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==Description==
==Description==
''Chrysiridia rhipheus'' (Drury, 1773) has a wingspan of {{nowrap|7 to 9 centimetres}} {{nowrap|(3 to 3½ in)}}, but sometimes up to {{nowrap|11 centimetres}} {{nowrap|(4 in)}}. Moths from high altitude, {{nowrap|900 to 1080 metres}} {{nowrap|(3000-3500 ft)}}, have a median wingspan of 7&nbsp;centimetres (3&nbsp;in); While moths from lower altitude, 600&nbsp;metres (2000&nbsp;ft), have a median wingspan of 9&nbsp;centimetres (3½&nbsp;in).<ref name=Catala/> As many of the [[Uraniinae|uraniine moths]], the sunset moth has an uncanny resemblance to [[Swallowtail butterfly|swallowtail butterflies]] because of its tails and colourful wings, both characteristics common to the latter. For those reasons it is easy to mistaken it for a [[butterfly]].<ref name = Klots>{{cite book |last=Klots |first=Alexander Barrett |coauthors=Elsie Broughton Klots |title=Living Insects of the World |origyear=1959 |url=http://books.google.com/books?id=odFFGQAACAAJ&dq=swallowtail+Chrysiridia&source=gbs_book_other_versions_r&cad=1_2 |accessdate=2008-02-19 |date=2007-05-25 |publisher=[[Doubleday (publisher)|Doubleday]]}}</ref>
''Chrysiridia rhipheus'' (Drury, 1773) has a wingspan of {{nowrap|7 to 9 [[centimetre|cm]]}} {{nowrap|(3 to 3½ [[inch|in]])}}, but sometimes up to {{nowrap|11 cm}} {{nowrap|(4 in)}}. Moths from high altitude, {{nowrap|900 to 1080 [[metre|m]]}} {{nowrap|(3000-3500 [[foot|ft]])}}, have a median wingspan of 7&nbsp;cm (3&nbsp;in); While moths from lower altitude, 600&nbsp;m (2000&nbsp;ft), have a median wingspan of 9&nbsp;cm (3½&nbsp;in).<ref name=Catala/> As many of the [[Uraniinae|uraniine moths]], the sunset moth has an uncanny resemblance to [[Swallowtail butterfly|swallowtail butterflies]] because of its tails and colourful wings, both characteristics common to the latter. For those reasons it is easy to mistaken it for a [[butterfly]].<ref name = Klots>{{cite book |last=Klots |first=Alexander Barrett |coauthors=Elsie Broughton Klots |title=Living Insects of the World |origyear=1959 |url=http://books.google.com/books?id=odFFGQAACAAJ&dq=swallowtail+Chrysiridia&source=gbs_book_other_versions_r&cad=1_2 |accessdate=2008-02-19 |date=2007-05-25 |publisher=[[Doubleday (publisher)|Doubleday]]}}</ref>


It is black with iridescent red, blue and green markings. A band of white scales fringes the wings, and is winder on the hindwings. It has six tails, that are very often lost during the moth's life.<ref name=Catala>{{cite journal |last=Catala |first=R. |year=1940 |title=Variations expérimentales de ''Chrysiridia madagascariensis'' Less. (Lep. Uraniidae) |journal=Archives du [[Muséum national d'histoire naturelle|Muséum National D'Histoire Naturelle]] |volume=17 |pages=Ph.D. Thesis}}</ref> Pattern variations are common, and the moth is often partly asymmetrical,<ref>{{cite web |url=http://nauka.relis.ru/30/0201/30201141.htm |title=Лицом к лицу с природой |accessdate=2008-03-18 |last=Kaabak |first=L. |date= |work= |publisher= |language=Russian}}</ref> One of the factors for this is temperature shocks during the pupal stage.<ref name=Catala/>
It is black with iridescent red, blue and green markings. A band of white scales fringes the wings, and is winder on the hindwings. It has six tails, that are very often lost during the moth's life.<ref name=Catala>{{cite journal |last=Catala |first=R. |year=1940 |title=Variations expérimentales de ''Chrysiridia madagascariensis'' Less. (Lep. Uraniidae) |journal=Archives du [[Muséum national d'histoire naturelle|Muséum National D'Histoire Naturelle]] |volume=17 |pages=Ph.D. Thesis}}</ref> Pattern variations are common, and the moth is often partly asymmetrical,<ref>{{cite web |url=http://nauka.relis.ru/30/0201/30201141.htm |title=Лицом к лицу с природой |accessdate=2008-03-18 |last=Kaabak |first=L. |date= |work= |publisher= |language=Russian}}</ref> One of the factors for this is temperature shocks during the pupal stage.<ref name=Catala/>

