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Ithomiini Doherty 1891

Clearwing Butterflies

Andrew V. Z. Brower and Chris Jiggins
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taxon links [up-->]Aeria [up-->]Velamysta [up-->]Olyras [up-->]Mcclungia [up-->]Athesis [up-->]Greta [up-->]Hypothyris [up-->]Haenschia [up-->]Ithomia [up-->]Ollantaya [up-->]Athyrtis mechanitis [up-->]Godyris [up-->]Forbestra [up-->]Paititia neglecta [up-->]Pachacutia [up-->]Veladyris [up-->]Melinaea [up-->]Episcada [up-->]Mechanitis [up-->]Scada [up-->]Eutresis [up-->]Hyalyris [up-->]Callithomia [up-->]Megoleria [up-->]Patricia [up-->]Hypoleria [up-->]Methona [up-->]Pagyris [up-->]Ceratinia [up-->]Elzunia [up-->]Hyposcada [up-->]Aremfoxia [up-->]Placidina [up-->]Pseudoscada [up-->]Heterosais [up-->]Tithorea [up-->]Brevioleria [up-->]Sais [up-->]Thyridia [up-->]Napeogenes [up-->]Pteronymia [up-->]Oleria [up-->]Dircenna [up-->]Hyalenna [up-->]Epityches Monophyly UncertainMonophyly UncertainMonophyly UncertainMonophyly UncertainMonophyly UncertainPhylogenetic position of group is uncertainPhylogenetic position of group is uncertainPhylogenetic position of group is uncertain[down<--]Danainae Interpreting the tree
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Containing group: Danainae


The ithomiine butterflies are a diverse neotropical group of approximately 370 species that show huge diversity of colour pattern and form (Lamas 2004). Most species are involved in Müllerian mimicry with one another and serve as models in mimicry complexes with other Lepidoptera, including the heliconiines and the danaine genus Lycorea.


The chemical ecology of these butterflies is fairly well known. Larvae of relatively basal Tithoreina feed on host plants in the family Apocynaceae, a host plant group shared with the related danaine butterflies. Most species, however feed on the Solanaceae (nightshades), which is a very speciose and chemically diverse plant family. Most ithomiines are thought to be distasteful to predators, but interestingly some species do not sequester chemicals from their larval host plants (Brown 1984a & b). Instead the males collect pyrrolizidine alkaloids from flowers (especially Asteraceae) and rotting leaves (mainly of Boraginaceae). This may be an adaptation to reduce dependence on larval host plant chemicals (Brown, 1987).

Discussion of Phylogenetic Relationships

The most recent revision of ithomiine sub-tribal and generic nomenclature is in the Checklist of Neotropical Lepidoptera (Lamas 2004), upon which the names in this tree and its subtrees are based. Lamas (2004), as well as Willmott and Freitas (2006) view Ithomiinae as a subfamily. Here, the clade is treated as a tribe, in order to accommodate the well-supported monophyly of (Danaini , Tellervini , Ithomiini) as a subfamily within Nymphalidae. Thus, Lamas' tribal names are listed here as subtribes.

The topology is derived from several recent works: the DNA sequence-based phylogenetic hypothesis of Brower et al. (2006), and the extensive morphological work of Willmott and Freitas (2006). It is based in part upon an unpublished combination of data from both. Relationships within Oleriina are based on de Silva et al. (2010). The phylogenetic affinities of three taxa (Eutresis, Haenschia and Aremfoxia) whose phylogenetic positions were not well resolved in Willmott and Freitas (2006) and were not sampled by Brower et al. (2006) are indicated as uncertain.

A few genera are likely paraphyletic with respect to others as currently circumscribed: molecular and morphological evidence both suggest that Pagyris is paraphyletic with respect to Placidina, Hypothyris is paraphyletic with respect to Hyalyris, Episcada is paraphyletic with respect to Ceratinia, Greta is paraphyletic with respect to Pseudoscada. The morphological data further suggests paraphyly of Godyris and Hypoleria.


Several generic names have been synonymized recently. These include:

Other Names for Ithomiini Doherty 1891


Brower A. V. Z., A. V. L. Freitas, M.-M. Lee, K. L. Silva-Brandao, A. Whinnett, and K. R. Willmott. 2006. Phylogenetic relationships among the Ithomiini (Lepidoptera: Nymphalidae) inferred from one mitchondrial and two nuclear gene regions. Syst. Ent. 31: 288-301.

