Ithomiini
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Ithomiini is a
butterfly Butterflies are insects in the macrolepidopteran clade Rhopalocera from the order Lepidoptera, which also includes moths. Adult butterflies have large, often brightly coloured wings, and conspicuous, fluttering flight. The group compris ...
tribe The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide usage of the term in English is in the discipline of anthropology. This definition is contested, in part due to confl ...
in the nymphalid subfamily
Danainae Danainae is a subfamily of the family Nymphalidae, the brush-footed butterflies. It includes the Daniadae, or milkweed butterflies, who lay their eggs on various milkweeds on which their larvae (caterpillars) feed, as well as the clearwing butt ...
. 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 Genus ( plural genera ) is a taxonomic rank used in the biological classifica ...
(Ithomiinae). These butterflies are exclusively
Neotropical The Neotropical realm is one of the eight biogeographic realms constituting Earth's land surface. Physically, it includes the tropical terrestrial ecoregions of the Americas and the entire South American temperate zone. Definition In bioge ...
, 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 The family Asteraceae, alternatively Compositae, consists of over 32,000 known species of flowering plants in over 1,900 genera within the order Asterales. Commonly referred to as the aster, daisy, composite, or sunflower family, Compositae ...
) and wilted borages (
Boraginaceae Boraginaceae, the borage or forget-me-not family, includes about 2,000 species of shrubs, trees and herbs in 146, to 156 genera with a worldwide distribution. The APG IV system from 2016 classifies the Boraginaceae as single family of the ...
). The slow-flying adults are Müllerian mimics of each other as well as of many other Lepidoptera.
Henry Walter Bates Henry Walter Bates (8 February 1825, in Leicester – 16 February 1892, in London) was an English naturalist and explorer who gave the first scientific account of mimicry in animals. He was most famous for his expedition to the rainforests of ...
referred to a "transparency group" of Amazon butterfly species. It was originally with seven species belonging to six different genera.
Reginald Punnett Reginald Crundall Punnett FRS (; 20 June 1875 – 3 January 1967) was a British geneticist who co-founded, with William Bateson, the ''Journal of Genetics'' in 1910. Punnett is probably best remembered today as the creator of the 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 ''Patia orise'' is a butterfly in the family Pieridae. It is found from Costa Rica to Brazil and the Guyanas. Description It is very similar to '' Thyridia psidii'' and '' Methona confusa'', especially in the female. The perfectly transparent ...
'' 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
androconia The external morphology of Lepidoptera is the physiological structure of the bodies of insects belonging to the order Lepidoptera, also known as butterflies and moths. Lepidoptera are distinguished from other orders by the presence ...
l 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 or ...
host plants. Exceptions are the more basal genera ''
Tithorea Tithorea ( el, Τιθορέα), is an ancient place with more than 4,000 years of human history. A part of the municipality of Amfikleia-Elateia, in Phthiotis, Greece, it had a population of 630 in 2011, and is situated 156 km from Athens. ...
'', '' Aeria'', and ''
Elzunia Elzunia is a Neotropical genus of butterflies of the family Nymphalidae, subfamily Danainae, and tribe Ithomiini Ithomiini is a butterfly tribe in the nymphalid subfamily Danainae. It is sometimes referred to as the tribe of clearwing butterf ...
'' that, like ''Tellervo'' and some
Danainae Danainae is a subfamily of the family Nymphalidae, the brush-footed butterflies. It includes the Daniadae, or milkweed butterflies, who lay their eggs on various milkweeds on which their larvae (caterpillars) feed, as well as the clearwing butt ...
, feed on Echiteae vines (
Apocynaceae Apocynaceae (from '' Apocynum'', Greek for "dog-away") is a family of flowering plants that includes trees, shrubs, herbs, stem succulents, and vines, commonly known as the dogbane family, because some taxa were used as dog poison Members ...
,
Apocynoideae Apocynoideae is a subfamily of the flowering plant family Apocynaceae (order Gentianales). It contains about 78 genera with roughly 860 species. Several genera are of pharmacological interest - notably those - such as Strophanthus - which have f ...