Revision as of 04:39, 18 June 2008

Madagascan sunset moth
Engraving captioned Urania riphaeus from Charles D. d'Orbigny’s Dictionnaire universel d'histoire naturelle (1849)
Scientific classification
Kingdom:
Phylum:
Class:
Order:
Superfamily:
Family:
Subfamily:
Genus:
Hübner, 1823
Species:
C. rhipheus
Binomial name
Chrysiridia rhipheus
(Drury, 1773) originally as Papilio rhipheus[1]
Contained in Madagascar
Synonyms

Urania ripheus var. madagascariensis Lesson, 1831[2]
Urania crameri Maassen, 1897[2]
Rhipheus dasycephalus Swainson, 1833[3] Leilus orientalis Swainson, 1833[2]
Chrysiridia riphearia Hübner, 1823[4]
Urania ripheus[5]
Urania rhiphaeus[6]
Urania rhipheus
Chrysiridia madagascariensis[6]
Chrysiridia rhiphaeus[7]
Chrysiridia ripheus

The Madagascan sunset moth, or simply sunset moth (Chrysiridia rhipheus, also known as Urania rhipheus, U. rhiphaeus,[6] C. rhiphaeus,[7] and C. madagascariensis[6]) is a diurnal moth of the Uraniidae family. It is very colourful, but the iridescent parts of the wings do not have pigment; the colors originate from refraction instead.[8] It is considered to be one of the most impressive and beautiful Lepidoptera,[9] for this reason it has gained an international reputation, is much sought by collectors,[10] and is featured in most coffee table books on the Lepidoptera.[11] It is the sole herbivore of the four species of the toxic shrubs and trees of the genus Omphalea in its native Madagascar; this toxicity is passed on by the feeding caterpillar to the pupa and the adult moth.[12]

Taxonomy

The Madagascan sunset moth was first described by British entomologist Dru Drury in 1773. Because of its resemblance to swallowtail butterflies, Drury had first described the species as Papilio rhipheus,[1] after he mistakenly thought its antennae were clubbed;[3] this characteristic is frequently used to differentiate moths from butterflies. Once Drury’s inaccuracy was found, it was placed in the genus Urania, until 1823 when German entomologist Jacob Hübner placed it in a new genus; Chrysiridia. It also has also been given other names, including U. crameri by Maassen in 1879 and U. ripheus var. madagascariensis by Lesson in 1831.[2]

Native Malagasy people call it Adriandolo or Lolonandriana,[6][13] from lolo for spirit or butterfly and andriana for noble or king,[14] therefore meaning noble butterfly,[6] noble spirit,[15] king butterfly[13] or king spirit.

The genus Chrysiridia is entirely African as the only other species is the closely related East African C. croesus. It is it, with Urania, sister-taxa to the most basal diurnal uraniine genus Alcides. All three genera feed on Omphalea, but Alcides also feeds on Endospermum.[12]

Description

Chrysiridia rhipheus (Drury, 1773) has a wingspan of 7 to 9 cm (3 to 3½ in), but sometimes up to 11 cm (4 in). Moths from high altitude, 900 to 1080 m (3000-3500 ft), have a median wingspan of 7 cm (3 in); While moths from lower altitude, 600 m (2000 ft), have a median wingspan of 9 cm (3½ in).[16] As many of the uraniine moths, the sunset moth has an uncanny resemblance to swallowtail butterflies because of its tails and colourful wings, both characteristics common to the latter. For those reasons it is easy to mistaken it for a butterfly.[17]

It is black with iridescent red, blue and green markings. A band of white scales fringes the wings, and is winder on the hindwings. It has six tails, that are very often lost during the moth's life.[16] Pattern variations are common, and the moth is often partly asymmetrical,[18] One of the factors for this is temperature shocks during the pupal stage.[16]

Chrysiridia rhipheus underside
Chrysiridia rhipheus topside

Wing microstructure

The iridescent parts of the wings do not have pigment; the colour originates from refraction of light by the microstructures of the ribbon-like scales covering the moth’s wings. These structural characteristics make this species and its neotropical relative Urania common subjects of optical sciences.[8]

The iridescence on the Madagascan sunset moth's wings is produced by the alliance of two optic phenomena:

  • The first mechanism is an air-cuticle-multilayer that creates optical interference. Each scale is composed of one to six cuticle layers, depending on the portion of the scale, separated by air and held by connecting blocks.
  • The second mechanism is more unusual among Lepidoptera: The scales are highly curved, which creates a mechanism of inter-scale reflection. The proximal part of each scale is almost flat, 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. Light is reflected from one row to the next, which makes the color more polarization dependent.

