Heliconius Melpomene
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''Heliconius melpomene'', the postman butterfly, common postman or simply postman, is a brightly colored, geographically variable butterfly species found throughout Central and
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...
. It was first described by
Carl Linnaeus Carl Linnaeus (23 May 1707 – 10 January 1778), also known after ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...
in his 1758 10th edition of ''Systema Naturae''. Its coloration coevolved with another member of the genus, '' H. erato'', as a warning to predators of its inedibility; this is an example of
Müllerian mimicry Müllerian mimicry is a natural phenomenon in which two or more well-defended species, often foul-tasting and sharing common predators, have come to mimicry, mimic each other's honest signal, honest aposematism, warning signals, to their mutuali ...
. ''H. melpomene'' was one of the first butterfly species observed to forage for pollen, a behavior that is common in other insect groups but rare in butterflies. Because of the recent rapid
evolutionary radiation An evolutionary radiation is an increase in taxonomic diversity that is caused by elevated rates of speciation, that may or may not be associated with an increase in morphological disparity. A significantly large and diverse radiation within ...
of the genus ''Heliconius'' and overlapping of its habitat with other related species, ''H. melpomene'' has been the subject of extensive study on
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 ...
and hybridization. These hybrids tend to have low fitness as they look different from the original species and no longer exhibit Müllerian mimicry. ''Heliconius melpomene'' possesses
ultraviolet Ultraviolet radiation, also known as simply UV, is electromagnetic radiation of wavelengths of 10–400 nanometers, shorter than that of visible light, but longer than X-rays. UV radiation is present in sunlight and constitutes about 10% of ...
vision which enhances its ability to distinguish subtle differences between markings on the wings of other butterflies. This allows the butterfly to avoid mating with other species that share the same geographic range.


Description

The postman butterfly is predominately black with either red or yellow bands across the
forewings Insect wings are adult outgrowths of the insect exoskeleton that enable insect flight, insects to fly. They are found on the second and third Thorax (insect anatomy), thoracic segments (the mesothorax and metathorax), and the two pairs are often ...
. The postman butterfly has large long wings (35–39 mm). It is poisonous, and the red patterns on its wings are an example of
aposematism Aposematism is the Advertising in biology, advertising by an animal, whether terrestrial or marine, to potential predation, predators that it is not worth attacking or eating. This unprofitability may consist of any defenses which make the pr ...
. They look similar to '' H. erato''. Two features found on the underside of the hind wings help to distinguish ''H. erato'' from ''H. melpomene''—''H. erato'' usually has four red dots where the wing attaches to the thorax while ''H. melpomene'' usually has three. In Mexico, Central America and the west coast of Colombia and Ecuador, the yellowish-white stripe on the underside reaches the margin of the hindwing in ''H. erato'' but ends before reaching the margin in ''H. melpomene''. There are many geographical races/subspecies/ morphs of this butterfly throughout Central and
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...
. The geographical variation in patterns has been studied using linkage mapping and it has been found that the patterns are associated with a small number of genetic loci called genomic "
hotspots Hotspot, Hot Spot or Hot spot may refer to: Places * Hot Spot, Kentucky, a community in the United States Arts, entertainment, and media Fictional entities * Hot Spot (comics), a name for the DC Comics character Isaiah Crockett * Hot Spot (Tr ...
". Hotspot loci for color patterning have been found homologous between co-mimics ''H. erato'' and ''H. melpomene'', strengthening evidence for
parallel evolution Parallel evolution is the similar development of a trait in distinct species that are not closely related, but share a similar original trait in response to similar evolutionary pressure.Zhang, J. and Kumar, S. 1997Detection of convergent and pa ...
between the two species, across morph patterns.


Geographic range and habitat

''Heliconius melpomene'' is found from
Central America Central America is a subregion of North America. Its political boundaries are defined as bordering Mexico to the north, Colombia to the southeast, the Caribbean to the east, and the Pacific Ocean to the southwest. Central America is usually ...
to
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...
, especially on the slopes of the
Andes The Andes ( ), Andes Mountains or Andean Mountain Range (; ) are the List of longest mountain chains on Earth, longest continental mountain range in the world, forming a continuous highland along the western edge of South America. The range ...
mountains. It most commonly inhabits open terrain and forest edges, although it can also be found near the edges of rivers and streams. It shares its range with other ''Heliconius'' species, and ''H. melpomene'' is usually less abundant than other species.


