Chrysiridia rhipheus

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Madagascan sunset moth
Urania riphaeus engraving.jpg
Engraving captioned Urania riphaeus from Charles D. d'Orbigny's Dictionnaire universel d'histoire naturelle (1849)
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Lepidoptera
Family: Uraniidae
Genus: Chrysiridia
Species:
C. rhipheus
Binomial name
Chrysiridia rhipheus
(Drury, 1773)
LocationMadagascar.svg
Endemic to Madagascar
Synonyms
  • Papilio rhipheusDrury, 1773 [1]
  • Urania ripheus var. madagascariensis Lesson, 1831 [2]
  • Urania crameriMaassen, 1897 [2]
  • Urania druryi Boisduval, 1874 [3]
  • Urania papageno Niepelt, 1931 [4]
  • Rhipheus dasycephalusSwainson, 1833 [5]
  • Leilus orientalisSwainson, 1833 [2]
  • Chrysiridia riphearia Hübner, 1823 [6]
  • Urania ripheus [7]
  • Urania rhiphaeus [8]
  • Urania rhipheus
  • Chrysiridia madagascariensis [8]
  • Chrysiridia rhiphaeus [9]
  • Chrysiridia ripheus

Chrysiridia rhipheus, the Madagascan sunset moth, is a species of day-flying moth of the family Uraniidae. It is considered one of the most impressive and appealing-looking lepidopterans. [10] Famous worldwide, it is featured in most coffee table books on Lepidoptera and is much sought after by collectors, though many older sources misspell the species name as "ripheus". [11] [12] The colours originate from optical interference in the iridescent parts of the wings, while the black parts are pigmented. [7] [13] Adults have a wingspan of 7–9 cm (2.8–3.5 in).

Contents

Dru Drury, who described the moth in 1773, placed it in the genus Papilio , considering it a butterfly. Jacob Hübner placed it in the moth genus Chrysiridia in 1823. Later redescriptions led to junior synonyms such as Chrysiridia madagascariensis (Lesson, 1831). [2]

At first the moth was thought to be from China or Bengal, [14] [15] but was later found to be endemic to Madagascar. It is found throughout the year in most parts of the island, with peak populations between March and August, and smallest numbers between October and December. Females lay about 80 eggs under the leaves of Omphalea spp. The caterpillars are whitish yellow with black spots and red feet and are covered in club-ended black setae. Silk spun from the mouth helps the caterpillars hold on to smooth leaves and climb back to the plant when they fall. After completing four instars, the caterpillars spin an open network cocoon. The pupal stage lasts from 17 to 23 days. [16] Chrysiridia rhipheus is the sole specialist herbivore of the four species of Omphalea in Madagascar. Omphalea is toxic: the toxins are sequestered by the feeding caterpillar and retained in the pupal and adult stages. [17] Thousands of these moths migrate between the eastern and western ranges of their host plants. [16] [17]

Taxonomy

The Madagascan sunset moth was first described by the British entomologist Dru Drury in 1773 from a specimen obtained from a Captain May of Hammersmith. Because of its resemblance to swallowtail butterflies, Drury described the species as Papilio rhipheus. [1] 1 The specimen Drury received had the head of a different species attached, probably that of a butterfly, which had clubbed antennae [5] —this trait is frequently used to differentiate moths from butterflies. Once the inaccuracy in Drury's specimen was found, the moth was placed in the genus Urania , until 1823 when the German entomologist Jacob Hübner placed it in a new genus, Chrysiridia. The moth has also been described under other names. Because Drury described his specimen as having clubbed antennae and being tailless, William Swainson thought it was a different species than the complete specimen described by Pieter Cramer. In 1833, Swainson named the butterfly Rhipheus dasycephalus and the moth Leilus orientalis. [5] Other synonyms include U. crameri by Maassen in 1879 and U. ripheus var. madagascariensis by Lesson in 1831. [2]

Native Malagasy people call it adriandolo or lolonandriana, [8] [18] from lolo for "spirit" or "butterfly" and andriana for "noble" or "king", [19] therefore meaning "noble butterfly", [8] "noble spirit", [20] "king butterfly" [18] or "king spirit".