Although the biological function of this polarization-dependent change in color has not been studied, it may work as a visual signal among the species. That is if the insects possess polarization and color vision.[5]

Distribution and habitat

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 from the dry deciduous forest in the west (O. ankaranensis, O. occidentalis, and O. palmata) to the eastern rainforest species (O. oppositifilia). The western species are largely protected. The eastern species on the other hand is mostly unprotected and is dispersed in widely scattered populations threatened by deforestation. Being the only evergreen species, O. oppositifilia is probably crucial for the moth’s survival. The Jamaican moth Urania sloanus, from the same subfamily (Uraniinae), probably became extinct after the loss of one species of its host plants.[12][16]

The moths migrate in response to changes in the host plants. Chrysiridia larvae defoliate the whole plant, and even eat flowers and fruit, thus having a considerable negative impact on the reproduction and survival of seedlings.[12] The plants probably react by changing the nutrient and secondary compound levels. They become toxic to the larvae and cause high mortality levels. Inversely Omphalea who do not sustain damage by predation for long periods have lowered moth-induced defences.[11] This allows for mass local emergences, followed by apparent population crashes. The population crashes are probably caused by the emigration of adults, or to a lesser probability, the high mortality levels in larvae. Through allelochemistry, the plant may recruit hymenopteran parasitoids as a protection, hence playing a role in the population dynamics of the moth.[12]

Host plants

Distribution of the four Madagascan species of Omphalea

Like C. croesus and species of the genus Urania, C. rhipheus is a specialist species whose larvae feed strictly on species of the pantropical genus Omphalea (Euphorbiaceae).[12] Four species of the genus Omphalea are endemic to Madagascar:

  • O. ankaranensis, a shrub from the limestone karst of northern Madagascar
  • O. palmata, a dry forest shrub, closely related to O. ankaranensis, but from western Madagascar
  • O. occidentalis, also a dry forest species of western Madagascar
  • O. oppositifilia, a tree from the east coast rainforest of Madagascar[19]

All Omphalea, in common with many other Euphorbiaceae, possess leaf nectaries attracting polistine wasps, which are known to be effective predators of early instar larvae. The leaf nectaries also attract ants. The ants usually protect their host plant, eating both the nectar and plant-eating insects on it. But they generally ignore completely Chrysiridia larvae, making it likely that the larvae possess a chemical deterrent as a primary defence.[12] This chemical deterrent comes from the host plants. The species from the genus contain polyhydroxy alkaloids potentially sequestered or excreted by the caterpillar, the pupa and the moth.[20]

Behaviour

The sunset moth is day flying while most moths are active at night. It is also toxic, as revealed by its aposematic colours, a common defence found in several 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.[12] The moth flies over the crown of trees and in clearings. Sometimes because of high winds it is carried about one hundred metres (300 ft), it then falls, inert and wings closed, to the ground.[16]

Life cycle

The moth has continuous generations and is present all year. The highest populations are found from March to August, while the lowest are from October to December.[16] The females lay their eggs late in the afternoon or at nightfall, and locate potential oviposition sites visually.[12] Like the eggs of other Uraniidae, the sunset moth’s are domed with projecting ribs.[21] There are usually 17 ribs, but sometimes 18 and less often 16. The eggs are usually laid under leaves of Omphalea, also but rarely on top. Eggs are laid in groups of 60 to 110, but more often 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 three to four days instars still eat leaves but also flowers, fruit, tendrils, petioles and young stems, 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 and it does not cause problems of mouthpart coagulation.[12][16] The caterpillars spin silk from their mouth with an ‘∞’ motion as they walk, this keeps them from falling on the smooth surface of the leaves. The silk also permits them to climb back to the plant when they fall. But a strong rain makes them fall despite the silk. There are four instars, and the caterpillar stage lasts form two months in the warm season and 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.[22] It has five pairs of prolegs on the segments 3 to 6 and 10, with its six legs it has a total of sixteen.[23]

After completing all but its last moult, the caterpillar spins a cocoon out of silk. The cocoon can also be in the crown of trees, between two leaves, but more often near the ground, between moss and bark. It is an open network cocoon with large and irregular mesh.[22][24][16] The cocoon takes about ten hours to spin, the metamorphosis takes place about twenty-nine hours later and lasts about six minutes. The times are slightly longer in the cold season. The chrysalis stage lasts seventeen days in November, the warmest month, and twenty-three days in July, the coldest month. Five to six days before eclosion the motives of the wings start to become visible. The moth emerges during the night or in low light, it splits the pupal case from the top. Once out of the pupal exuvia (the pupal exoskeleton), the moth looks for a horizontal surface to suspend itself with its four anterior legs. The wings are deployed in about ten minutes, by pumping haemolymph into the wing veins. Once the wings are deployed the moth beats them a few times, waits forty-five minutes, then beats them lightly again. The moth finally takes flight one hour and a half to 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). But not all white flowers elicit a response: the white and showy flowers of Omphalea oppositifolia aren’t 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.[12]