Origins

A recent study, using
amplified fragment length polymorphism Amplified fragment length polymorphism (AFLP-PCR or AFLP) is a PCR-based tool used in genetics research, DNA fingerprinting, and in the practice of genetic engineering. Developed in the early 1990s by Pieter Vos, AFLP uses restriction enzymes t ...
(AFLP) and
mitochondrial DNA Mitochondrial DNA (mtDNA and mDNA) is the DNA located in the mitochondrion, mitochondria organelles in a eukaryotic cell that converts chemical energy from food into adenosine triphosphate (ATP). Mitochondrial DNA is a small portion of the D ...
datasets, places the origin of ''H. melpomene'' to 2.1 million years ago. ''H. melpomene'' shows clustering of AFLPs by geography suggesting that the species originated in eastern South America.


Food resources


Caterpillars

Caterpillars of ''Heliconius'' exhibit a behavior known as
monophagy Feeding is the process by which organisms, typically animals, obtain food. Terminology often uses either the suffixes -vore, -vory, or -vorous from Latin ''vorare'', meaning "to devour", or -phage, -phagy, or -phagous from Greek φαγε ...
, meaning they feed on only one kind of plant, specifically the genus ''
Passiflora ''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. ''Passiflora'' species are widely cultivated for their striking flowers, fla ...
''. In ''H. melpomene'', the host plants are limited to '' Passiflora oerstedii'' and '' Passiflora menispermifolia''. Species of the genus ''Passiflora'' have evolved many chemical defenses, but ''Heliconius'' caterpillars have developed adaptations that allow them to continue to feed on the plants and actually incorporate the toxic compounds into their systems and make them unpalatable to predators. These interactions make ''Heliconius'' and ''Passiflora'' a model system for studies of
coevolution In biology, coevolution occurs when two or more species reciprocally affect each other's evolution through the process of natural selection. The term sometimes is used for two traits in the same species affecting each other's evolution, as well a ...
.


Adults


Diet

Unlike most other butterflies, several ''Heliconius'' species have been observed eating
pollen Pollen is a powdery substance produced by most types of flowers of seed plants for the purpose of sexual reproduction. It consists of pollen grains (highly reduced Gametophyte#Heterospory, microgametophytes), which produce male gametes (sperm ...
as well as
nectar Nectar is a viscous, sugar-rich liquid produced by Plant, plants in glands called nectaries, either within the flowers with which it attracts pollination, pollinating animals, or by extrafloral nectaries, which provide a nutrient source to an ...
. The exact mechanism by which the butterfly digests the pollen is uncertain; it was originally thought that once the pollen was soaked in nectar after ingestion, it would then be able to be digested by the butterfly. Recently, however, the
enzyme An enzyme () is a protein that acts as a biological catalyst by accelerating chemical reactions. The molecules upon which enzymes may act are called substrate (chemistry), substrates, and the enzyme converts the substrates into different mol ...
protease A protease (also called a peptidase, proteinase, or proteolytic enzyme) is an enzyme that catalysis, catalyzes proteolysis, breaking down proteins into smaller polypeptides or single amino acids, and spurring the formation of new protein products ...
was discovered in the butterfly's saliva, which implies an adaptation for breaking down pollen. This enzyme was found in higher concentrations in the saliva of female butterflies, likely due to the greater need of nutrition associated with reproduction. These adaptations allow the butterflies to extract important
amino acid Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although over 500 amino acids exist in nature, by far the most important are the 22 α-amino acids incorporated into proteins. Only these 22 a ...
s from the pollen, which, in addition to general nutrition benefits, allows ''H. melpomene'' to have brighter colors and be more distasteful to predators than their non-pollen-foraging counterparts. It is thought that this foraging adaptation and subsequent enhancement of coloration contributed to the speciation of ''Heliconius''.


