Birdwings are butterflies in the swallowtail family, that belong to the genera Trogonoptera, Troides, and Ornithoptera. Most recent authorities recognise 36 species, however, this is debated, and some authorities include additional genera. Birdwings are named for their exceptional size, angular wings, and birdlike flight. They are found across tropical Asia, mainland and archipelagic Southeast Asia, and Australasia.
Due to their size and brightly coloured males, they are popular among collectors of butterflies, but all birdwings are now listed by CITES,[1] thereby limiting (and in the case of O. alexandrae completely banning) international trade.
After mating, females immediately begin to seek appropriate host plants; climbing vines of the genera Aristolochia and Pararistolochia (both in the family Aristolochiaceae) are sought exclusively. The female lays her spherical eggs under the tips of the vine's leaves, one egg per leaf.
Larva
Early instar larva of Troides minos
The caterpillars are voracious eaters but move very little; a small group will defoliate an entire vine. If starved due to overcrowding, the caterpillars may resort to cannibalism. Fleshy spine-like tubercles line the caterpillars' backs, and their bodies are dark red to brown and velvety black. Some species have tubercles of contrasting colours, often red, or pale "saddle" markings. Like other members of their family, birdwing caterpillars possess a retractable organ behind their heads called an osmeterium. Shaped like the forked tongue of a snake, the osmeterium excretes a fetid terpene-based compound and is deployed when the caterpillar is provoked. The caterpillars are also unappealing to most predators due to their toxicity: the vines which the caterpillars feed upon contain aristolochic acid, a poisonous compound known to be carcinogenic in rats. The feeding caterpillars incorporate and concentrate the aristolochic acid into their tissues, where the poison will persist through metamorphosis and into adulthood.
Pupa
Pupa of Ornithoptera victoriae
Birdwing chrysalids are camouflaged to look like a dead leaf or twig. Before pupating, the caterpillars may wander considerable distances from their host plants. In O. alexandrae, it takes about four months to get from egg to adult. Barring predation, this species can also survive up to three months as an adult.
Imago
Birdwings inhabit rainforests and adults are usually glimpsed along the forest periphery. They feed upon—and are important long-range pollinators of—nectar-bearing flowers of the forest canopy, as well as terrestrial flowers, such as lantana. They are strong flyers and seek sunlit spots in which to bask.
Breeding behaviour varies little between species; the female's role is relatively passive, slowly fluttering from perch to perch while the male performs an elaborate, quivering yet stationary dance 20–50cm above her.
Copulating pair of Ornithoptera euphorion (female above, male below). The sexes are quite dissimilar in appearance, as is typical of species in the genus Ornithoptera.
Birdwings are typified by large size (up to a maximum body length of 7.6cm or 3inches and a wingspan of 28cm or 11inches in O. alexandrae), showy colouration (in contrasting shades of green, yellow, black, white, and sometimes blue or orange), and slender, lanceolate forewings. With few exceptions (i.e., the New GuineanO. meridionalis and O. paradisea), the hindwings lack tails. Sexual dimorphism is strong in Ornithoptera species only, where males are black combined with bright iridescent green, blue, orange, or yellow while the larger and less colourful females are overall black or dark brownish with white, pale brown, or yellow markings.
Males and females of most Troides birdwings are similar and have jet black to brown dorsal forewings, often with the veins bordered in grey to creamy white. At least one of these darkly-coloured species (T. rhadamantus) possesses thermoreceptors on the anal veins (A2 and A3) of the wings and on the antennal clubs. The antennal receptors of the clubs—which also possess hygroreceptors that measure atmospheric humidity—are known as sensilla basiconica. The thermoreceptors are sensitive to sudden increases in temperature; they are thought to help the butterfly thermoregulate and avoid overheating while basking.