The genus Chrysiridia is entirely African and the only other species in the genus is the East African C. croesus . Chrysiridia is one of three diurnal uraniine genera. The other two genera are Urania, its sister taxon, and Alcides, the most basal. In the group, the use of Endospermum is an ancestral state (a plesiomorphy). The more basal Alcides feed on Endospermum and Omphalea , while Urania and Chrysiridia feed only on Omphalea. [17]

Description

Both sides of Chrysiridia rhipheus Chrysiridia rhipheus MHNT.jpg
Both sides of Chrysiridia rhipheus

Chrysiridia rhipheus has a wingspan of 7–9 centimetres (2.8–3.5 in), and sometimes up to 11 centimetres (4.3 in). Moths from the highlands, 900–1,080 metres (2,950–3,540 ft), have a median wingspan of 7 cm (2.8 in); moths from lower altitudes, 600 m (2,000 ft), have a median wingspan of 9 cm (3.5 in). [16] Like many other uraniine moths, the sunset moth has an uncanny resemblance to swallowtail butterflies, especially in its tails and colourful wings, and can easily be mistaken for a butterfly. [21]

The sunset moth is black with iridescent red, blue and green markings. There is a fringe of white scales on the wing edges, wider on the hindwings. The moth has six tails, very often lost or damaged during its life. [16] Pattern variations are common, and the moth is often partly asymmetrical; [22] one of the factors causing this is temperature shock during the pupal stage. [16]

Wing microstructure

Chrysiridia rhipheus scale detail Urania ripheus scale detail - 35x - Flickr - Thomas Shahan 3.jpg
Chrysiridia rhipheus scale detail

Unlike in many other moths, the colour of C. rhipheus's wings does not come from pigments. The colour originates from coherent scattering and interference of light by the microstructure of the ribbon-like scales covering the moth's wings. [7] [13] These structural characteristics make this species and its Neotropical relative Urania common subjects of research in optical science. [13]

The colours on the Madagascan sunset moth's wings are produced by the conjunction of two optical phenomena:

  1. An air-cuticle multilayer in the scales creates optical interference. Each scale contains cuticle layers with randomly located blocks of cuticle that hold them in place and maintain an air gap between them. The layers and air gaps are narrower than the wavelength of visible light. The structure varies from one layer at the proximal end of each scale, to about six layers at the distal end. This multilayer structure strongly reflects certain wavelengths of light, which are determined by the thicknesses of the layers and the angle at which the light hits the scale.
  2. The scales are highly curved, which creates inter-scale reflection. This mechanism is unusual among Lepidoptera. The proximal part of each scale is almost flat; the scale then gradually curves up and then steeply bends down at the distal end, covering the proximal part of the next row of scales. Because of this shape, adjacent rows of scales have valley-like grooves between them. This allows light to strike one scale, reflect at a roughly 90° angle, strike the next scale, and be reflected away from the wing. Because the angle of each reflection is far from normal incidence, the interference effect favours reflection of different colours of light than are seen when light strikes near the top of the curved part of the scale.

The colour seen in each part of the wing is the combination of the colours reflected by these two effects.

Because the cuticle layers are arranged in rows, with a cylindrical curve, the amount of the second type of reflection is dependent on the polarization of the incoming light. This makes the moth's overall colour polarization-dependent. Many insects can detect polarization of light, so it has been proposed that this feature may be used as a visual signal between moths. [7] This has not yet been studied, however.

