''Heliconius'' comprises a colorful and widespread genus of brush-footed butterflies commonly known as the longwings or heliconians. This genus is distributed throughout the tropical and subtropical regions of the New World, from South America as far north as the southern United States. The larvae of these butterflies eat

passion flower ''Passiflora'', known also as the passion flowers or passion vines, is a genus of about 550 species of flowering plants, the type genus of the family Passifloraceae. They are mostly tendril-bearing vines, with some being shrubs or trees. They ...

vines (Passifloraceae). Adults exhibit bright wing color patterns which signal their distastefulness to potential predators. Brought to the forefront of scientific attention by Victorian naturalists, these butterflies exhibit a striking diversity and

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 ...

, both amongst themselves and with species in other groups of butterflies and moths. The study of ''Heliconius'' and other groups of mimetic butterflies allowed the English naturalist Henry Walter Bates, following his return from Brazil in 1859, to lend support to Charles Darwin, who had found similar diversity amongst the

Galápagos finches Darwin's finches (also known as the Galápagos finches) are a group of about 18 species of passerine birds. They are well known for their remarkable diversity in beak form and function. They are often classified as the subfamily Geospizinae or t ...

Model for evolutionary study

''Heliconius'' butterflies have been a subject of many studies, due partly to their abundance and the relative ease of breeding them under laboratory conditions, but also because of the extensive mimicry that occurs in this group. From the nineteenth century to the present day, their study has helped scientists to understand how new species are formed and why nature is so diverse. In particular, the genus is suitable for the study of both

Batesian mimicry Batesian mimicry is a form of mimicry where a harmless species has evolved to imitate the warning signals of a harmful species directed at a predator of them both. It is named after the English naturalist Henry Walter Bates, after his work on bu ...

and Müllerian mimicry. Because of the type of plant material that ''Heliconius'' caterpillars favor and the resulting poisons they store in their tissues, the adult butterflies are usually unpalatable to predators. This warning is announced, to the mutual benefit of both parties, by bright colors and contrasting wing patterns, a phenomenon known as

aposematism Aposematism is the advertising by an animal to potential predators that it is not worth attacking or eating. This unprofitability may consist of any defences which make the prey difficult to kill and eat, such as toxicity, venom, foul taste or ...

. ''Heliconius'' butterflies are thus Müllerian mimics of one another, and are also involved in Müllerian mimicry with various species of Ithomiini, Danaini, Riodinidae ('' Ithomeis'' and ''

Stalachtis ''Stalachtis'' is a genus of metalmark butterflies (family Riodinidae). It is currently the only member of the tribe Stalachtini, but many metalmark butterflies are yet to be unequivocally assigned to tribes, so this might change eventually. Th ...

'') and Acraeini as well as pericopine arctiid moths. They are probably the models for various palatable Batesian mimics, including ''

Papilio zagreus ''Papilio zagreus'' is a butterfly of the family Papilionidae (swallowtails). It is found in South America, including Venezuela, Colombia, Ecuador, Peru, Bolivia and western Brazil. Description A powerfully built insect with strong neuration in ...

'' and various Phyciodina.

Convergence

''Heliconius'' butterflies such as ''

Heliconius numata ''Heliconius numata'', the Numata longwing, is a brush-footed butterfly species belonging to the family Nymphalidae, subfamily Heliconiinae. Distribution and habitat This species is native to most of South America, from Venezuela to southern B ...

'' are famous practitioners of Müllerian mimicry, and benefit from mimicking other unpalatable species of butterfly in their local habitat, such as ''

Melinaea ''Melinaea'' is a genus of clearwing ( ithomiine) butterflies. They are in the brush-footed butterfly family, Nymphalidae. Species Arranged alphabetically:convergent evolution, whereby many (sometimes unrelated) species become protected by similar patterns or coloration. This is a distinct strategy from the better-known