The colours of most species are pigmentary (via papiliochrome); but two species, Troides magellanus and the much rarer T. prattorum, are noted for their use of limited-view iridescence: the yellow of the dorsal hindwings is modified by bright blue-green iridescence which is only seen when the butterfly is viewed at a narrow, oblique angle. This "grazing iridescence" is brought about through diffraction of light (after back-reflection) by the wings' extremely steeply-set, multilayered rib-like scales (rather than the ridge-lamellae of most other iridescent butterflies, such as Morpho species). Such limited-view iridescence was previously only known from one other species, the riodinidAncyluris meliboeus. In A. meliboeus, however, the iridescence is produced by ridge-lamellar scales and features a wider range of colours.
The close evolutionary relationship between Troides and Ornithoptera butterflies is well demonstrated by the fact that commercial breeders have produced numerous hybrids between the two.
The final and smallest genus is Trogonoptera with just two species. They resemble each other, being overall black with iridescent green markings and a red head. Females are duller than males.
O. alexandrae has a very small distribution and is the only birdwing on Appendix I of CITES
With the exception of Queen Alexandra's birdwing (O. alexandrae), all birdwings are listed in Appendix II of CITES,[1] and accordingly their trade is restricted in countries that have signed the CITES convention. Exceptions are made for captive-reared specimens, which mainly originate from ranches in Papua New Guinea and Indonesia.[6] Most species of all three genera have now been reared in captivity, though with significant differences in the quantities reared of each species.[6]O. alexandrae is listed on Appendix I[1] and therefore cannot legally be traded internationally. At the 2006 meeting of the CITES Animals Committee some suggested O. alexandrae should be moved to Appendix II, as the conservation benefits of sustainable management perhaps are higher than those of the trade ban.[7]
Three Troides and eight Ornithoptera species have been given assessments by the IUCN Red List, with classifications ranging from "least concern" to "endangered".[8]
Richmond birdwings (O. richmondia) depend on the plant Aristolochia praevenosa which they need for their caterpillars. However, the very similar Aristolochia elegans (Dutchman's pipe) which can be found in many Australian backyards, kills the caterpillars.
Reproduction
Ornithoptera, or the genus of birdwing butterflies, usually reproduce sexually and are oviparous.[9] In butterflies sex is determined by a WW/WZ system, with a heterogametic female, reverse of that found in mammals and many other insects, which have a heterogametic male.[10] During copulation males will transfer an ejaculate containing both sperm and accessory substances that can make up to fifteen percent of a males body mass.
Mating systems
Mating systems, first explored in evolutionary terms by Darwin, includes all behaviours associated with sexual reproduction.[11] Mating systems include all costs and benefits, pre- and postcopulatory competitions, displays and mate choice. Butterfly mating systems have great variation, including strict monandry, one male and one female, to polyandry, having many mates of the opposite sex. Typically Ornithoptera tend to be polygamous, mating with more than one individual.
Female choice
Female choice can have a serious impact on mate selection and successful reproduction. Several species of Ornithoptera have been known to create hybrids if they have no access to their own species.[9]Troides oblongamaculatus females have been known to choose to mate with other species such as Ornithoptera priamus poseidon, which will attempt mating if their own species is not to be found near by. The females will typically resist mating attempts by covering their abdomen with their forewings or dropping to the ground, making mating near impossible. Although the females usually resist these mating attempts, they have been noted to be more susceptible if they have not had previous encounters with males of their own species.
Male courtship
Some male Ornithoptera species demonstrate courtship behaviour. Ornithoptera priamus posedion males will approach a female carefully, and examine the female for several minutes. After consideration, the male may choose to hover twenty to thirty centimeters above the female, displaying the bright yellow marking on its hindwings. Meanwhile, the forewings will move forward, exposing the abdomen and androconial hair tufts. Mating is only attempted when the female has ceased to flap her wings.[12] After about thirty seconds of the display, the male will attempt copulation.