Distribution and habitat

Distribution of the four species of Omphalea in Madagascar Omphalea distribution Madagascar.svg
Distribution of the four species of Omphalea in Madagascar

Drury's specimen was given to him by Captain May of Hammersmith and believed to have come from China. [14] Cramer believed the specimen came from Chandernagore in Bengal, however, giving rise to the French name "page de Chadernagor". [15] It is now known that Chrysiridia rhipheus is endemic to Madagascar. Thousands of adult moths seasonally migrate between geographically isolated populations of their host plant Omphalea spp. They can be found almost everywhere on the island, except in the south-west and the extreme subdesertic south of the Androy where their host plant is absent. They migrate from the three species in the dry deciduous forest in the west (O. ankaranensis, O. occidentalis and O. palmata) to the eastern rainforest species (O. oppositifolia). The western species are largely in protected areas. The eastern species, on the other hand, is mostly unprotected and dispersed in widely scattered populations threatened by deforestation. Being the only evergreen species, O. oppositifolia is probably crucial for the moth's survival. The Jamaican moth Urania sloanus , from the same subfamily (Uraniinae), most likely became extinct after the loss of one of its host plant species. [16] [17]

The moths migrate in response to changes in the host plants. Chrysiridia larvae defoliate the whole plant, and even eat the flowers and fruit, and thus have a considerable negative impact on the reproduction and survival of seedlings. [17] The plants probably react by changing their nutrient and secondary compound levels, becoming toxic to the larvae and causing high mortality. Omphalea populations that are not damaged by moths for long periods of time have lower toxicity. [11] These factors cause mass increases in local population, followed by sudden crashes. The population crashes might result from increased larval mortality, but are more likely caused by the emigration of the adult moths. Through semiochemicals, the plant may recruit hymenopteran parasitoids as a protection, hence playing a role in the population dynamics of the moth. [17]

Host plants

Like C. croesus and species of the genus Urania, C. rhipheus is a specialist species whose caterpillars feed strictly on species of the pantropical genus Omphalea (Euphorbiaceae). [17] Four species of the genus Omphalea are endemic to Madagascar:

Omphalea, like many other members of the Euphorbiaceae, possesses leaf nectaries that attract polistine wasps, which are predators of early instar caterpillars. The leaf nectaries also attract ants. The ants usually protect their host plant, eating both the nectar and plant-eating insects on it. However, they generally completely ignore Chrysiridia caterpillars, making it likely that the caterpillars possess a chemical deterrent as a primary defence. [17] This chemical deterrent comes from the host plants. Omphalea species contain polyhydroxy alkaloids potentially sequestered or excreted by the caterpillar, the pupa and the adult moth. [24]

Behaviour

Unlike most moths, the sunset moth is day flying and the bright aposematic colours warn predators of its toxicity, a strategy seen in many diurnal moths. Another habit the moth has in common with many butterflies is the night resting posture—the wings are held vertically over the back. During their migrations they roost in a group for the night. [17] The moth flies over the crown of trees and in clearings. Sometimes because of high winds it is carried about 100 m (300 ft), it then falls, inert and wings closed, to the ground. [16]

Life cycle

Final instar larva, shown here on the leaf of Omphalea oppositifolia Chrysiridia rhipheus larva.jpg
Final instar larva, shown here on the leaf of Omphalea oppositifolia
Adult Chrysiridia rhipheus at Madagascar near Mahajanga Chrysiridia rhipheus-mada-alive=vivant.jpg
Adult Chrysiridia rhipheus at Madagascar near Mahajanga

Continuous generations of the moth are present all year. The highest populations are found from March to August (fall and winter), while the lowest are from October to December (spring). [16] The females lay their eggs late in the afternoon or at nightfall, and locate potential oviposition sites visually. [17] Like the eggs of other Uraniidae, the sunset moth's eggs are domed with projecting ribs. [25] A single egg weighs about 1 milligram (3.5×10−5  oz ) and usually has 17 ribs, but sometimes 18 or less often 16. The eggs are usually laid on the lower surface of Omphalea leaves, but occasionally on the upper surface. Eggs are laid in groups of 60 to 110, usually about 80. [16]