), 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 The Nymphalidae are the largest family of butterflies, with more than 6,000 species distri ...
'' and ''
Hyposcada ''Hyposcada'' is a genus of clearwing ( ithomiine) butterflies, named by Frederick DuCane Godman and Osbert Salvin in 1879. They are in the brush-footed butterfly family, Nymphalidae The Nymphalidae are the largest family of butterflies, with ...
'' that feed on
Gesneriaceae Gesneriaceae, the gesneriad family, is a family of flowering plants consisting of about 152 genera and ca. 3,540 species in the tropics and subtropics of the Old World (almost all Didymocarpoideae) and the New World (most Gesnerioideae), ...
. The local abundance of ithomiine butterflies in the Amazon forest, the lack of observations of predation, and their "peculiar smell" led
Henry Walter Bates Henry Walter Bates (8 February 1825, in Leicester – 16 February 1892, in London) was an English naturalist and explorer who gave the first scientific account of mimicry in animals. He was most famous for his expedition to the rainforests of ...
in 1867 to suggest that these organisms should be chemically defended. This was first experimentally demonstrated in 1889 when
Thomas Belt Thomas Belt (183221 September 1878), an English geologist and naturalist, was born at Newcastle-on-Tyne in 1832, and educated in that city. He is remembered for his work on the geology of gold bearing minerals, glacial geology, and for his desc ...
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 Colombian white-faced capuchin (''Cebus capucinus''), also known as the Colombian white-headed capuchin or Colombian white-throated capuchin, is a medium-sized New World monkey of the family Cebidae, subfamily Cebinae. It is native to the ext ...
''. 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 ...
''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 ''Trichonephila clavipes'' (formerly known as ''Nephila clavipes''), commonly known as the golden silk orb-weaver, golden silk spider, or banana spider (a name shared with several others), is an orb-weaving spider species which inhabits forests ...
'' 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 ''Heliotropium'' is a genus of flowering plants in the heliotrope family, Heliotropiaceae. There are around 325 species in this almost cosmopolitan genus, which are commonly known as heliotropes (sg. ). It is highly toxic for dogs and cats. ...
''),
Asteraceae The family Asteraceae, alternatively Compositae, consists of over 32,000 known species of flowering plants in over 1,900 genera within the order Asterales. Commonly referred to as the aster, daisy, composite, or sunflower family, Compositae ...
(mostly in the tribe Eupatorieae, and rarely on ''Senecio'' species),
Apocynaceae Apocynaceae (from '' Apocynum'', Greek for "dog-away") is a family of flowering plants that includes trees, shrubs, herbs, stem succulents, and vines, commonly known as the dogbane family, because some taxa were used as dog poison Members ...
(''Prestonia'', belonging to the tribe Echiteae) and
Orchidaceae Orchids are plants that belong to the family Orchidaceae (), a diverse and widespread group of flowering plants with blooms that are often colourful and fragrant. Along with the Asteraceae, they are one of the two largest families of flowerin ...
(''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 Danainae is a subfamily of the family Nymphalidae, the brush-footed butterflies. It includes the Daniadae, or milkweed butterflies, who lay their eggs on various milkweeds on which their larvae (caterpillars) feed, as well as the clearwing butt ...
, Ctenuchidae, and
Arctiidae The Arctiinae (formerly called the family Arctiidae) are a large and diverse subfamily of moths with around 11,000 species found all over the world, including 6,000 neotropical species.Scoble, MJ. (1995). ''The Lepidoptera: Form, Function and ...
) 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'' that contain pyrrolizidine alkaloids from their larval host plant, ''
Crotalaria ''Crotalaria'' is a genus of flowering plants in the family Fabaceae (subfamily Faboideae) commonly known as rattlepods. The genus includes over 700 species of herbaceous plants and shrubs. Africa is the continent with the majority of ''Crotalari ...
'' (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 goddess of forests. She was the daughter of Tapio, an East Finnish forest spirit. Finnish goddesses Nature goddesses Characters in the Kalevala {{Finland-myth-stub ...
