Elymniini

Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Protostomia
Cladus: Ecdysozoa
Cladus: Panarthropoda
Phylum: Arthropoda
Subphylum: Hexapoda
Classis: Insecta
Cladus: Dicondylia
Subclassis: Pterygota
Cladus: Metapterygota
Infraclassis: Neoptera
Cladus: Eumetabola
Cladus: Endopterygota
Superordo: Panorpida
Cladus: Amphiesmenoptera
Ordo: Lepidoptera
Subordo: Glossata
Cladus: Coelolepida
Cladus: Myoglossata
Cladus: Neolepidoptera
Infraordo: Heteroneura
Cladus: Eulepidoptera
Cladus: Ditrysia
Cladus: Apoditrysia
Cladus: Obtectomera
Superfamilia: Papilionoidea

Familia: Nymphalidae
Subfamilia: Satyrinae
Tribus: Elymniini
Subtribus: Elymniina
Genera not assignet to a subtribus

Celebina – Debis – Zophoessa –

Name

Elymniini Herrich-Schäffer, 1864
Vernacular names
English: True browns

The Elymniini is one of the tribes of the subfamily Satyrinae. If the subfamily Satyrinae elevate to family status, this tribe shall be treated as subfamily Elymniinae. Elymniini was formerly a large group, but recently, it is considered to be include only one genus, Elymnias, according to molecular phylogenetic analyses.

Systematics and taxonomy

The systematics and taxonomy of Satyrinae began to be heavily revised around the 2000s, and this revision continues to this day. Consequently, the subdivisions of this tribe have also undergone significant changes in recent years.

The higher level classification of Satyrine butterfly developed by Miller (1968) became the basis for later taxonomic studies.[2][3] The work which based on adult morphology, recognised seven subfamilies in the family Satyridae: Haeterinae, Brassolinae, Biinae, Elymniinae, Eritinae, Ragadiinae and Satyrinae, and four tribes within Elymniinae: Lethini, Zetherini, Elymniini and Mycalesini.[4] A later work, Harvey (1991), largely followed Miller's classification, but downranked the family Satyridae to the subfamily Satyrinae, with the consequent downranking of the subdivisions : subfamilies to tribes, tribes to subtribes, such as Elymniinae to the tribe Elymniini and Elymniini to the subtribe Elymniiti.[3] In these classifications, Elymniinae/Elymniini was the second largest subgroup of Satyrine butterflies after Satyrinae/Satyrini. However, the morphological key characters that defined this group were not entirely definitive and the phylogenetic relationship of this group with other groups in the family/subfamily was confused.[3][5]

The confusion over the taxonomy and phylogeny of Satyrinae is gradually being resolved by recent developments in molecular phylogenetic studies.[2] Peña et al. (2006) was the first to develop a comprehensive phylogenetic hypothesis of Satyrinae since Miller, and showed that most of the families previously recognised were paraphyletic or polyphyletic. In the classification proposed in the work, Elymniini included only genus Elymnias, and the genera formerly included in that tribe, such as Lethe, Pararge and Mycalesis, were transferred to several tribes and subtribes, such as Lethina, Parargina and Mycalesina in Satyrini.[3] Subsequent analyses have largely followed this same classification scheme,[6][7][8] and also, all of the genera included in Elymniini by Miller were synonymized with Elymnias.[1][9] Thus, the tribe Elymniini should include only a single genus Elymnias, according to current phylogenetic understanding.[6]

However, this classification is still new and may not always be used in catalogs, picture books, etc.[5] Moreover, phylogenetic analyses have not yet resolved the problems. For example, some analyses relate Hyantis and Morphopsis to this tribe, but others do not, therefore, it is not clear to which tribe these two genera should be placed in.[6] In addition, several different hypotheses have been proposed for the phylogenetic relationship between the tribe Elymniini and other tribes within Satyrinae,[3][7][10] thus there is no consensus on the phylogenetic position of this tribe in the subfamily.[6]
Zethera incerta, Indonesia
Pararge aegeria male, Portugal
Lethe anthedon, Virginia, USA
Mycalesis perseus, Kerala, India
Table for representative higher level classifications of satyrines in Peña et al. (2006)
(excerpts with some modifications[a])
Notes

Only tribes related to this tribe are extracted and the format of the table are changed. In addition, Lamas (2004) was excluded because it only listed genera distributed in the Neotropics. Moreover, the genera Aeropetes and Paralethe which formerly related to this tribe, are not included here, because they have been re-transferred to another tribe (Dirini) since this study.[11]
Elymniopsis has been synonymized with Elymnias.[9][11]

Orsotriaena have been re-transferred to another subtribe (Eritina) in Satyrini since this study.[12]

References
Citations

Wahlberg 2018.
Peña & Wahlberg (NYMPHALIDAE.net).
Peña et al. 2006.
Miller 1968.
YIN et al. 2007.
Marín et al. 2011.
Yang & Zhang 2015.
Chen et al. 2020.
Wei et al. 2017.
Peña & Wahlberg 2008.
Wahlberg 2019.

Kodandaramaiah et al. 2010.

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.

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.

Kodandaramaiah, Ullasa; Peña, Carlos; Braby, Michael F.; Grund, Roger; Müller, Chris J.; Nylin, Sören; Wahlberg, Niklas (2010). "Phylogenetics of Coenonymphina (Nymphalidae: Satyrinae) and the problem of rooting rapid radiations" (PDF). Molecular Phylogenetics and Evolution. 54 (2): 386–394. doi:10.1016/j.ympev.2009.08.012. PMID 19686856.

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. 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.

Wahlberg, Niklas. "The Nymphalidae Systematics Group".
Peña, Carlos; Wahlberg, Niklas. "Satyrinae". NYMPHALIDAE.net.
Wahlberg, Niklas (2018). "Elymniini". NYMPHALIDAE.net.
Wahlberg, Niklas (2019). "The higher classification of Nymphalidae". NYMPHALIDAE.net.

Wei, Chia-Hsuan; Lohman, David J.; Peggie, Djunijanti; Yen, Shen-Horn (2017). "An illustrated checklist of the genus Elymnias Hübner, 1818 (Nymphalidae, Satyrinae)". ZooKeys (676): 47–152. doi:10.3897/zookeys.676.12579. PMC 5523206. PMID 28769686.

Yang, Mingsheng; Zhang, Yalin (2015). "Phylogenetic utility of ribosomal genes for reconstructing the phylogeny of five Chinese satyrine tribes (Lepidoptera, Nymphalidae)". ZooKeys (488): 105–120. doi:10.3897/zookeys.488.9171. PMC 4389127. PMID 25878526.

YIN, Xian-bing; HAO, Jia-sheng; XU, Li; ZHU, Guo-ping; HUANG, Dun-yuan; PAN, Hong-chun (2007). "基于线粒体 ND1 和COI 基因序列探讨锯眼蝶 亚科主要类群的系统发生关系" [Phylogentic Relationships of Butterflies in the Subfamily Elymninae (Lepidoptera: Satyridae) Based on Mitochondrial ND1 and COI Gene Sequences]. 动物学研究 Zoological Research (in Chinese). 28 (5): 477–484. ISSN 0254-5853.