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Ithomiini is a butterfly tribe in the
nymphalid The Nymphalidae are the largest family of butterflies, with more than 6,000 species distributed throughout most of the world. Belonging to the superfamily Papilionoidea, they are usually medium-sized to large butterflies. Most species have a red ...
subfamily Danainae. It is sometimes referred to as the tribe of clearwing butterflies or glasswing butterflies. Some authors consider the group to be a
subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end subfamily botanical names with "-oideae", and zoologi ...
(Ithomiinae). These butterflies are exclusively Neotropical, found in humid forests from sea level to 3000 m, from Mexico to Argentina. There are around 370 species in some 40–45 genera.


Ithomiini biology

Ithomiines are unpalatable because their adults seek out and sequester
pyrrolizidine alkaloid Pyrrolizidine alkaloids (PAs), sometimes referred to as necine bases, are a group of naturally occurring alkaloids based on the structure of pyrrolizidine. Pyrrolizidine alkaloids are produced by plants as a defense mechanism against insect he ...
s from plants that they visit, especially composite 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 The Solanaceae , or nightshades, are a family of flowering plants that ranges from annual and perennial herbs to vines, lianas, epiphytes, shrubs, and trees, and includes a number of agricultural crops, medicinal plants, spices, weeds, and orn ...
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 ''Megoleria'' is a genus of clearwing ( ithomiine) butterflies, named by Constantino in 1999. They are in the brush-footed butterfly family, Nymphalidae. Species Arranged alphabetically: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 ''Nephila'' is a genus of araneomorph spiders noted for the impressive webs they weave. ''Nephila'' consists of numerous species found in warmer regions around the world. They are commonly called golden silk orb-weavers, golden orb-weavers, gian ...
'', and the white faced monkey '' Cebus capucinus''. The butterflies were consistently rejected, but other insects were eaten.
Lincoln P. Brower Lincoln Pierson Brower (September 10, 1931 – July 17, 2018) was an American entomologist and ecologist, known for his work on monarch butterflies through six decades, including on their automimicry, chemical ecology and conservation. G. Pasteu ...
in 1964 also showed that adults of ''Ithomia drymo pellucida'' were rejected by the
blue jay The blue jay (''Cyanocitta cristata'') is a passerine bird in the family Corvidae, native to eastern North America. It lives in most of the eastern and central United States; some eastern populations may be migratory. Resident populations are a ...
''Cyanocitta cristata bromia'', and Haber showed that nine species of birds also rejected several ithomiine species. Besides, João Vasconcellos-Neto and Thomas M. 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 alkaloid Pyrrolizidine alkaloids (PAs), sometimes referred to as necine bases, are a group of naturally occurring alkaloids based on the structure of pyrrolizidine. Pyrrolizidine alkaloids are produced by plants as a defense mechanism against insect he ...
s, indicating their role as chemical sources for sequestration. Other butterfly and moth species that sequester pyrrolizidine alkaloids ( Danainae,
Ctenuchidae The Ctenuchina are a subtribe of moths in the family Erebidae. Taxonomy The Ctenuchina were previously classified as the subfamily Ctenuchinae of the family Arctiidae. That subfamily contained three tribes: Ctenuchini, Euchromiini (wasp moths) ...
, 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 Thomas Eisner (June 25, 1929 – March 25, 2011) was a German-American entomologist and ecologist, known as the "father of chemical ecology." He was a Jacob Gould Schurman Professor of Chemical Ecology at Cornell University, and Director of the ...
, who showed that the spiders ''
Nephila ''Nephila'' is a genus of araneomorph spiders noted for the impressive webs they weave. ''Nephila'' consists of numerous species found in warmer regions around the world. They are commonly called golden silk orb-weavers, golden orb-weavers, gian ...
'' and '' Argiope'' rejected adults of the arctiid moth ''
Utetheisa ornatrix ''Utetheisa ornatrix'', also called the bella moth, ornate moth or rattlebox moth is a moth of the subfamily Arctiinae. It is aposematically colored ranging from pink, red, orange and yellow to white coloration with black markings arranged in va ...
'' 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.