Relationship with humans

A cigarette card featuring a Chrysiridia madagascariensis fairy in 1928

The spectacular moth is featured in most "coffee table" books on the Lepidoptera,[11] and has an international reputation.[10] It is considered to be one of the most impressive and beautiful Lepidoptera,[9] rivalling almost any of the butterflies in brilliance of coloring and form.[17] For these reasons it is much sought by collectors.[10] It is collected in the wild, and raised commercially for the international butterfly trade. Only one of the four species of host plants, Omphalea oppositifolia, is used to raise the moth commercially. Using mainly wild collected plants, but also cultivated at a small scale.[19]

It was known by Victorians who used its wings to make jewellery.[22]

In Malagasy lolo is polysemous for ‘butterfly’ or ‘moth’ and ‘soul’, there is little doubt that this is because a chrysalis resembles a covered corpse and that the butterfly or moth emerges from it, like the soul from body of the dead.[25][26] They believe the soul of the dead or of ancestors appears in the form of a Lepidoptera, to attack it is to attack the ancestors.[27][28] The Madagascan sunset moth in particular is called Adriandolo or Lolonandriana, from Andriana meaning ‘sovereign, prince, noble' or 'king’ and lolo.[13] Therefore meaning, depending on the translation, noble butterfly,[6] noble spirit,[15] king butterfly or king spirit.[13]