Pollination

Pollen is a rarely utilized but efficient protein source for ''Lepidoptera'' species. While
foraging Foraging is searching for wild food resources. It affects an animal's fitness because it plays an important role in an animal's ability to survive and reproduce. Foraging theory is a branch of behavioral ecology that studies the foraging behavi ...
for pollen, adults accumulate pollen on the end of their
proboscis A proboscis () is an elongated appendage from the head of an animal, either a vertebrate or an invertebrate. In invertebrates, the term usually refers to tubular arthropod mouthparts, mouthparts used for feeding and sucking. In vertebrates, a pr ...
and the grains stay there for long periods of time. These pollen grains are transferred to the stamen of another plant the butterfly visits while foraging. While there are many plants in ''H. melpomene'' range that provide suitable nutrients, only a few of these are visited by the butterfly. This makes the butterfly an efficient pollinator for the flowers it visits as there is a low likelihood of a plant receiving the wrong kind of pollen.


Parental care

By foraging for pollen while developing eggs, female ''H. melpomene'' butterflies provide valuable amino acids and proteins to their offspring. This reduces the amount of time that the offspring must spend foraging during the larval stage, and thus decreases the chances of larval predation. While this extra foraging behavior on the part of the female increases her likelihood of being eaten, the warning colors highlighting her distaste protect her from would be predators.


Oviposition

Female ''H. melpomene'' butterflies recognize host plants by identifying the corresponding chemical compound using
chemoreceptor A chemoreceptor, also known as chemosensor, is a specialized sensory receptor which transduces a chemical substance ( endogenous or induced) to generate a biological signal. This signal may be in the form of an action potential, if the chemorece ...
s located on her forelegs. When searching for a plant, the butterfly will drum her legs on the plant in order to detect the chemical compounds the plant releases. Once she has found the correct host plant, she will lay eggs singly on separate young leaves. Finding the correct host plant is crucial as ''H. melpomene'' larvae are adapted to only feed on certain ''Passiflora'' plants.


Life cycle

The eggs of ''H. melpomene'' are yellow and approximately 1.5 x 1 mm. They are mostly laid singly or rarely in small clusters on the young leaves of ''Passiflora'' plants. Caterpillars live in groups of two to three individuals and are white with black spots. Pupae are spiny and dark brown in color. The adults have black bodies with bright yellow or orange patterns on the wings. Female ''H. melpomene'' produce oocytes continuously throughout their life; this is due to the high nutrient diet the butterfly obtains from eating pollen. Closely related ''Heliconius'' species have been reported to have a maximum life span of six months, and it is likely that ''H. melpomene'' lives for a similar length of time.


Protective coloration and behavior


Mimicry

''Heliconius melpomene'' coevolved with its sister species, '' H. erato'', each developing similar bright color patterns. The ''H. melpomene'' patterns correspond to at least 20 of the 27 ''H. erato'' subspecies. This coloration warns potential predators that the butterflies are distasteful and should be avoided; this is an example of
aposematism Aposematism is the Advertising in biology, advertising by an animal, whether terrestrial or marine, to potential predation, predators that it is not worth attacking or eating. This unprofitability may consist of any defenses which make the pr ...
. Since both species possess this acrid taste, they display what is known as
Müllerian mimicry Müllerian mimicry is a natural phenomenon in which two or more well-defended species, often foul-tasting and sharing common predators, have come to mimicry, mimic each other's honest signal, honest aposematism, warning signals, to their mutuali ...
. Despite their easily confused coloration, these two species are able to exist in the same habitat range because they are
reproductively isolated The mechanisms of reproductive isolation are a collection of evolutionary mechanisms, behaviors and physiological processes critical for speciation. They prevent members of different species from producing offspring, or ensure that any offsprin ...
due to adaptations in the eyes of the butterflies that allow for better distinction between individuals.


Chemical defense

Both males and females release a strong odor detectable even to humans when handled in order to deter predation. Additionally, ''H. melpomene'' butterflies render themselves unpalatable to predators such as birds by producing
cyanogenic glycosides In chemistry, a glycoside is a molecule in which a sugar is bound to another functional group via a glycosidic bond. Glycosides play numerous important roles in living organisms. Many plants store chemicals in the form of inactive glycosides. ...
in both the larval and adult stages. These glycosides are incorporated into the insect's system by feeding on host plants that produce the compounds as a defense against herbivory.