. In Batesian mimics defensive coloration or patterns are a bluff, mimicking those of actually poisonous or foul-tasting species. In Müllerian mimicry all species of the set have honest warnings, but the similarity between members of a set allows a single encounter between a predator and one member of the set to deter that predator in all future encounters with all members of the set. In this way multiple, often unrelated species, effectively cooperate with one another to educate their mutual predators. Work has been done to understand the genetic changes responsible for the convergent evolution of wing patterns in comimetic species. Molecular work on two distantly related ''Heliconius'' comimics, ''Heliconius melpomene'' and ''Heliconius erato'', has revealed that homologous genomic regions in the species are responsible for the convergence in wing patterns. Also, Supple had found evidence of two co-mimics ''H. erato'' and ''H. melpomene'' having no shared single-nucleotide polymorphisms (SNPs), which would be indicative of introgression, and hypothesized the same regulatory genes for color/pattern had comparably changed in response to the same selective forces.Supple, M., Hines, H., Dasmahapatra, K., Lewis J., Nielsen D., Lavoie, C., Ray, D., Salavar, C., Mcmillan, O., Counterman, B. 2103. Genomic architecture of adaptive color pattern divergence and convergence in ''Heliconius'' butterflies. Genome research (2013): gr-150615. Similarly, molecular evidence indicates that ''

'' shares the same patterning homologues, but that these loci are locked into a wing patterning supergene that results in a lack of recombination and a finite set of wing pattern morphs. One puzzle with Müllerian mimicry/convergence is that it would be predicted the butterflies to all eventually converge on the same color and pattern for the highest predator education. Instead, ''Heliconius'' butterflies are greatly diverse and even form multiple 'mimicry rings' within the same geographical area. Additional evolutionary forces are likely at work.

Speciation

''Heliconius'' butterflies are models for the study of

speciation Speciation is the evolutionary process by which populations evolve to become distinct species. The biologist Orator F. Cook coined the term in 1906 for cladogenesis, the splitting of lineages, as opposed to anagenesis, phyletic evolution within ...

. Hybrid speciation has been hypothesized to occur in this genus and may contribute to the diverse mimicry found in ''Heliconius'' butterflies. It has been proposed that two closely related species, ''H. cydno'' and ''H. melpomene'', hybridized to create the species ''H. heurippa''. In addition, the

clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...

containing ''Heliconius erato'' radiated before ''Heliconius melpomene'', establishing the wing pattern diversity found in both species of butterfly. In a DNA sequencing comparison involving species ''H. m. aglope'', ''H. timareta'', and ''H. m. amaryllis'', it was found that gene sequences around mimicry loci were more recently diverged in comparison with the rest of the genome, providing evidence for speciation by hybridization over speciation by ancestral polymorphism. Hybridization is correlated with

introgression Introgression, also known as introgressive hybridization, in genetics is the transfer of genetic material from one species into the gene pool of another by the repeated backcrossing of an interspecific hybrid with one of its parent species. Intr ...

. Results from Supple and her team have shown SNP's being polymorphic mostly around hybrid zones of a genome, and they claimed this supported the mechanism of introgression over ancestral variation for genetic material exchange for certain species. Selection factors can drive introgression to revolve around genes correlated with wing pattern and color. Research has shown introgression centering on two known chromosomes that contain mimicry alleles.

Assortive mating Assortative mating (also referred to as positive assortative mating or homogamy (sociology), homogamy) is a mating system, mating pattern and a form of sexual selection in which individuals with similar phenotypes or genotypes mating, mate with on ...

reproductively isolates ''H. heurippa'' from its parental species. Melo did a study on the hybrid ''H. heurippa'' to determine its mating habits regarding preference between other hybrids and its parental species. The results showed ''H. heurippa'' chose to reproduce via

backcrossing Backcrossing is a crossing of a hybrid with one of its parents or an individual genetically similar to its parent, to achieve offspring with a genetic identity closer to that of the parent. It is used in horticulture, animal breeding, and product ...

