The chemical ecology of these butterflies is fairly well known. Larvae of 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, 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).
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.
Several generic names have been synonymized recently. These include:
- Ceratiscada Brown & d'Almeida 1970 (synonymized with Episcada)
- Dygoris Fox 1945 (synonymized with Godyris)
- Garsauritis d'Almeida 1938 (synonymized with Hypothyris)
- Hypomenitis Fox 1945 (synonymized with Greta)
- Prittwitzia Brown & Ebert 1970 (synonymized with Episcada)
- Rhodussa d'Almeida 1939 (synonymized with Hypothyris)
- Roswellia Fox 1948 (synonymized with Athesis)
- Talamancana Haber, Brown & Freitas 1994 (synonymized with Pteronymia)
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).
Evolution and Systematics
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.
Ithomiini is a butterfly tribe in the nymphalid subfamily Danainae. Some authors consider the group to be a subfamily (Ithomiinae). These butterflies are exclusively Neotropical, found in humid forests from sea-level to 3000 m, from the southwestern United States to Argentina. There are around 370 species in some 40–45 genera.
Ithomiines are unpalatable because their adults seek out and sequester pyrrolizidine alkaloids from plants that they visit, especially composit flowers (Asteraceae) and wilted borages (Boraginaceae). The slow-flying adults are Müllerian mimics of each other as well as of many other Lepidoptera. Henry Walter Bates referred to a "transparency group" of Amazon butterfly species. It was originally with seven species belonging to six different genera. Reginald Punnett suggested 28 species of this peculiar facies are known, though some are excessively rare. The majority are Ithomiines, but two species of the Danaine genus Lycorea, the Pierine Dismorphia orise the Swallow-tail Parides hahneli, and several species of diurnal moths belonging to different families also enter into the combination.Identification of adult ithomiines relies on hindwing venation and male androconial scales (sex brushes located on the hindwing costa).
The group has repeatedly been proposed as biological indicators of ecological conditions or biological diversity within neotropical forests, but individual sites harbor between 10 and 50 species, for the most part, and beta diversity is often great, even over relatively short distances.
Ithomiine larvae feed mostly on Solanaceae host plants. Exceptions are the more basal genera Tithorea, Aeria, and Elzunia that, like Tellervo and some Danainae, feed on Echiteae vines (Apocynaceae, Apocynoideae), as well as Megoleria and Hyposcada that feed on Gesneriaceae.
The local abundance of ithomiine butterflies in the Amazon forest, the lack of observations of predation, and their “peculiar smell” led Henry Walter Bates in 1867 to suggest that these organisms should be chemically defended. This was first experimentally demonstrated in 1889 when Thomas Belt fed ithomiines (that he called “Heliconii”) to birds, the spider Nephila, and the white faced monkey Cebus capucinus. The butterflies were consistently rejected, but other insects were eaten. Lincoln P. Brower in 1964 also showed that adults of Ithomia drymo pellucida were rejected by the bluejay Cyanocitta cristata bromia, and Haber showed that nine species of birds also rejected several ithomiine species. Besides, Vasconcellos-Neto and Lewinsohn demonstrated that the neotropical orb-weaving spider Nephila clavipes released unharmed 14 species of field-caught ithomiine butterflies.
The source of the protecting chemicals in the bodies of adult ithomiines proved not to be their larval host plants, as was first suggested, but rather in plants visited by the butterflies. Adults of ithomiine, mainly males, visit flowers of some Boraginaceae, (Tournefortia, Heliotropium), Asteraceae (mostly in the tribe Eupatorieae, and rarely on Senecio species), Apocynaceae (Prestonia, belonging to the tribe Echiteae) and Orchidaceae (Epidendrum paniculatum). Dead or withered plants are also visited and, when feeding on these plants, the butterflies scratch the tissues with their legs and suck the oozing sap. These plants are known to contain pyrrolizidine alkaloids, indicating their role as chemical sources for sequestration. Other butterfly and moth species that sequester pyrrolizidine alkaloids (Danainae, Ctenuchidae, and Arctiidae) also visit similar sources. The first demonstration that pyrrolizidine alkaloids were involved in the chemical defense of insects was given by Thomas Eisner, who showed that the spiders Nephila and Argiope rejected adults of the arctiid moth Utetheisa ornatrix that contain pyrrolizidine alkaloids from their larval host plant, Crotalaria (Fabaceae: Crotalarieae). Eisner's best-selling popular science book For love of insects tells the story of this exciting discovery.