Ithomiini classification

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 Tellervo () is the Finnish mythology, Finnish goddess of forests. She was the daughter of Tapio (spirit), Tapio, an East Finnish forest spirit. Finnish goddesses Nature goddesses Characters in the Kalevala {{Finland-myth-stub ...
'') or the larger tribe Danaini. The relationships of the three tribes in the subfamily Danainae are still unclear. * Source
"The higher classification of Nymphalidae"
''Nymphalidae.net''. Archived February 20, 2009. * Note: A species list with proposed new tribes for subfamily Ithomiinae is available from
Keith Willmott Keith may refer to: People and fictional characters * Keith (given name), includes a list of people and fictional characters * Keith (surname) * Keith (singer), American singer James Keefer (born 1949) * Baron Keith, a line of Scottish barons i ...
a

* Note: Names preceded by an equal sign (=) are synonyms, homonyms, rejected names or invalid names. Tribe Ithomiini Godman & Salvin, 1879 * Subtribe
Tithoreina Ithomiini is a butterfly Tribe (biology), tribe in the Nymphalidae, nymphalid subfamily Danainae. It is sometimes referred to as the tribe of clearwing butterflies or glasswing butterflies. Some authors consider the group to be a subfamily (Ithom ...
Fox, 1940 ** '' Elzunia'' Bryk, 1937 ** '' Tithorea'' Doubleday, 1847 (= ''Hirsutis'' Haensch, 1909) ** '' Aeria'' Hübner, 1816 * Subtribe
Melinaeina Ithomiini is a butterfly tribe in the nymphalid subfamily Danainae. It is sometimes referred to as the tribe of clearwing butterflies or glasswing butterflies. Some authors consider the group to be a subfamily (Ithomiinae). These butterflies are ...
Clark, 1947 ** '' Athesis'' Doubleday, 1847 (= ''Roswellia'' Fox, 1948) ** ''
Eutresis ''Eutresis'' is a genus of clearwing ( ithomiine) butterflies, named by Edward Doubleday in 1847. They are in the brush-footed butterfly family, Nymphalidae The Nymphalidae are the largest family of butterflies, with more than 6,000 species d ...
'' Doubleday, 1847 ** ''
Athyrtis ''Athyrtis'' is a butterfly genus of the Ithomiini tribe in the brush-footed butterfly family, Nymphalidae. It is a monotypic genus, containing only ''Athyrtis mechanitis''. The genus and the species were named by father and son entomologists Bar ...
'' C. Felder & R. Felder, 1862 ** ''
Paititia ''Paititia'' is a butterfly genus in the tribe Ithomiini in the brush-footed butterfly family, Nymphalidae. The genus is monotypic In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon ...
'' Lamas, 1979 ** ''
Olyras ''Olyras'' is a genus of clearwing (Ithomiini, ithomiine) butterflies, named by Henry Doubleday (entomologist), Henry Doubleday in 1847. They are in the brush-footed butterfly family, Nymphalidae. Species Arranged alphabetically:Patricia Patricia is a female given name of Latin language, Latin origin. Derived from the Latin word ''Patrician (ancient Rome), patrician'', meaning "noble"; it is the feminine form of the masculine given name Patrick (given name), Patrick. The name Patr ...
'' Fox, 1940 ** ''
Melinaea ''Melinaea'' is a genus of clearwing ( ithomiine) butterflies. They are in the brush-footed butterfly family, Nymphalidae. Species Arranged alphabetically:Mechanitina Ithomiini is a butterfly tribe in the nymphalid subfamily Danainae. It is sometimes referred to as the tribe of clearwing butterflies or glasswing butterflies. Some authors consider the group to be a subfamily (Ithomiinae). These butterflies are ...
Bar, 1878 ** '' Methona'' Doubleday, 1847 (= ''Gelotophye'' d'Almeida, 1940) ** ''
Thyridia ''Thyridia'' is a monotypic genus of clearwing ( ithomiine) butterflies, named by Jacob Hübner in 1816. Its only species is ''Thyridia psidii'', the Melantho tigerwing or clapping ticlear. It is in the brush-footed butterfly family, Nymphalidae ...
'' Hübner, 1816 (= ''Xanthocleis'' Boisduval, 1870; = ''Aprotopus'' Kirby, 1871; = ''Aprotopos'' Kirby, 1871) ** ''
Scada Supervisory control and data acquisition (SCADA) is a control system architecture comprising computers, networked data communications and graphical user interfaces for high-level supervision of machines and processes. It also covers sensors and ...
'' Kirby, 1871 (= homonym ''Salacia'' Hübner, 1823; = ''Heteroscada'' Schatz, 1886) ** '' Sais'' Hübner, 1816 ** '' Forbestra'' Fox, 1967 ** ''