, while the parental species were highly unlikely to reproduce with the backcrosses. This is significant, because hybrids' mating behavior would relatively quickly isolate itself from its parental species, and eventually form a species itself, as defined by lack of gene flow. His team also hypothesized that along with a mixed inheritance of color and pattern, the hybrids also obtained a mixed preference for mates from their parental species genes. The ''H. heurippa'' likely had a genetic attraction for other hybrids, leading to its reproductive isolation and speciation. Although rare, ''Heliconius'' butterflies are an example of homoploid hybrid speciation, i.e. hybridization without changing the number of chromosomes. Aposematism, using warning colors, has been noted to improve species diversification, which may also contribute to the wide range of ''Heliconius'' butterflies.

Sexual selection of aposematic colors

For aposematism and mimicry to be successful in the butterflies, they must continually evolve their colours to warn predators of their unpalatability. Sexual selection is important in maintaining aposematism, as it helps to select for specific shades of colours rather than general colors. A research team used techniques to determine some the color qualities of a set of butterflies. They found that color was more vivid on the dorsal side of the butterflies than on the ventral. Also, in comparing the sexes, females appeared to have differing brightness in specific spots. It is important to select for specific colors to avoid subtle shades in any of the species involved in the mimicry. Unsuccessful warning colors will reduce the efficiency of the aposematism. To select for specific colours, neural receptors in the butterflies' brains give a disproportionate recognition and selection to those shades. To test the importance of these neural and visual cues in the butterflies, researchers conducted an experiment wherein they eliminated colours from butterflies' wings. When a colour was eliminated, the butterfly was less successful in attracting mates and therefore did not reproduce as much as its counterparts

Sexual Selection of Pheromones

In order to attract mates female ''Heliconius'' secrete pheromones from a yellow like sac that they secrete the scent to appear more attractive to the males. They found that typically it is virgin female ''Heliconius'' that secrete these pheromones, The males are able to attach themselves using their denticles to these secretion sacs during mating in order to ensure secretion. Pheromones are vital when it comes to mate choice it determines the more likely chance that there will be a success in mating between the ''Heliconius.'' There is an reproductive isolation between populations so while mates are attracted by pheromones they still will choose to similar patterned winged ''Heliconius.''

Mating and offspring

''Heliconius'' has evolved two forms of mating. The main form is standard sexual reproduction. Some species of ''Heliconius'', however, have converged evolutionarily in regard to pupal mating. One species to exhibit this behavior is ''

Heliconius charithonia ''Heliconius charithonia'', the zebra longwing or zebra heliconian, is a species of butterfly belonging to the subfamily Heliconiinae of the family Nymphalidae. It was first described by Carl Linnaeus in his 1767 12th edition of ''Systema Natura ...

''. In this form of mating, the male ''Heliconius'' finds a female pupa and waits until a day before she is moulted to mate with her. With this type of mating there is no sexual selection present. ''H. erato'' has a unique mating ritual, in which males transfer anti- aphrodisiac pheromones to females after copulation so that no other males will approach the mated females. No other Lepidoptera exhibit this behavior. ''Heliconius'' female butterflies also disperse their eggs much more slowly than other species of butterflies. They obtain their nutrients for egg production through pollen in the adult stage rather than the larval stage. Due to nutrient collection in the adult rather than larval stage, adult females have a much longer life than other species, which allows them to better disperse their eggs for survival and speciation. This form of egg production is helpful because larvae are much more vulnerable than adult stages, although they also utilize aposematism. Because many of the nutrients needed to produce eggs are obtained in the adult stage, the larval stage is much shorter and less susceptible to predation.

Cyanic characteristics

In order to be unpalatable, the ''Heliconius'' butterflies use cyanic characteristics, meaning they produce substances that have a cyanide group attached to them, ultimately making them harmful. Research has found that the amino acids needed to make the cyanic compounds come from feeding on pollen. Although feeding on pollen takes longer than nectar feeding, the aposematic characteristics help to warn predators away and give them more time for feeding. While ''Heliconius'' larvae feed on Passifloraceae which also have cyanic characteristics, the larvae have evolved the ability to neutralize cyanic molecules to protect them from the negative effects of the plant.

