Red postman | |
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H. e. petiverana, dorsal view | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Class: | Insecta |
Order: | Lepidoptera |
Family: | Nymphalidae |
Genus: | Heliconius |
Species: | H. erato |
Binomial name | |
Heliconius erato | |
Subspecies | |
Many, see text | |
Synonyms | |
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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. It was described by Carl Linnaeus in his 1758 10th edition of Systema Naturae. [1]
H. erato exhibits Müllerian mimicry with other Heliconius butterflies such as Heliconius melpomene in order to warn common predators against attacking, which contributes to its surprising longevity. [2] [3] It also has a unique mating ritual involving the transfer of anti-aphrodisiacs from males to females. [4]
Recent field work has confirmed the relative abundance of this butterfly. [5]
H. erato is a neotropical species, found from southern Texas to northern Argentina and Paraguay, and resides on the edges of tropical rainforests. [6] [7] It is philopatric, having a particularly restricted home range. [8] In areas of dense population in Trinidad, some home ranges are only separated by 30 yards, but H. erato rarely travels to neighboring home ranges. [9] [6]
Larvae feed on the host plant, first consuming the terminal bud. After they have exhausted the resources of the plant they have hatched on, later instars may move to another plant. [7]
H. erato is a pollen-feeding species, collecting from the Lantana camara flower. They do not spend much time or energy collecting nectar (only remaining for a few seconds). Instead, they collect pollen in a mass on the ventral side of their proboscis. They then agitate the pollen by coiling and uncoiling their proboscis in order to release its nutrients. H. erato is then able to extract nitrogenous compounds in a clear liquid, including amino acids like arginine, leucine, lysine, valine, proline, histidine, isoleucine, methionine, phenylalanine, threonine, and tryptophan. Females typically carry larger loads of pollen than males as females require more amino acids for egg production. [2]
Previous studies have shown that host plants, such as Passiflora , have coevolved with Heliconius butterflies. Passiflora plants are usually found in low densities with even less plants in fruiting or flower conditions due to caterpillar feeding. [10] To increase chances of survival and cross-pollination, Passiflora plants synthesize toxins in leaves to deter Heliconius. Passiflora species produce different toxins, leading to different preferences for oviposition among Heliconius species. This leads to a lower chance of herbivore damage for individual Passiflora species and thus helps protect Passiflora plants. Chemical composition of toxins in such plants have not been studied widely. Studies have identified cyanogenic glycosides and alkaloids as potential chemicals that drive distasteful reactions among Heliconius. [11] [12] Toxin variation among Passiflora is one of the reasons for host specificity among Heliconius butterflies.
Studies have shown that H. erato species that feed on specific Passiflora species tend to spend more time on the host plant and are thus exposed to the toxins for a longer period. [12] [13] Accumulation of toxins such as cyanogenic glycosides leads to a low survival rate among H. erato larvae. Increasing exposure to parasitoids due to longer time spent on the host plant also contributes to the high mortality rate. One recent study showed that mortality increased among H. erato larvae which fed on cyanide-releasing Passiflora. Survived butterflies were capable of excreting higher levels of cyanides, suggesting a defense mechanism in H. erato. [14] H. erato species with more mechanisms to detoxify and secrete ingested toxins are the result of genetic differences among H. erato subspecies. [13] Toxin excretion, from previous studies, results in changes in wing pattern and body size. Consequences include decreased fecundity, egg size, and survival rate. [15] [16]
Nectar excretion from Passiflora has also been studied as one factor which contributes to coevolution. Passiflora nectar is known to produce aggressive behaviors among ants, wasps, and egg parasitoids. Ehrlich and Gilbert have estimated that parasitoids are capable of destroying most Heliconius eggs under nectar influence. [17] Therefore, host plants such as Passiflora are believed to have self-defense mechanisms that utilize predators against Heliconius butterflies. [10]
H. erato subspecies have innate, localized host plant preferences for oviposition. These predilections do not vary based on one's own larval host plant or with experimental conditioning. Adult females have been observed to oviposit on the meristem of their host species. Individual plant choice is based on internode length, terminal bud presence, shoot size, and leaf area, in order to confer greater larval survival advantage. In H. erato phyllis, plant choice is contingent upon terminal bud presence and condition. However, selection by quality generally depends on host plant abundance and availability. [7]