References

  1. ^ a b Belouino, Paul (1852). Dictionnaire général et complet des persécutions souffertes par l'Église (in French). Paris: J.P. Migne Éditeur. pp. 1784–1786. Retrieved 2008-02-16.
  2. ^ a b c d Kirby, William Forsell (1897). A Hand-book to the Order Lepidoptera (PDF). Baldwin & Cradock. pp. 50–53. Retrieved 2008-03-21.
  3. ^ a b Swainson, William S. (1833). Zoological Illustrations. Vol III. Second series. London: Baldwin & Cradock. pp. 130–131. Retrieved 2008-02-19.
  4. ^ Savela, Markku (2007-03-15). "Chrysiridia". Lepidoptera and some other life forms. Retrieved 2008-06-16.
  5. ^ a b Yoshioka, Shinya (March 5, 2007). "Polarization-sensitive color mixing in the wing of the Madagascan sunset moth" (PDF). Optics Express. 15 (5). Washington: Optical Society of America: 2691–2701. doi:10.1364/OE.15.002691. Retrieved 2008-02-18. {{cite journal}}: Check date values in: |date= (help); Unknown parameter |coauthors= ignored (|author= suggested) (help); Unknown parameter |laysource= ignored (help); Unknown parameter |laysummary= ignored (help); Unknown parameter |quotes= ignored (help)
  6. ^ a b c d e f g Decary, Raymond (1950). La Faune Malgache (in French). Paris: Payot.
  7. ^ a b Biosecurity New Zealand (2005-08-24). "Import Health Standard for the Importation of Tropical Butterfly and Moth Pupae into New Zealand". Commercial Imports. New Zealand Government. Retrieved 2006-11-08.
  8. ^ a b Prum, Richard O. (January 31, 2006). "Anatomically diverse butterfly scales all produce structural colours by coherent scattering" (PDF). The Journal of Experimental Biology. 209. Cambridge: The Company Of Biologists: 748–765. doi:10.1242/jeb.02051. Retrieved 2008-02-16. {{cite journal}}: Check date values in: |date= (help); Unknown parameter |coauthors= ignored (|author= suggested) (help); Unknown parameter |quotes= ignored (help)
  9. ^ a b Tait, Malcolm (2006-08-28). "1". Animal Tragic: Popular Misconceptions of Wildlife Through the Centuries. Think Books. p. 38. ISBN 184525015X. Retrieved 2008-02-19. {{cite book}}: External link in |chapterurl= (help); Unknown parameter |chapterurl= ignored (|chapter-url= suggested) (help)
  10. ^ a b c Griveaud, Paul (1963). "4". The Invertebrates of Madagascar (PDF). pp. 81–82. Retrieved 2008-02-19.
  11. ^ a b c Smith, Neal G. (1983). "Host Plant Toxicity and Migration in the Dayflying Moth Urania". The Florida Entomologist. 66 (1). Florida Entomological Society: 76–85. doi:10.2307/3494552. {{cite journal}}: Unknown parameter |laysource= ignored (help); Unknown parameter |laysummary= ignored (help); Unknown parameter |month= ignored (help); Unknown parameter |quotes= ignored (help)
  12. ^ a b c d e f g h i j k Lees, David (1991). "Foodplants of the Uraniinae (Uraniidae) and their Systematic, Evolutionary and Ecological Significance" (PDF). The Journal of the Lepidopterists' Society. 45 (4). The Lepidopterists' Society: 296–347. Retrieved 2006-10-29. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help); Unknown parameter |laysummary= ignored (help); Unknown parameter |quotes= ignored (help)
  13. ^ a b c d Oberthür, Charles. Et́udes de lépidoptérologie comparée (in French). Retrieved 2008-03-20.
  14. ^ Webber, Joseph. Dictionnaire français-malgache (PDF) (in French). Retrieved 2008-03-20.
  15. ^ a b Shoumatoff, Alex (1988-03-07). "Our Far Flung Correspondents (Madagascar)". The New Yorker. New York. p. 62. Retrieved 2008-02-16.
  16. ^ a b c d e f g h i j k Catala, R. (1940). "Variations expérimentales de Chrysiridia madagascariensis Less. (Lep. Uraniidae)". Archives du Muséum National D'Histoire Naturelle. 17: Ph.D. Thesis.
  17. ^ a b Klots, Alexander Barrett (2007-05-25) [1959]. Living Insects of the World. Doubleday. Retrieved 2008-02-19. {{cite book}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  18. ^ Kaabak, L. "Лицом к лицу с природой" (in Russian). Retrieved 2008-03-18.
  19. ^ a b Gillespie, Lynn J. (1997). "Omphalea (Euphorbiaceae) in Madagascar: A New Species and a New Combination". Novon. 7 (2). St. Louis: Missouri Botanical Garden: 127–136. doi:10.2307/3392184. {{cite journal}}: Unknown parameter |laysource= ignored (help); Unknown parameter |laysummary= ignored (help); Unknown parameter |quotes= ignored (help)
  20. ^ Kite, G.C. (1991). "Alkaloidal glycosidase inhibitors in nocturnal and diurnal uraniine moths and their respective foodplant genera, Endospermum and Omphalea". Biochemical Systematics and Ecology. 19 (6). Oxford: Pergamon Press: 441–445. doi:10.1016/0305-1978(91)90063-6. {{cite journal}}: Cite has empty unknown parameter: |laydate= (help); Unknown parameter |coauthors= ignored (|author= suggested) (help); Unknown parameter |laysource= ignored (help); Unknown parameter |laysummary= ignored (help); Unknown parameter |quotes= ignored (help)
  21. ^ Minet, J. (1986). "Ébauche d'une classification moderne de l'ordre des Lépidoptères". Alexanor. 14 (7): 291–313. {{cite journal}}: Unknown parameter |quotes= ignored (help)
  22. ^ a b c Carter, David (2000). Butterflies and Moths (Eyewitness Handbook). Dorling Kindersley Publishing. ISBN 1-56458-062-8.
  23. ^ Common, I. F. B. Moths of Australia. Brill Publishers. p. 382. ISBN 9-004-09227-7. Retrieved 2008-03-18.
  24. ^ "Chrysiridia rhipheus". Collection naturalisée (in French). Insectarium de Montréal. 2005-05-10. Retrieved 2006-10-29.
  25. ^ Mannoni, Octave (1991). Prospero and Caliban: The Psychology of Colonization. Contributor Maurice Bloch. University of Michigan Press. p. 53. ISBN 0472064304. Retrieved 2008-06-17. {{cite book}}: line feed character in |publisher= at position 23 (help)
  26. ^ Bradshaw, Joel (2007-10-03). Issues in Austronesian Morphology: A Focusschrift for Byron W. Bender. Pacific Linguistics, Research School of Pacific and Asian Studies, The Australian National University. pp. 55 and 61. ISBN 0858834855. Retrieved 2008-06-17. {{cite book}}: Unknown parameter |coauthors= ignored (|author= suggested) (help); Unknown parameter |origdate= ignored (|orig-date= suggested) (help)
  27. ^ Académie malgache (2006-04-19) [1919]. Bulletin de l'Académie malgache. 1915-1921 (in French). Vol. 2–5. Imp. officielle de la colonie. p. 30. Retrieved 2008-06-17.
  28. ^ Eliane, Itti (2003). L'Image des civilisations francophones dans les manuels scolaires (in French). Editions Publibook. p. 141. ISBN 274832482X. Retrieved 2008-06-17.