After they hatch, the small caterpillars only eat the tissue (parenchyma) between the veins of the leaves. They do this to avoid the sticky and toxic latex produced by the plant's laticifers and transported in the veins. After 3–4 days, the caterpillars also eat flowers, fruit, tendrils, petioles and young stems (as well as continuing to eat leaves), defoliating the entire plant. They are particularly fond of the glands at the base of the leaf, near the petiole. They can deal with the chemical defences in the latex, which does not cause the problem of mouthpart coagulation. [16] [17] The caterpillars spin silk from their mouth with an '∞' motion of the head as they walk, this keeps them from falling from the smooth surface of the leaves. The silk also permits them to climb back to the plant should they fall. Strong rain makes them fall despite the silk. There are four instars, and the caterpillar stage lasts from two months in the warm season to two and a half to three months in the cold season. [16] The caterpillar is whitish yellow with black spots and red feet and is covered in club-ended black setae. [26] It has five pairs of prolegs on segments 3 to 6 and 10, and six true legs attached to the thorax. [27]

After completing all but its last moult, the caterpillar spins a cocoon out of silk. The cocoon can be in the tree crown or between two leaves, but is most often near the ground, between moss and bark. It is an open network cocoon with large and irregular mesh. [16] [26] [28] In the warm season, the cocoon takes about 10 hours to spin, the metamorphosis takes place about 29 hours later and lasts about 6 minutes. These durations are slightly longer in the cold season. The chrysalis stage lasts 17 days in November, the warmest month, and 23 days in July, the coldest month. Five to six days before eclosion, the motifs of the wings start to become visible. The moth emerges during the night or in low light, by splitting the pupal case from the top. Once out of the pupal exuvia (the pupal exoskeleton), the moth finds a horizontal surface, from which it suspends itself by its four anterior legs. The wings are deployed in about 10 minutes, by pumping haemolymph into the wing veins. The moth then beats them a few times, waits 45 minutes to let them harden, then beats them lightly again. The moth finally takes flight between one-and-a-half and two hours later. [16]

Nectar sources

Adult moths prefer white or whitish-yellow flowers as a nectar source, which indicates that visual cues play a large role in their selection. Most flowers visited are inflorescences of small flowers or have dense filaments, giving them the appearance of a bottle brush, often because of the conspicuous projecting stamens as in many Leguminosae (Mimosoideae, Myrtaceae and Combretaceae). However, not all white flowers elicit a response: the white and showy flowers of Omphalea oppositifolia are not visited by the adult moths. Nectar sources include the flowers of:

All these flowers are white, with the exception of Camellia sinensis which has a yellow centre, and all either have dense filaments or are formed of tight clusters of small flowers. [17]

In culture

A cigarette card featuring a "Chrysiridia madagascariensis" fairy in 1928 Chrysiridia Cigarette card.jpg
A cigarette card featuring a "Chrysiridia madagascariensis" fairy in 1928

This spectacular moth is considered one of the most impressive and beautiful Lepidoptera, [10] rivalling almost any of the butterflies in brilliance of colouring and form. [21] It is featured in most coffee table books on the Lepidoptera, [11] and is much sought after by collectors. [12] It is collected in the wild, and raised commercially for the international butterfly trade; its wings were used to make jewellery in the Victorian era. [26] The Madagascan sunset moth appeared on a 6 maloti postage stamp in the Lesotho Postal Services Butterflies of Africa issue of 20 August 2007. [29] Only one of the four species of host plants, Omphalea oppositifolia, is used to raise the moth commercially, mainly using plants collected in the wild, but also some cultivated for the purpose. [23]

In Malagasy, lolo is polysemous for "butterfly" or "moth" and "soul". There is little doubt that this is because a pupa resembles a covered corpse and that the adult emerges from it—like the soul from body of the dead. [30] [31] The Malagasy people believe the soul of the dead or of ancestors appears in the form of a lepidopteran, and thus to attack it is to attack the ancestors. [32] [33]

Footnotes

^1 The meaning of the specific epithet rhipheus was not given by the author Dru Drury. [14] Possibilities include the Latin Montes Rhipheaus, the Ural Mountains [34] or Ripheus, "the most just of all the Trojans". [35]

Related Research Articles

<span class="mw-page-title-main">Lepidoptera</span> Order of insects including moths and butterflies

Lepidoptera or lepidopterans is an order of winged insects that includes butterflies and moths. About 180,000 species of the Lepidoptera have been described, representing 10% of the total described species of living organisms, making it the second largest insect order with 126 families and 46 superfamilies, and one of the most widespread and widely recognizable insect orders in the world.