Host plants include a wide variety of passion flower ( Passiflora ) vines, including:
The red postman returns to a communal roost every night that contains members of the same species and of other heliconids. [6] The roost is typically situated about 2–10 meters from the ground on twigs and tendrils and is occupied by a small group of butterflies. [8] Adults who have just emerged from the pupa typically roost alone for a few days before roosting with others. [6]
The red postman has been observed to live in the wild for at least 20 days. [6] In captivity, they live for more than a month and have been recorded to live up to 186 days. [2] This is significantly longer than other temperate and tropical butterflies, which live for a month at best in captivity. H. erato's longevity can be explained by its benign climate and undoubted unpalatability, as well as the benefits from digesting pollen. [6]
The H. erato female lays one to four yellow eggs a day that average 1.5 mm in height and 0.9 mm in diameter. [8] [4] The eggs have a unique texture, with about 16 vertical and 11 horizontal ridges. Some plants mimic this in order to discourage females from ovipositing on them. [8]
The caterpillar appearance is very discrete when young and has a small, dark prothoracic plate. As it matures, its appearance grows more colorful. Caterpillars of H. erato chestertonii have a unique dark stripe on their side. In its fifth instar, it has a white body with black and orange spots, black spikes, and a yellow head. [8]
Pupae reside on the stem of host plants. Heliconius pupae are usually camouflaged and have defensive spikes. Pupae may be light or dark. [8]
Adult males have androconial scales on the subcostal region of their hindwings and on their median membrane. [8] Adult wingspans range from about 6.7 to 8.0 cm.
Adults have a variety of phenotypes, all with red coloration. These include: dennis-ray pattern ("dennis" refers to a red patch on the forewing; "ray" refers to red lines on the hindwing); [19] red on the forewing with yellow on the hindwing; yellow on the forewing and red on the hindwing; and white or yellow on the hindwing and forewing. [8] H. erato chestertonii is the only subspecies without any red markings, instead displaying blue.
H. erato is preyed on by birds, lizards, monkeys, and mantids, but is relatively safe due to its unpalatability and protective coloration. [9] [6]
H. erato is particularly distasteful to predators. Subspecies have evolved as Müllerian mimics, sharing aposematic patterns with other species in order to deter common predators. They typically co-mimic with other species of Heliconius, most often H. melpomene , which matches with at least 20 of the 27 subspecies. [2] [3] Subspecies have region-specific patterns that correspond to their regional mimics. H. erato chestertonii is unique as it displays blue on its wings while most other subspecies have red markings. It is the only subspecies that lacks a H. melpomene co-mimic: instead, its pattern corresponds with a subspecies of H. cydno, H. cydno gustavi. [9]
Variations from the geographical phenotype of subspecies are penalized by increased predation. In one study, researchers painted H. erato petiverana in Costa Rica to look like H. erato chestertonii from Colombia. These two subspecies successfully warn predators in their own regions with Müllerian patterns with H. melpomene rosina and H. cydno gustavi, respectively. However, the painted H. erato petiverana subjects suffered from increased predation: the H. erato chestertonii phenotype was found to be unfavorable in Costa Rica. This is because their markings did not match the Müllerian pattern of the area, so predators could not recognize their distastefulness. [9]
Listed alphabetically: [1]
The optix gene encodes the complex red coloration of Heliconius wings. An approximately 50-kb area in the intergenic region near the gene is shared by H. erato and other Heliconius, which contains cis-regulatory elements that control expression of optix. [19]
The clade containing Heliconius erato radiated before Heliconius melpomene, establishing the wing pattern diversity found in both species of butterfly. [3]
A genetic divide exists between the subspecies on either side of the Andes mountains, resulting in two distinct clades. The eastern clade is from Amazonia, southeastern Brazil, and Guiana, and consists of the subspecies dingus, emma, lativitta, phyllis, notabilis, favorinus, erato, hydara, and venustus. The western clade is from Central America and the Pacific slope of South America and consists of petiverana, hydara, venus, guarica, and cyrbia. This distinction is confirmed by sequence divergence: there is more divergence between the clades and less divergence within each clade. In addition, while there are similar haplotypes between the clades, they result in drastically different phenotypes - likely due to changes in genetic pathways for wing pattern during independent evolution. Mitochondrial DNA invariability also suggests recent radiation of these clades, probably within the last 200,000 years. These findings are consistent with the Pleistocene refugia hypothesis: in the late Pleistocene epoch, climate change reduced once widespread habitable forest areas, resulting in allopatric speciation. [3]
