Journal of Research on the Lepidoptera 24(4): 392.
  25. Torres, E., D. C. Lees, R. I. Vane-Wright, C. Kremen, J. A. Leonard, and R. K. Wayne. 2001. Examining monophyly in a large radiation of Madagascan butterflies (Lepidoptera: Satyrinae: Mycalesina) based on mitochondrial DNA data. Molecular Phylogenetics and Evolution 20:460-473.
  26. Weingartner, E., N. Wahlberg, and S. Nylin. 2006. Speciation in Pararge (Satyrinae: Nymphalidae) butterflies - North Africa is the source of ancestral populations of all Pararge species. Systematic Entomology 31:621-632.