Mechanitis ''Mechanitis'' is a genus of butterflies in the tribe Ithomiini, commonly known as tigerwings. They were named by Johan Christian Fabricius in 1807. They are in the brush-footed butterfly family, Nymphalidae. Members of the genus ''Mechanitis'' ...
'' Fabricius, 1807 (= homonym ''Nereis'' Hübner, 1806; = unavailable name ''Hymenitis'' Illiger, 1807; = ''Epimetes'' Billberg, 1820) * Subtribe
Napeogenina Ithomiini is a butterfly Tribe (biology), tribe in the Nymphalidae, nymphalid subfamily Danainae. It is sometimes referred to as the tribe of clearwing butterflies or glasswing butterflies. Some authors consider the group to be a subfamily (Ithom ...
** ''
Aremfoxia ''Aremfoxia'' is a monotypic genus of clearwing ( ithomiine) butterflies in the brush-footed butterfly family, Nymphalidae. The genus was named by Herman G. Real in 1971. Its one species, ''Aremfoxia ferra'', was described by Richard Haensch Ric ...
'' Réal, 1971 ** ''
Epityches ''Epityches'' is a monotypic genus of clearwing ( ithomiine) butterflies, named by d'Almeida in 1938. They are in the brush-footed butterfly family, Nymphalidae The Nymphalidae are the largest family of butterflies, with more than 6,000 speci ...
'' d'Almeida, 1938 (= homonym ''Tritonia'' Geyer, 1832) ** ''
Hyalyris ''Hyalyris'' is a genus of clearwing ( ithomiine) butterflies, named by Jean Baptiste Boisduval in 1870. They are in the brush-footed butterfly family, Nymphalidae. Species Arranged alphabetically:Napeogenes ''Napeogenes'' is a genus of clearwing ( ithomiine) butterflies, named by Henry Walter Bates in 1862. They are in the brush-footed butterfly family, Nymphalidae. Species Arranged alphabetically:Hypothyris'' Hübner, 1821 (= ''Mansueta'' d'Almeida, 1922; = ''Pseudomechanitis'' Röber, 1930; = ''Garsauritis'' d'Almeida, 1938; = ''Rhodussa'' d'Almeida, 1939) * Subtribe Ithomiina Godman & Salvin, 1879 ** ''
Placidina ''Placidina'' is a genus of clearwing ( ithomiine) butterflies, named by d'Almeida in 1928. They are in the brush-footed butterfly family, Nymphalidae. It is a monotypic genus, containing only ''Placidina euryanassa'' described by father-and-son ...
'' d'Almeida, 1928 (= ''Placidula'' d'Almeida, 1922) ** '' Pagyris'' Boisduval, 1870 (= ''Miraleria'' Haensch, 1903) ** ''
Ithomia ''Ithomia'' is a genus of clearwing ( ithomiine) butterflies, named by Jacob Hübner in 1816. They are in the brush-footed butterfly family, Nymphalidae. Species Arranged alphabetically: *''Ithomia agnosia'' (Hewitson, 855 – agnosia glasswin ...
'' Hübner, 1816 (= ''Dynothea'' Reakirt, 1866) * Subtribe
Oleriina Ithomiini is a butterfly tribe in the nymphalid subfamily Danainae. It is sometimes referred to as the tribe of clearwing butterflies or glasswing butterflies. Some authors consider the group to be a subfamily (Ithomiinae). These butterflies are ...
** ''
Megoleria ''Megoleria'' is a genus of clearwing ( ithomiine) butterflies, named by Constantino in 1999. They are in the brush-footed butterfly family, Nymphalidae. Species Arranged alphabetically:Hyposcada'' Godman & Salvin, 1879 ** ''
Oleria ''Oleria'' is a genus of clearwing ( ithomiine) butterflies, named by Jacob Hübner in 1816. They are in the brush-footed butterfly family, Nymphalidae. Species Arranged alphabetically within species groups: *'' Oleria aegle'' (Fabricius, 1776) ...
'' Hübner, 1816 (= ''Leucothyris'' Boisduval, 1870; = ''Ollantaya'' Brown & Freitas, 1994) * Subtribe Dircennina d'Almeida, 1941 ** ''
Ceratinia ''Ceratinia'' is a genus of clearwing ( ithomiine) butterflies, named by Jacob Hübner in 1816. They are in the brush-footed butterfly family, Nymphalidae. Species Arranged alphabetically:Callithomia ''Callithomia'' is a genus of clearwing (Ithomiini, ithomiine) butterflies, named by Henry Walter Bates in 1862. They are in the brush-footed butterfly family, Nymphalidae. Species Arranged alphabetically:Dircenna'' Doubleday, 1847 ** ''
Hyalenna ''Hyalenna'' is a genus of clearwing (Ithomiini, ithomiine) butterflies described by William Trowbridge Merrifield Forbes in 1942. They are in the brush-footed butterfly family, Nymphalidae. Species Arranged alphabetically: *''Hyalenna alidella' ...
'' Forbes, 1942 ** '' Episcada'' Godman & Salvin, 1879 (= ''Ceratiscada'' Brown & d'Almeida, 1970; = ''Prittwitzia'' Brown, Mielke & Ebert, 1970) ** ''