The subtribes in the Ithomiini help to organize the 43 recognized genera, but this group is the subject of ongoing molecular, phylogenetic, and morphological research, and the classification presented below will no doubt be refined in the near future.
The sister group to the tribe Ithomiini is either the small tribe Tellervini (containing the single Australasian genus Tellervo) or the larger tribe Danaini. The relationships of the three tribes in the subfamily Danainae are still unclear.
- Source: The higher classification of Nymphalidae, at Nymphalidae.net
- Note: A species list with proposed new tribes for subfamily Ithomiinae is available from Keith Willmott at .
- Note: Names preceded by an equal sign ( = ) are synonyms, homonyms, rejected names or invalid names.
Tribe Ithomiini Godman & Salvin, 1879
- (Subtribe Tithoreina Fox, 1940)
- (Subtribe Melinaeina Clark, 1947)
- (Subtribe Mechanitina Bar, 1878)
- Methona Doubleday 1847 ( = Gelotophye d’Almeida 1940)
- Thyridia Hübner 1816 ( = Xanthocleis Boisduval 1870; = Aprotopus Kirby 1871; = Aprotopos Kirby 1871)
- Scada Kirby 1871 ( = homonym Salacia Hübner 1823; = Heteroscada Schatz 1886)
- Sais Hübner 1816
- Forbestra Fox 1967
- Mechanitis Fabricius 1807 ( = homonym Nereis Hübner 1806; = unavailable name Hymenitis Illiger 1807; = Epimetes Billberg 1820)
- (Subtribe Napeogenina)
- Aremfoxia Réal 1971
- Epityches d'Almeida 1938 ( = homonym Tritonia Geyer 1832)
- Hyalyris Boisduval 1870 ( = Oreogenes Stichel 1899)
- Napeogenes Bates 1862 ( = homonym Ceratonia Boisduval 1870; = Choridis Boisduval 1870)
- Hypothyris Hübner 1821 ( = Mansueta d'Almeida 1922; = Pseudomechanitis Röber 1930; = Garsauritis d'Almeida 1938; = Rhodussa d'Almeida 1939)
- (Subtribe Ithomiina Godman & Salvin, 1879)
- (Subtribe Oleriina)
- (Subtribe Dircennina d'Almeida, 1941)
- Ceratinia Hübner 1816 ( = Calloleria Godman & Salvin 1879; = Epileria Rebel 1902; = Teracinia Röber 1910)
- Callithomia Bates 1862 ( = Cleodis Boisduval 1870; = Epithomia Godman & Salvin 1879; = Corbulis Boisduval 1870; = Leithomia Masters 1973)
- Dircenna Doubleday 1847
- Hyalenna Forbes 1942
- Episcada Godman & Salvin 1879 ( = Ceratiscada Brown & d’Almeida 1970; = Prittwitzia Brown, Mielke & Ebert 1970)
- Haenschia Lamas 2004
- Pteronymia Butler & Druce 1872 ( = Ernicornis Capronnier 1874; = Parapteronymia Kremky 1925; = Talamancana Haber, Brown & Freitas 1994)
- (Subtribe Godyridina)
- Velamysta Haensch 1909
- Godyris Boisduval 1870 ( = Dismenitis Haensch 1903; = Dygoris Fox 1945)
- Veladyris Fox 1945
- Hypoleria Godman & Salvin 1879 ( = homonym Pigritia d’Almeida 1922; = homonym Pigritina Hedicke 1923; = homonym Heringia d’Almeida 1924)
- Brevioleria Lamas 2004
- Mcclungia Fox 1940
- Greta Hemming 1934 ( = homonym Hymenitis Hübner 1819; = Hypomenitis Fox 1945)
- Heterosais Godman & Salvin 1880 ( = Rhadinoptera d'Almeida 1922)
- Pseudoscada Godman & Salvin 1879 ( = Languida d’Almeida 1922)
- Lamas, G. (1999). Nymphalidae II Pt. 3 Ithomiinae. In: E. Bauer & T. Frankenbach (Eds.), Butterflies of the World. (pp. 1–17). Keltern, Germany: Goecke & Evers. ISBN 978-3-931374-66-2 16 colour plates - illustrates 252 specimens covering subset of the 320 known species, many of which are divided into subspecies.
- Brown, Jr., K. S., and A. V. L. Freitas. 1994. Juvenile stages of Ithomiinae: overview and systematics (Lepidoptera: Nymphalidae). Tropical Lepidoptera 5(1): 9-20.pdf
- See Links.
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