'') or the larger tribe
Danaini The Danaini are a tribe of brush-footed butterflies ( family Nymphalidae). The tribe's type genus ''Danaus'' contains the well-known monarch butterfly (''D. plexippus'') and is also the type genus of the tribe's subfamily, the milkweed butterflie ...
. The relationships of the three tribes in the subfamily
Danainae Danainae is a subfamily of the family Nymphalidae, the brush-footed butterflies. It includes the Daniadae, or milkweed butterflies, who lay their eggs on various milkweeds on which their larvae (caterpillars) feed, as well as the clearwing butt ...
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 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 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 ...
Fox, 1940 ** ''
Elzunia Elzunia is a Neotropical genus of butterflies of the family Nymphalidae, subfamily Danainae, and tribe Ithomiini Ithomiini is a butterfly tribe in the nymphalid subfamily Danainae. It is sometimes referred to as the tribe of clearwing butterf ...
'' Bryk, 1937 ** ''
Tithorea Tithorea ( el, Τιθορέα), is an ancient place with more than 4,000 years of human history. A part of the municipality of Amfikleia-Elateia, in Phthiotis, Greece, it had a population of 630 in 2011, and is situated 156 km from Athens. ...
'' Doubleday, 1847 (= ''Hirsutis'' Haensch, 1909) ** '' Aeria'' Hübner, 1816 * Subtribe Melinaeina Clark, 1947 ** '' Athesis'' Doubleday, 1847 (= ''Roswellia'' Fox, 1948) ** '' Eutresis'' Doubleday, 1847 ** '' Athyrtis'' C. Felder & R. Felder, 1862 ** '' Paititia'' Lamas, 1979 ** '' Olyras'' Doubleday, 1847 ** ''
Patricia Patricia is a female given name of Latin origin. Derived from the Latin word '' patrician'', meaning "noble"; it is the feminine form of the masculine given name Patrick. The name Patricia was the second most common female name in the United Stat ...
'' Fox, 1940 ** ''
Melinaea ''Melinaea'' is a genus of clearwing ( ithomiine) butterflies. They are in the brush-footed butterfly family, Nymphalidae. Species Arranged alphabetically:Mechanitina Bar, 1878 ** '' Methona'' Doubleday, 1847 (= ''Gelotophye'' d'Almeida, 1940) ** '' Thyridia'' 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 o ...
'' Kirby, 1871 (= homonym ''Salacia'' Hübner, 1823; = ''Heteroscada'' Schatz, 1886) ** ''
Sais Sais ( grc, Σάϊς, cop, Ⲥⲁⲓ) was an ancient Egyptian city in the Western Nile Delta on the Canopic branch of the Nile,Mish, Frederick C., Editor in Chief. "Saïs." '' Webster's Ninth New Collegiate Dictionary''. 9th ed. Springfield ...
'' 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 ** '' Placidina'' 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 glasswi ...
'' Hübner, 1816 (= ''Dynothea'' Reakirt, 1866) * Subtribe Oleriina ** ''
Megoleria ''Megoleria'' is a genus of clearwing ( ithomiine) butterflies, named by Constantino in 1999. They are in the brush-footed butterfly family, Nymphalidae The Nymphalidae are the largest family of butterflies, with more than 6,000 species distri ...
'' Constantino, 1999 ** ''
Hyposcada ''Hyposcada'' is a genus of clearwing ( ithomiine) butterflies, named by Frederick DuCane Godman and Osbert Salvin in 1879. They are in the brush-footed butterfly family, Nymphalidae The Nymphalidae are the largest family of butterflies, with ...
'' 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'' 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 & 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 ar ...
** '' 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)


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 ...


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
TOL




In Japanese but
binomial Binomial may refer to: In mathematics *Binomial (polynomial), a polynomial with two terms *Binomial coefficient, numbers appearing in the expansions of powers of binomials *Binomial QMF, a perfect-reconstruction orthogonal wavelet decomposition * ...
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