Cryptic choice: sperm competition and postcopulatory guarding
In many animals, females often mate with more than one male.[13] Males who are able will adapt strategies such as postcopulatory guarding to ensure the paternity of the offspring. Following insemination, it is common for the male Ornithoptera to produce a mating plug, which will seal the ostium bursae and prevent remating by the female, as new sperm is unable to enter the opening. The plug does not impede oviposition and may stay in place for the duration of the female's life.
Sexual dimorphism
Sexual dimorphism is very prominent in Ornithoptera species, the males being black with brightly colored markings of blue, green, orange or yellow and the females are overall black or dark brown.[14] The sexual dichromatism functions in mate recognition by the use of photoreceptors. Due to the protected nature of Ornithoptera it has been difficult to study the spectral sensitivities of the sexes although this difference in coloration alludes to the idea of sensory exploitation of the female's photoreceptors. The sensory bias of females to select for males with brighter wings has yet to be studied in Ornithoptera.
Gyanandromorphism is a very rare condition in which an organism simultaneously expresses both male and female phenotypes. It is only observed in species that express strong sexual dimorphism. Gynandromorphs are suspected to be due to genetic errors associated with cell division such as nondisjunction, as well as fertilization of binucleate ova and fertilisation of multiple sperm that may fuse and act as a second nucleus. Ornithoptera is known to commonly exhibit this phenomenon, but little to no research has been successful in determining why. Those who experience this phenomenon, usually females, show male-pigmented tissues on their wings.
1 2 Straatman, R. Hybridisation of Birdwing Butterflies (Lepidoptera: Papilionidae) In Papua New Guinea. The Lepidopterological Society of Japan. Tyô to Ga (Transactions of the Lepidopterological Society of Japan) Vol. 27, No, 4 1976 156-162.
↑ Nielsen, J. (2010). A Review of Synandromorphism in the Genus Ornithoptera Boisduval, (Lepidoptera: Papilionidae). Australian Entomologist, 37(3): 105-112.
↑ Emlen ST, Oring LW. (1977). Ecology, sexual selection, and the evolution of mating systems. Science 197:215-223.
↑ Orr, A. (1999). Possible Postcopulatory Mate Guarding in Ornithopter Euphorion (Gray) (Lepidoptera: Papilionidae). Australian Entomologist, 26 (3): 71-76.
↑ Satoh, Y. Yano, S. Takafuji, A. Mating strategy of spider mite, Tetranychus urticae (Acari: Tetranychidae) males: postcopulatory guarding to assure paternity. (2001). Applied Entomology, 36 (1): 41-45.
↑ Wilts, B. D., Matsushita, A., Arikawa, K., & Stavenga, D. G. (2015). Spectrally tuned structural and pigmentary coloration of birdwing butterfly wing scales. Journal of the Royal Society Interface, 12(111), 20150717. http://doi.org/10.1098/rsif.2015.0717
Other references
d'Abrera, Bernard. (1975). Birdwing Butterflies of the World Hill House Publishers ISBN0947352422
Igarashi, S. (1979). Papilionidae and their early stages. Volume I Text (in Japanese), Volume 2 Plates. Kodansha, Tokyo.
Parsons, M.J. (1996). A phylogenetic reappraisal of the birdwing genus Ornithoptera (Lepidoptera: Papilionidae: Troidini) and a new theory of its evolution in relation to Gondwanan vicariance biogeography Journal of Natural History Volume 30, Issue 11:1707-1736.