Species

Most current researchers agree that there are some 39 ''Heliconius'' species. These are listed alphabetically here, according to Gerardo Lamas' (2004) checklist. Note that the subspecific nomenclature is incomplete for many species (there are over 2000 published names associated with the genus, many of which are subjective synonyms or infrasubspecific names). * ''Heliconius'' Kluk, 1802 :* '' Heliconius antiochus'' (Linnaeus, 1767) – Antiochus longwing :* '' Heliconius aoede'' (Hübner,

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– Aoede longwing :* '' Heliconius astraea'' Staudinger, 1897 :* ''

Heliconius atthis ''Heliconius atthis'', the false zebra longwing or Atthis longwing, is a species of '' Heliconius'' butterfly. It is endemic to western Ecuador.Heliconius besckei'' Ménétriés, 1857 :* ''

Heliconius burneyi left, ''H. b. catharinae''variation ''serpensis''Cristalino River, Southern Amazon, Brazil ''Heliconius burneyi'', the Burney's longwing, is a butterfly of the family Nymphalidae. It was described by Jacob Hübner in 1831. It is found in the Ama ...

'' (Hübner, 1816) – Burney's longwing :* ''

'' (Linnaeus, 1767) – zebra longwing :* '' Heliconius clysonymus'' Latreille, 1817 – Clysonymus longwing, montane longwing :* ''

Heliconius congener ''Heliconius'' comprises a colorful and widespread genus of brush-footed butterflies commonly known as the longwings or heliconians. This genus is distributed throughout the tropical and subtropical regions of the New World, from South America a ...

'' Weymer, 1890 :* ''

Heliconius cydno ''Heliconius cydno'', the cydno longwing, is a nymphalid butterfly that ranges from Mexico to northern South America. It is typically found in the forest understory and deposits its eggs on a variety of plants of the genus ''Passiflora''. It is ...

'' (Doubleday, 1847) – cydno longwing :* ''

Heliconius demeter ''Heliconius demeter'', the Demeter longwing, is a butterfly of the family Nymphalidae. It was described by Otto Staudinger in 1897. It is found in the Northern and Eastern part of the Amazon basin, from Guyana to Northern Brazil and Peru. The h ...

'' Staudinger, 1897 – Demeter longwing :* '' Heliconius doris'' (Linnaeus, 1771) – Doris longwing or Doris :* ''

Heliconius egeria ''Heliconius'' comprises a colorful and widespread genus of brush-footed butterflies commonly known as the longwings or heliconians. This genus is distributed throughout the tropical and subtropical regions of the New World, from South America ...

'' (Cramer, 1775) :* ''

Heliconius eleuchia ''Heliconius eleuchia'', the white-edged longwing, is a species of '' Heliconius'' butterfly described by William Chapman Hewitson in 1853. Subspecies Subspecies include: * ''Heliconius eleuchia eleuchia'' - present in Costa Rica and in Colom ...

'' Hewitson, 1853 – white-edged longwing or eleuchia longwing :* ''

Heliconius elevatus ''Heliconius elevatus'' is a butterfly of the family Nymphalidae. It was described by Nöldner in 1901. It is found in the Amazon Basin. The habitat consists of riparian forests. The larvae are gregarious and mostly feed on ''Passiflora ...

'' Nöldner, 1901 :* ''

Heliconius erato ''Heliconius erato'', or the red postman, is one of about 40 neotropical species of butterfly belonging to the genus ''Heliconius''. It is also commonly known as the small postman, the red passion flower butterfly, or the crimson-patched longwing ...

'' (Linnaeus, 1764) – crimson-patched longwing, red postman :* ''

Heliconius eratosignis ''Heliconius eratosignis'', the eratosignis longwing, is a butterfly of the family Nymphalidae. It was described by James John Joicey and George Talbot in 1925. It is found in the south-western Amazon basin, from Brazil to Peru and Bolivia. Th ...