A common classification of the Lepidoptera involves their differentiation into butterflies and moths. Butterflies are a natural monophyletic group, often given the suborder Rhopalocera, which includes Papilionoidea, Hesperiidae (skippers), and Hedylidae. In this taxonomic scheme, moths belong to the suborder Heterocera. Other taxonomic schemes have been proposed, the most common putting the butterflies into the suborder Ditrysia and then the "superfamily" Papilionoidea and ignoring a classification for moths.

<span class="mw-page-title-main">Moth</span> Group of mostly-nocturnal insects in the order Lepidoptera

Moths are a group of insects that includes all members of the order Lepidoptera that are not butterflies. They were previously classified as suborder Heterocera, but the group is paraphyletic with respect to butterflies and neither subordinate taxon is used in modern classifications. Moths make up the vast majority of the order. There are approximately 160,000 species of moth, many of which have yet to be described. Most species of moth are nocturnal, although there are also crepuscular and diurnal species.

<span class="mw-page-title-main">Pupa</span> Life stage of some insects undergoing transformation

A pupa is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their life cycle, the stages thereof being egg, larva, pupa, and imago. The processes of entering and completing the pupal stage are controlled by the insect's hormones, especially juvenile hormone, prothoracicotropic hormone, and ecdysone. The act of becoming a pupa is called pupation, and the act of emerging from the pupal case is called eclosion or emergence.

<span class="mw-page-title-main">Uraniidae</span> Family of moths

The Uraniidae are a family of moths containing four subfamilies, 90 genera, and roughly 700 species. The family is distributed throughout the tropics of the Americas, Africa and Indo-Australia. Some of the tropical species are known for their bright, butterfly-like colors and are called sunset moths. Such moths are apparently toxic and the bright colors are a warning to predators.

<i>Omphalea</i> Genus of flowering plants

Omphalea is a plant genus of the family Euphorbiaceae first described as a genus in 1759. It is native to tropical parts of the Americas, the West Indies, Asia, Australia, and Africa.

<i>Papilio demoleus</i> Species of butterfly

Papilio demoleus is a common and widespread swallowtail butterfly. The butterfly is also known as the lime butterfly, lemon butterfly, lime swallowtail, and chequered swallowtail. These common names refer to their host plants, which are usually citrus species such as the cultivated lime. Unlike most swallowtail butterflies, it does not have a prominent tail. When the adult stage is taken into consideration, the lime swallowtail is the shortest-lived butterfly, with male adults dying after four days and females after a week. The butterfly is native to Asia and Australia, and can be considered an invasive pest in other parts of the world. The butterfly has spread to Hispaniola island in the Western Hemisphere, and to Mahé, Seychelles.

<span class="mw-page-title-main">Insect migration</span> Seasonal movement of insects

Insect migration is the seasonal movement of insects, particularly those by species of dragonflies, beetles, butterflies and moths. The distance can vary with species and in most cases, these movements involve large numbers of individuals. In some cases, the individuals that migrate in one direction may not return and the next generation may instead migrate in the opposite direction. This is a significant difference from bird migration.

<span class="mw-page-title-main">Uraniinae</span> Subfamily of moths

The Uraniinae or uraniine moths are a subfamily of moths in the family Uraniidae. It contains seven genera that occur in the tropics of the world.