Males scout out females during the day and often mate with females as they emerge from the chrysalis. [8] Many males sit at female pupae waiting for them to emerge and are undisturbed by any commotion. Females mate with only one male at a time and can reproduce throughout life. [4] All subspecies can potentially mate across subspecies, but interspecies offspring are not common. These offspring only survive well in extremely specific hybrid regions and are unsuccessful elsewhere because their unusual recombinant phenotype attracts more predators. [3]
Adult males have androconial scales which disseminate pheromones to attract mates. [8] Males transfer an anti-aphrodisiac to females during copulation, which repulses and repels other potential mates from the female. It smells similar to phenylcarbylamine, or witch hazel. It emanates from two external protrusions on the abdomen of the female, which are adjacent to yellow glands that are thought to store the pheromone. The pheromone is rarely detected in males as they store it internally. The odor on females can last for weeks, even months, and is advantageous as neither sex wastes time or risks injury in subsequent matings. H. erato chestertonii has an odor distinct from other subspecies. No other Lepidoptera exhibit this behavior. [4]
H. erato has compound eyes, meaning that each eye consists of many individual photoreceptor units. H. erato eyes are unique in that they have at least five different kinds of photoreceptors and are sexually dimorphic, despite having sexually monomorphic wing patterns. (Butterflies with sexually dimorphic eyes typically have sexually dimorphic wing patterns.) The males lack protein expression of one of the SW (short-wave) opsins, which are light-sensitive proteins found in the retina. The UV discrimination conferred by this missing protein may cause males to mistake female co-mimics of other species. While inefficient, this option may have evolved because it is less costly than producing and using the UV machinery. Females, on the other hand, use this ability discriminate H. erato males from other co-mimics because they eventually invest more into egg production and can only mate with a few males. [21]
One study used amplified fragment length polymorphism (AFLP) and mitochondrial DNA (mtDNA) data sets to place the origins of H. erato at 2.8 million years ago. H. erato also shows clustering of AFLPs by geography revealing that H. erato originated in western South America. [22]
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. In the simplest case, as in Batesian mimicry, a mimic resembles a model, so as to deceive a dupe, all three being of different species. A Batesian mimic, such as a hoverfly, is harmless, while its model, such as a wasp, is harmful, and is avoided by the dupe, such as an insect-eating bird. Birds hunt by sight, so the mimicry in that case is visual, but in other cases mimicry may make use of any of the senses. Most types of mimicry, including Batesian, are deceptive, as the mimics are not harmful, but Müllerian mimicry, where different harmful species resemble each other, is honest, as when species of wasps and of bees all have genuinely aposematic warning coloration. More complex types may be bipolar, involving only two species, such as when the model and the dupe are the same; this occurs for example in aggressive mimicry, where a predator in wolf-in-sheep's-clothing style resembles its prey, allowing it to hunt undetected. Mimicry is not limited to animals; in Pouyannian mimicry, an orchid flower is the mimic, resembling a female bee, its model; the dupe is the male bee of the same species, which tries to copulate with the flower, enabling it to transfer pollen, so the mimicry is again bipolar. In automimicry, another bipolar system, model and mimic are the same, as when blue lycaenid butterflies have 'tails' or eyespots on their wings that mimic their own heads, misdirecting predator dupes to strike harmlessly. Many other types of mimicry exist.
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, who worked on butterflies in the rainforests of Brazil.
The viceroy is a North American butterfly. It was long thought to be a Batesian mimic of the monarch butterfly, but since the viceroy is also distasteful to predators, it is now considered a Müllerian mimic instead.
Polygonia c-album, or the “comma”, is a food generalist (polyphagous) butterfly species belonging to the family Nymphalidae. The angular notches on the edges of the forewings are characteristic of the genus Polygonia, which is why species in the genus are commonly referred to as anglewing butterflies. Comma butterflies can be identified by their prominent orange and dark brown/black dorsal wings.
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 Naturae. The boldly striped black and white wing pattern is aposematic, warning off predators. It is the state butterfly of Florida.