'' (Joicey & Talbot, 1925) :* '' Heliconius ethilla'' (Godart, 1819) – Ethilia longwing :* ''

Heliconius godmani ''Heliconius'' comprises a colorful and widespread genus of brush-footed butterflies commonly known as the longwings or heliconians. This genus is distributed throughout the tropical and subtropical regions of the New World, from South America ...

'' Staudinger, 1882 :* ''

Heliconius hecale ''Heliconius hecale'', the tiger longwing, Hecale longwing, golden longwing or golden heliconian, is a heliconiid butterfly that occurs from Mexico to the Peruvian Amazon. Hecale, was an old woman who gave shelter to Theseus on his way to captu ...

'' (Fabricius, 1775) – tiger longwing or Hecale longwing :* '' Heliconius hecalesia'' Hewitson, 1853 – five-spotted longwing :* '' Heliconius hecuba'' (Hewitson,

858 __NOTOC__ Year 858 ( DCCCLVIII) was a common year starting on Saturday (link will display the full calendar) of the Julian calendar. Events By place Europe * Summer – King Louis the German, summoned by the disaffected Frankish ...

– Hecuba longwing :* '' Heliconius hermathena'' (Hewitson, 1853) – Hermathena longwing :* ''

Heliconius heurippa ''Heliconius heurippa'' is a butterfly of the genus '' Heliconius'' that is believed by some scientists to be a separate species from—but a hybrid of—the species ''Heliconius cydno'' and ''Heliconius melpomene'', making ''H. heurippa' ...

'' (Hewitson, 1853) :* ''

Heliconius hewitsoni ''Heliconius'' comprises a colorful and widespread genus of brush-footed butterflies commonly known as the longwings or heliconians. This genus is distributed throughout the tropical and subtropical regions of the New World, from South America a ...

'' Staudinger, 1875 :* '' Heliconius hierax'' Hewitson, 1869 :* ''

Heliconius himera ''Heliconius'' comprises a colorful and widespread genus of Nymphalidae, brush-footed butterflies commonly known as the longwings or heliconians. This genus is distributed throughout the tropical and subtropical regions of the New World, from So ...

'' Hewitson, 1867 :* '' Heliconius hortense'' Guérin,

844 __NOTOC__ Year 844 ( DCCCXLIV) was a leap year starting on Tuesday (link will display the full calendar) of the Julian calendar. Events By place Byzantine Empire * Spring – Battle of Mauropotamos: A Byzantine expedition under ...

/small> – Mexican longwing or mountain longwing :* ''

Heliconius ismenius ''Heliconius ismenius'', the Ismenius tiger or tiger heliconian, is a butterfly of the family Nymphalidae found in Central America and northern South America. They are abundant as far south as Ecuador and Venezuela and as far north as southern Mex ...

'' Latreille,

817 __NOTOC__ Year 817 ( DCCCXVII) was a common year starting on Thursday (link will display the full calendar) of the Julian calendar. Events By place Europe * Summer – Emperor Louis I issues an ''Ordinatio Imperii'', an imperial dec ...

/small> – Ismenius tiger or tiger helconian :* '' Heliconius lalitae'' Brévignon, 1996 :* ''

Heliconius leucadia ''Heliconius'' comprises a colorful and widespread genus of brush-footed butterflies commonly known as the longwings or heliconians. This genus is distributed throughout the tropical and subtropical regions of the New World, from South America a ...

'' (Bates, 1862) – Leucadia longwing :* ''

Heliconius melpomene ''Heliconius melpomene'', the postman butterfly, common postman or simply postman, is a brightly colored butterfly found throughout Central and South America. It was first described by Carl Linnaeus in his 1758 10th edition of ''Systema Naturae' ...

'' (Linnaeus, 1758) – (common) postman :* ''

Heliconius metharme ''Heliconius metharme'' is a species of butterfly of the family Nymphalidae. It was described by Wilhelm Ferdinand Erichson in 1849. It is widespread in the Amazon basin, Venezuela and the Guianas. The habitat consists of deep forests. It is pa ...