Communal roosting

Individuals of the genus ''Heliconius'' form large communal roosts which they return to each night after foraging. The reason for this behavior was not well characterized until recently when it was determined that the large aggregations of butterflies provided protection from predators. Butterflies fare better in these groups for two reasons. First, the prey dilution effect lowers the likelihood that one particular individual will be eaten because of the large number of other individuals that are in the area. Second, the congregation of the brightly colored individuals is more likely to deter predators by making the warning coloration more prominent.


Genetics


Hybridization

Due to its overlapping range with many closely related species, ''H. melpomene'' sometimes hybridizes in nature despite adaptations meant to counteract this. Females resulting from the cross of ''H. melpomene'' and '' H. cydno'' are sterile. While hybrid males are not sterile, they exhibit patterns that are intermediate between the crossed species and thus the males are unlikely to be recognized as mates by either species. Furthermore, the patterns on both sexes will be non-mimetic, meaning they will not be recognized by predators as displays of distaste. Therefore, the hybrids resulting from the cross of ''H. melpomene'' with other ''Heliconius'' species have low fitness and are not likely to persist.


Hybrid speciation

Hybridisation of ''Heliconius melpomene'' with '' Heliconius pardalinus'' resulted in the hybrid speciation of '' Heliconius elevatus''.A new butterfly species was created by two species hybridising 200,000 years ago. (2024, April 17). Leibniz Institute for the Analysis of Biodiversity Change (LIB). Retrieved April 21, 2024, from https://leibniz-lib.de/en/2024-04-17-new-butterfly-species/Rosser, N., Seixas, F., Queste, L.M. et al. Hybrid speciation driven by multilocus introgression of ecological traits. Nature (2024). https://doi.org/10.1038/s41586-024-07263-w


Subspecies

Subspecies of ''H. melpomene'' include:Wahlberg N. (last change 1 August 2022)
Heliconiini
Nymphalidae.net. Retrieved 5 February 2010.
* ''H. m. aglaope'' (C. & R. Felder, 1862) (Amazonian foothills of Ecuador and Peru) * ''H. m. amandus'' (Grose-Smith & Kirby, 1892) (Bolivia) * ''H. m. amaryllis'' (C. & R. Felder, 1862) (Tarapoto Valley, Peru) * ''H. m. cythera'' (Hewitson, 1869) (western slopes od Ecuador) * ''H. m. euryades'' (Riffarth, 1900) (southeastern Peru) * ''H. m. malleti'' (Lamas, 1988) (western Amazonia) * ''H. m. melpomene'' (Linnaeus, 1758) (northern coast of South America from Panama to the Amazon) * ''H. m. meriana'' (Turner, 1967) (Guianas) * ''H. m. nanna'' (Stichel, 1899) (Atlantic coastal forest of Brazil) * ''H. m. penelope'' (Staudinger, 1894) (Bolivia) * ''H. m. plesseni'' (Riffarth, 1907)(Andean foothills of eastern Ecuador) * ''H. m. rosina'' (Boisduval, 1870) (Central America) * ''H. m. thelxiope'' (Hübner,
806 __NOTOC__ Year 806 ( DCCCVI) was a common year starting on Thursday of the Julian calendar, the 806th year of the Common Era (CE) and Anno Domini (AD) designations, the 806th year of the 1st millennium, the 6th year of the 9th century, and the ...
(eastern Brazil, south of the Amazon) * ''H. m. vicinus'' (Ménétriés, 1847) (Rio Negro, Brazil and southern Venezuela) * ''H. m. vulcanus'' (Butler, 1865) (Choco (western slope of Colombia)) * ''H. m. xenoclea'' (Hewitson,
853 __NOTOC__ Year 853 ( DCCCLIII) was a common year starting on Sunday of the Julian calendar. Events By place Byzantine Empire * May 22 – A Byzantine fleet (85 ships and 5,000 men) sacks and destroys the port city of Damietta, lo ...
/small> (Rio Perene. eastern Peru)


Mating


Mate searching

When searching for mates, males of ''H. melpomene'' exhibit patrolling behavior, which involves searching for potential mates while flying around the range that the species inhabits. This requires the ability to distinguish ''H. melpomene'' females from those of other species, a key adaptation of the butterfly.