<i>Hemaris thysbe</i> Species of moth

Hemaris thysbe, the hummingbird clearwing, is a moth of the family Sphingidae (hawkmoths). Coloration varies between individuals, but typically the moth is olive green and burgundy on its back, and white or yellow and burgundy on the underside. Its wings are transparent with a reddish-brown border. It has light-colored legs, which combined with the lack of striping on the underside is diagnostic. Beating its wings rapidly, H. thysbe hovers to collect nectar from a variety of flowers. The combination of its appearance and its behavior commonly leads to it being confused with a hummingbird or bumblebee.

<i>Callosamia promethea</i> Species of moth

Callosamia promethea, commonly known as the promethea silkmoth, is a member of the family Saturniidae, which contains approximately 1,300 species. It is also known as the spicebush silkmoth, which refers to one of the promethea silkmoth's common host plants, spicebush. C. promethea is classified as a silk moth, which stems from its ability to produce silk, which it does in the formation of its cocoon. C. promethea lives in forests in the eastern U.S. and does not damage the trees on which it lives. The species was first described by Dru Drury in 1773.

<i>Urania sloanus</i> Extinct species of moth

Urania sloanus, or Sloane's urania, was a species of moth of the family Uraniidae endemic to Jamaica. It was last reported in 1894 or 1895, but possibly survived until at least 1908. The species was first described by Pieter Cramer in 1779.

<i>Acronicta rumicis</i> Species of moth

Acronicta rumicis, the knot grass moth, is a species of moth which is part of the genus Acronicta and family Noctuidae. It was first described by Carl Linnaeus in his 1758 10th edition of Systema Naturae. It is found in the Palearctic region. A. rumicis lives and feeds on plants located in wide-open areas. At its larval stage, as a caterpillar, it causes such a large impact as a crop pest that it has received much attention and research. A. rumicis feeds on maize, strawberries and other herbaceous plants.

<i>Sarisa <span style="font-style:normal;">(moth)</span></i> Genus of moths

Sarisa is a monotypic moth genus in the family Geometridae and was first described by David Stephen Fletcher in 1979. The genus contains only one species, Sarisa muriferata, the hook-tip fern looper, which is endemic to New Zealand and surrounding islands. This species was described by Francis Walker in 1862. It is widespread in the North and South Islands, and has been recorded from Stewart Island, Big South Cape Island, the Chatham Islands and the Auckland Islands.

<span class="mw-page-title-main">External morphology of Lepidoptera</span> External features of butterflies and moths

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 of scales on the external parts of the body and appendages, especially the wings. Butterflies and moths vary in size from microlepidoptera only a few millimetres long, to a wingspan of many inches such as the Atlas moth. Comprising over 160,000 described species, the Lepidoptera possess variations of the basic body structure which has evolved to gain advantages in adaptation and distribution.

<span class="mw-page-title-main">Lepidoptera migration</span>

Many populations of Lepidoptera migrate, sometimes long distances, to and from areas which are only suitable for part of the year. Lepidopterans migrate on all continents except Antarctica, including from or within subtropical and tropical areas. By migrating, these species can avoid unfavorable circumstances, including weather, food shortage, or over-population. In some lepidopteran species, all individuals migrate; in others, only some migrate.

C. madagascariensis may refer to:

<i>Chrysiridia</i> Genus of moths

Chrysiridia is a genus of uraniid moths from Africa including Madagascar. The genus was erected by Jacob Hübner in 1823. They are diurnal and strongly marked with iridescent colours.

<i>Urania boisduvalii</i> Species of moth

Urania boisduvalii is a day-flying moth of the family Uraniidae. It was first described by Félix Édouard Guérin-Méneville in 1829. A genetic analysis of Urania moths gave rise to a phylogenetic tree which places U. boisduvalii as sister to the in-group that includes U. fulgens spp. poeyi, U. fulgens, U. sloanus, U. sloanus, U. leilus and U. leilus spp. brasiliensis.

<i>Lyssa menoetius</i> Species of moth

Lyssa menoetius is a moth of the family Uraniidae. The species was first described by German entomologist Hopffer in 1856.

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