Brown, K. S. 1984. Adult-obtained pyrrolizidine alkaloids defend ithomiine butterflies against a spider predator. Nature (London) 309:707-709.

Brown, K. S. 1984. Chemical ecology of dehydropyrrolizidine alkaloids in adult Ithomiinae (Lepidoptera: Nymphalidae). Rev. Brasil. Biol. 44:435-460.

Brown, K. S. 1987. Chemistry at the Solanaceae/Ithomiinae interface. Annals of the Missouri Botanical Garden 74:341-358.

Brown, K. S., and A. V. L. Freitas. 1994. Juvenile stages of Ithomiinae: overview and systematics (Lepidoptera: Nymphalidae). Tropical Lepidoptera 5:9-20.

De Silva, D. L., Day, J. J., Elias, M., Willmott, K., Whinnett, A. & Mallet, J. 2010. Molecular phylogenetics of the neotropical butterfly subtribe Oleriina (Nymphalidae: Danaini: Ithomiini). Mol. Phhylogenet. Evol. 55, 1032-1041.

Fox, R. M. 1940. A generic review of the Ithomiinae (Lepidoptera: Nymphalidae). Transactions of the American Entomological Society 66:161-207.

Fox, R. M. 1968. Ithomiidae (Lepidoptera: Nymphaloidea) of Central America. Transactions of the American Entomological Society 94:155-208.

Lamas, G. 2004. Ithomiinae in J. B. Heppner, ed. Atlas of Neotropical Lepidoptera. Checklist: Part 4A. Hesperioidea - Papilionoidea. Association for Tropical Lepidoptera/Scientific Publishers, Gainsville.

Mallarino, R., E. Bermingham, K. R. Willmott, A. Whinnett, and C. D. Jiggins. 2004. Molecular systematics of the butterfly genus Ithomia (Lepidoptera: Ithomiinae): a composite phylogenetic hypothesis based on seven genes. Molecular Phylogenetics and Evolution 34:625-644.

Willmott, K. R. and A. L. V. Freitas. 2006. Higher-level phylogeny of the Ithomiinae (Lepidoptera: Nymphalidae) : classification, patterns of larval hostplant colonization and diversification. Cladistics 22: 297-368.

Information on the Internet

The butterflies of Monteverde, Costa Rica
Illustrated checklist of Ithomiinae
Title Illustrations
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Scientific Name Aeria eurimedia
Location Amazonia Lodge, Atalaya, Peru
Comments Hairpencils are androconial organs on the costal margin of the hindwing in male ithomiines. They represent a synapomorphy or the tribe.
Specimen Condition Live Specimen
Identified By Kim Garwood
Behavior fanning hairpencils in courtship display
Sex Male
Life Cycle Stage adult
View lateral
Source Yellow-striped clearwing
Source Collection Neotropical Butterflies
Copyright © 2004 Kim Garwood
Scientific Name Greta nero
Location Fortuna, Chiriqui, Panama
Specimen Condition Live Specimen
Identified By C Jiggins
Sex Male
Image Use creative commons This media file is licensed under the Creative Commons Attribution-NonCommercial-ShareAlike License - Version 3.0.
Copyright © 2005 Chris Jiggins
About This Page

Middle Tennessee State University, Murfreesboro, Tennessee, USA

Chris Jiggins
University of Cambridge, Cambridge, UK

Correspondence regarding this page should be directed to Andrew V. Z. Brower at and Chris Jiggins at

Page: Tree of Life Ithomiini Doherty 1891. Clearwing Butterflies. Authored by Andrew V. Z. Brower and Chris Jiggins. The TEXT of this page is licensed under the Creative Commons Attribution License - Version 3.0. Note that images and other media featured on this page are each governed by their own license, and they may or may not be available for reuse. Click on an image or a media link to access the media data window, which provides the relevant licensing information. For the general terms and conditions of ToL material reuse and redistribution, please see the Tree of Life Copyright Policies.

Citing this page:

Brower, Andrew V. Z. and Chris Jiggins. 2010. Ithomiini Doherty 1891. Clearwing Butterflies. Version 21 July 2010 (under construction). http://tolweb.org/Ithomiini/27570/2010.07.21 in The Tree of Life Web Project, http://tolweb.org/

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