Female/male interactions

Male ''H. melpomene'' possess abdominal
clasper In biology, a clasper is a male anatomical structure found in some groups of animals, used in mating. Male cartilaginous fish have claspers formed from the posterior portion of their pelvic fin which serve to channel semen into the female's ...
s that are used to grasp females for forced copulations. During mating, the male passes nutrients in a
spermatophore A spermatophore, from Ancient Greek σπέρμα (''spérma''), meaning "seed", and -φόρος (''-phóros''), meaning "bearing", or sperm ampulla is a capsule or mass containing spermatozoa created by males of various animal species, especiall ...
; the female can use this
nuptial gift Formally, a nuptial gift is a material presentation to a recipient by a donor during or in relation to sexual intercourse that is not simply gametes in order to improve the reproductive fitness of the donor. Often, such a gift will improve the fit ...
to nourish the fertilizing eggs inside her. In addition to the spermatophore, males also deliver a
pheromone A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
to the female that is an antiaphrodisiac to other males. This increases the likelihood of the male's reproductive success by preventing the female from mating with any other males, which ensures that only the original male's sperm will be used to fertilize the female's eggs.. The pheromone is produced only by males and is secreted to identify themselves to other males, so the antiaphrodisiac works by making the female smell like a male. After a period of time the pheromone wears off and the female is able to mate again, which she will do several times throughout her life.


Physiology


Vision

Due to a duplication in a gene for UV light detection, ''H. melpomene'' individuals are capable of distinguishing between a wider range of yellow shades than other butterfly species. Additionally, when looking for mates, the butterflies distinguish conspecifics from hybrids and heterospecifics by detecting subtle changes in marking patterns on wings. These adaptations allow the butterflies to avoid genetically costly mates, as hybrid females are sterile and hybrid males in this system are less fit due to disruptive sexual selection.


Gustation/tasting

While both sexes of ''H. melpomene'' possess
taste receptor A taste receptor or tastant is a type of cellular receptor that facilitates the sensation of taste. When food or other substances enter the mouth, molecules interact with saliva and are bound to taste receptors in the oral cavity and other locat ...
s on their hindlegs, only the female butterflies have the receptors on the forelegs; this is an example of
sexual dimorphism Sexual dimorphism is the condition where sexes of the same species exhibit different Morphology (biology), morphological characteristics, including characteristics not directly involved in reproduction. The condition occurs in most dioecy, di ...
. The taste receptors are used by both sexes in order to find food and mates, but the female also uses the sense to find suitable host plants for her eggs. These taste receptors are highly specialized due to the coevolution with the ''Passiflora'' plant.


Gallery

File:Heliconius melpomene penelope MHNT dos Male.jpg, ''H. m. penelope'', male, dorsal File:Heliconius melpomene penelope MHNT ventre Male.jpg, ''H. m. penelope'', male, ventral File:Heliconius melpomene penelope MHNT dos Femelle.jpg, ''H. m. penelope'' female, dorsal File:Heliconius melpomene penelope MHNT ventre Femelle.jpg, ''H. m. penelope'' female, ventral


References


Further reading

* *


External links


''Heliconius melpomene''
at the
Tree of Life Web Project The Tree of Life Web Project (ToL) is an Internet project providing information about the diversity and phylogeny of life on Earth. This collaborative peer reviewed project began in 1995, and is written by biologists from around the world. The s ...
*
"''Heliconius'' Kluk, 1780"
at Markku Savela's ''Lepidoptera and Some Other Life Forms'' {{Taxonbar, from=Q2697054
melpomene Melpomene (; ) is the Muse of tragedy in Greek mythology. She is described as the daughter of Zeus and Mnemosyne (and therefore of power and memory) along with the other Muses, and she is often portrayed with a tragic theatrical mask. Etymolog ...
Butterflies of Central America Nymphalidae of South America Butterflies of North America Lepidoptera of Brazil Lepidoptera of Colombia Lepidoptera of Venezuela Fauna of the Amazon Butterflies described in 1758 Taxa named by Carl Linnaeus