Dryas iulia, commonly called the Julia butterfly, Julia heliconian, the flame, or flambeau, is a species of brush-footed butterfly. The sole representative of its genus Dryas, it is native from Brazil to southern Texas and Florida, and in summer can sometimes be found as far north as eastern Nebraska. Over 15 subspecies have been described.
Müllerian mimicry is a natural phenomenon in which two or more well-defended species, often foul-tasting and sharing common predators, have come to mimic each other's honest warning signals, to their mutual benefit. The benefit to Müllerian mimics is that predators only need one unpleasant encounter with one member of a set of Müllerian mimics, and thereafter avoid all similar coloration, whether or not it belongs to the same species as the initial encounter. It is named after the German naturalist Fritz Müller, who first proposed the concept in 1878, supporting his theory with the first mathematical model of frequency-dependent selection, one of the first such models anywhere in biology.
The queen butterfly is a North and South American butterfly in the family Nymphalidae with a wingspan of 80–85 mm. It is orange or brown with black wing borders and small white forewing spots on its dorsal wing surface, and reddish ventral wing surface fairly similar to the dorsal surface. The ventral hindwings have black veins and small white spots in a black border. The male has a black androconial scent patch on its dorsal hindwings. It can be found in meadows, fields, marshes, deserts, and at the edges of forests.
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 a member of the Heliconiinae subfamily of Central and South America, and it is the only heliconiine that can be considered oligophagous. H. cydno is also characterized by hybridization and Müllerian mimicry. Wing coloration plays a key role in mate choice and has further implications in regards to sympatric speciation. Macrolide scent gland extracts and wing-clicking behavior further characterize this species.
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 vines (Passifloraceae). Adults exhibit bright wing color patterns which signal their distastefulness to potential predators.
Heliconius melpomene, the postman butterfly, common postman or simply postman, is a brightly colored, geographically variable butterfly species found throughout Central and South America. It was first described by Carl Linnaeus 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. 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 of the genus Heliconius and overlapping of its habitat with other related species, H. melpomene has been the subject of extensive study on speciation and hybridization. These hybrids tend to have low fitness as they look different from the original species and no longer exhibit Müllerian mimicry.
In evolutionary biology, Gilbertian mimicry is a rare type of mimicry in plants involving only two species, a host or prey animal which is the mimic, and its parasite or predator, which is both the model for the mimicry, and the dupe that is deceived by it. The mechanism provides a measure of protection for the mimic, as parasites and predators rarely attack their own species.
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 Mexico, Guatemala and Belize. H. ismenius are more commonly called the tiger-striped long wing butterfly. H. ismenius's nickname is derived from its long wing structure as well as the beautiful burnt orange and black stripes. Pierre André Latreille, a French zoologist, described Heliconius ismenius in 1817. H. ismenius resembles a number of other butterflies, an example of Müllerian mimicry.
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 an example of hybrid speciation.
Chemical mimicry is a type of biological mimicry involving the use of chemicals to dupe an operator.
Heliconius numata, the Numata longwing, is a brush-footed butterfly species belonging to the family Nymphalidae, subfamily Heliconiinae.
In evolutionary biology, mimicry in plants is where a plant evolves to resemble another organism physically or chemically. Mimicry in plants has been studied far less than mimicry in animals. It may provide protection against herbivory, or may deceptively encourage mutualists, like pollinators, to provide a service without offering a reward in return.
The evo-devo gene toolkit is the small subset of genes in an organism's genome whose products control the organism's embryonic development. Toolkit genes are central to the synthesis of molecular genetics, palaeontology, evolution and developmental biology in the science of evolutionary developmental biology (evo-devo). Many of them are ancient and highly conserved among animal phyla.
Heliconius eleuchia, the white-edged longwing, is a species of Heliconius butterfly described by William Chapman Hewitson in 1853.
Heliconius erato petiverana is a subspecies or geographical race of the red postman butterfly. It is characterized by a red forewing patch and a yellow transverse stripe on the hindwing, on an otherwise black or dark brown background.The type locality is "Mexico", and the form occurs from southern Texas to central Panama. Some lepidopterists distinguish the named forms H. erato cruentus and H. erato demophoon as separate races, based on the size of the red FW patch and the thickness of the yellow HW stripe.