'' (Erichson,

849 __NOTOC__ Year 849 (Roman numerals, DCCCXLIX) was a common year starting on Tuesday (link will display the full calendar) of the Julian calendar. Events By place Europe * Summer – Battle of Ostia: A Saracen Arab fleet from Sardi ...

:* ''

Heliconius metis ''Heliconius'' comprises a colorful and widespread genus of brush-footed butterflies commonly known as the longwings or heliconians. This genus is distributed throughout the tropical and subtropical regions of the New World, from South America ...

'' (Moreira & Mielke, 2010) :* ''

Heliconius nattereri ''Heliconius nattereri'' (Natterer's longwing) is a species of butterfly in the family Nymphalidae. It is endemic to the Atlantic forest of Brazil. The butterfly has been found in the Feliciano Miguel Abdala Private Natural Heritage Reserve in ...

'' Felder, 1865 – Natterer's longwing :* ''

'' (Cramer, 1780) – Numata longwing :* ''

Heliconius pachinus ''Heliconius'' comprises a colorful and widespread genus of brush-footed butterflies commonly known as the longwings or heliconians. This genus is distributed throughout the tropical and subtropical regions of the New World, from South America ...

'' Salvin, 1871 – pachinus longwing :* '' Heliconius pardalinus'' (Bates, 1862) :* '' Heliconius peruvianus'' Felder – Peruvian longwing :* ''

Heliconius ricini ''Heliconius ricini'', the ricini longwing, is a butterfly of the family Nymphalidae. It was described by Carl Linnaeus in 1758. It is found from Venezuela and Trinidad to the Guianas and northern Brazil. The habitat consists of savanna-typ ...

'' (Linnaeus, 1758) – ricini longwing :* ''

Heliconius sapho ''Heliconius sapho'', the Sapho longwing, is a butterfly of the family Nymphalidae. It was described by Dru Drury in 1782. It is found from Mexico southward to Ecuador. Name Drury left no notes on the origin of the name, but the spelling (and ...

'' (Drury, 1782) – Sapho longwing :* ''

Heliconius sara The Sara longwing (''Heliconius sara'') is a species of neotropical heliconiid butterfly found from Mexico to the Amazon Basin and southern Brazil. It is a colourful species: the dorsal wing surface is black with a large medial patch of metalli ...

'' (Fabricius, 1793) – Sara longwing :* '' Heliconius sergestus'' (Weymer, 1894) :* ''

Heliconius telesiphe ''Heliconius telesiphe'', the telesiphe longwing, is a butterfly of the family Nymphalidae. It was described by Edward Doubleday in 1847. It is found at mid-elevations in the Andes. Its habitat is cloud forests. The wingspan is 65–80 ...

'' Doubleday, 1847 – telesiphe longwing :* ''

Heliconius timareta ''Heliconius'' comprises a colorful and widespread genus of brush-footed butterflies commonly known as the longwings or heliconians. This genus is distributed throughout the tropical and subtropical regions of the New World, from South America ...

'' (Hewitson, 1867) :* ''

Heliconius tristero ''Heliconius'' comprises a colorful and widespread genus of brush-footed butterflies commonly known as the longwings or heliconians. This genus is distributed throughout the tropical and subtropical regions of the New World, from South America a ...

'' Brower, 1996 :* ''

Heliconius wallacei ''Heliconius wallacei'', the Wallace's longwing, is a butterfly of the family Nymphalidae. It was described by Tryon Reakirt in 1866. It is found from Venezuela and Trinidad to southern Brazil and Peru. The habitat consists of lowland rainforests ...

'' Reakirt, 1866 – Wallace's longwing :* '' Heliconius xanthocles'' Bates, 1862

References

External links

''Heliconius'' Research WorldwideMichel Cast La La diversité des ''Heliconius''

Tree of Life: ''Heliconius''

Neil Rosser et al.: Source geographic distribution data for the species
** {{Taxonbar, from=Q655665 Heliconius, Heliconiini Nymphalidae of South America Nymphalidae genera Taxa named by Jan Krzysztof Kluk