Haenschia ''Haenschia'' is a genus of clearwing ( ithomiine) butterflies, named by Lamas in 2004. They are in the brush-footed butterfly family, Nymphalidae. The name honours German entomologist and insect dealer Richard Haensch Richard Haensch (active fr ...
'' Lamas, 2004 ** ''
Pteronymia ''Pteronymia'' is a genus of clearwing ( ithomiine) butterflies, named by Arthur Gardiner Butler and Herbert Druce in 1872. They are in the brush-footed butterfly family, Nymphalidae. Species Arranged alphabetically: *'' Pteronymia alcmena'' ...
'' Butler & H. Druce, 1872 (= ''Ernicornis'' Capronnier, 1874; = ''Parapteronymia'' Kremky, 1925; = ''Talamancana'' Haber, Brown & Freitas, 1994) * Subtribe
Godyridina Ithomiini is a butterfly tribe in the nymphalid subfamily Danainae. It is sometimes referred to as the tribe of clearwing butterflies or glasswing butterflies. Some authors consider the group to be a subfamily (Ithomiinae). These butterflies are ...
** ''
Velamysta ''Velamysta'' is a genus of clearwing ( ithomiine) butterflies, named by Richard Haensch in 1909. They are in the brush-footed butterfly family, Nymphalidae The Nymphalidae are the largest family of butterflies, with more than 6,000 species d ...
'' Haensch, 1909 ** '' Godyris'' Boisduval, 1870 (= ''Dismenitis'' Haensch, 1903; = ''Dygoris'' Fox, 1945) ** ''
Veladyris ''Veladyris'' is a genus of clearwing ( ithomiine) butterflies, named by Fox in 1945. They are in the brush-footed butterfly family, Nymphalidae. It contains only one species, ''Veladyris pardalis'', which is found in Ecuador and Peru. Subspe ...
'' Fox, 1945 ** ''
Hypoleria ''Hypoleria'' is a genus of clearwing (Ithomiini, ithomiine) butterflies, named by Frederick DuCane Godman and Osbert Salvin in 1879. They are in the brush-footed butterfly family, Nymphalidae. Species Arranged alphabetically:Brevioleria ''Brevioleria'' is a genus of clearwing ( ithomiine) butterflies, named by Lamas in 2004. They are in the brush-footed butterfly family, Nymphalidae. Species Arranged alphabetically:Mcclungia ''Mcclungia'' is a genus of clearwing (Ithomiini, ithomiine) butterflies, named by Fox in 1940. They are in the brush-footed butterfly family, Nymphalidae. The genus is monotypic, containing only ''Mcclungia cymo'' named by Jacob Hübner in 1806. ...
'' Fox, 1940 ** ''
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'' Hemming, 1934 (= homonym ''Hymenitis'' Hübner, 1819; = ''Hypomenitis'' Fox, 1945) ** '' Heterosais'' Godman & Salvin, 1880 (= ''Rhadinoptera'' d'Almeida, 1922) ** ''
Pseudoscada ''Pseudoscada'' is a genus of clearwing ( ithomiine) butterflies, named by Frederick DuCane Godman and Osbert Salvin in 879 They are in the brush-footed butterfly family, Nymphalidae The Nymphalidae are the largest family of butterflies, with ...
'' Godman & Salvin, 1879 (= ''Languida'' d'Almeida, 1922)


See also

*
Mimicry In evolutionary biology, mimicry is an evolved resemblance between an organism and another object, often an organism of another species. Mimicry may evolve between different species, or between individuals of the same species. Often, mimicry f ...
*
Animal coloration Animal coloration is the general appearance of an animal resulting from the reflection or emission of light from its surfaces. Some animals are brightly coloured, while others are hard to see. In some species, such as the peafowl, the male h ...


References

* 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. . 16 color plates - illustrates 252 specimens covering subset of the 320 known species, many of which are divided into subspecies. *Brown Jr., K. S. & Freitas, A. V. L. (1994)
"Juvenile stages of Ithomiinae: overview and systematics (Lepidoptera: Nymphalidae)"
''Tropical Lepidoptera''. 5 (1): 9-20.


External links

*

Florida Museum
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In Japanese but
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names. Three pages. This page is ''Mechanitis''.
Learn about butterflies
Excellent photos of mimicry in ''Methona confusa'' * Richard Haensch, in Seitz, A. (1907). ''The Macrolepidoptera of the World''
5: The Macrolepidoptera of the American faunistic region. Danaidae: 113-171
{{Taxonbar , from=Q291207 - Butterfly tribes