Parsons, M.J. (1996). Gondwanan evolution of the troidine swallowtails (Lepidoptera: Papilionidae): Cladistic reappraisals using mainly immature stage characters, with focus on the birdwings Ornithoptera Boisduval Bulletin of the Kitakyushu Museum of Natural History 15: 43-118, 34 figures, 2 tables pdf
Parsons, M.J. (1992). The butterfly farming and trading industry in the Indo-Australian region and its role in tropical forest conservation. Tropical Lepidoptera 3 (Suppl. 1): 1-31.pdf Full text
Haugum, Jan. (1981). Notes on the Aristolochia of the Papuan Region, with particular reference to the larval foodplants of the Ornithoptera. Lep. Group Newsl. 2(10), pp.171–178
Haugum, Jan; & Low, A. M. (1978). A Monograph of the Birdwing Butterflies. Volume 1, Part 1. Introduction, Ornithoptera (Aetheoptera)., Klampenborg, Denmark, Scandinavian Science Press 1(1)
Haugum, Jan; & Low, A. M. (1979). A Monograph of the Birdwing Butterflies. Volume 1, Part 2. Ornithoptera (Ornithoptera)., Klampenborg, Denmark, Scandinavian Science Press 1(2)
Haugum, Jan; & Low, A. M. (1980). A Monograph of the Birdwing Butterflies. Volume 1, Part 3. Ornithoptera (Schoenbergia)., Klampenborg, Denmark, Scandinavian Science Press 1(3)
Haugum, Jan; & Low, A. M. (1981). A Monograph of the Birdwing Butterflies. Volume 2, Part 1. Trogonoptera & Ripponia., Klampenborg, Denmark, Scandinavian Science Press 2(1)
Haugum, Jan; & Low, A. M. (1982). A Monograph of the Birdwing Butterflies. Volume 2, Part 2. Troides; amphrysus & haliphron groups., Klampenborg, Denmark, Scandinavian Science Press 2(2)
Haugum, Jan; & Low, A. M. (1983). A Monograph of the Birdwing Butterflies. Volume 2, Part 3. Troides; helena and aeacus groups., Klampenborg, Denmark, Scandinavian Science Press 2(3)
Kiyotaro Kondo, Tsutomu Shinkawa & Hirotaka Matsuka. (2003). Molecular systematics of birdwing butterflies (Papilionidae) inferred from mitochondrial ND5 gene Journal of the Lepidopterists' Society 57:17-24 pdf[permanent dead link]
Robert Henry Fernando Rippon (1898 to 1906) Icones Ornithopterorum [London] Published by the author at Upper Norwood, London, S.E.
Schäffler, Oliver . (2001). Schmetterlinge der Erde, Butterflies of the World Part XII (12), Papilionidae VI: Ornithoptera Edited by Erich Bauer and Thomas Frankenbach Keltern: Goecke & Evers; Canterbury: Hillside Books. ISBN9783931374839 Supplement to von Knötgen, 1997
von Knötgen, Béla. (1997). Ornithoptera: Ornithoptera Schönbergia, Aetheoptera Wangen (Allemagne): MGG Verlag, 1997. Parallel text in German, English and French.
Darby, A.W. (1982). "The female genitalia of the Birdwing Butterflies, part 1 Lepidoptera Group 68. Vejle. 1982. Showing female genitalia of T. helena cerberus, O. priamus richmondia, O. priamus arruana, T. brookiana albescens.
Darby, A.W. (1983). "The female genitalia of the Birdwing Butterflies, part 2. Lepidoptera Group 68. Vejle. 1983. Showing female genitalia of O. goliath procus, T. amphrysus ruficollis, T. a. flavicollis, T. miranda miranda, T. m. neomiranda, T. cuneifera paeninsulae, T. helena cerberus, T. h. hephaestus, T. oblongomaculatus oblongomaculatus, T. o. bouruensis, T. o. papuensis, T. aeacus aeacus, T. a. thomsonii, T. aeacus formosanus, T. rhadamantus rhadamantus, T. r. dohertyi, T. r. plateni, T. vandepolli vandepolli, T. v. honrathiana, T. criton, T. darsius, T. haliphron haliphron, T. h. socrates, T. h. iris, T. h. naias, T. h. pallens, T. prattorum, T. magellanus sonani, T. hypolitus hypolitus.
Ornithoptera alexandrae, the Queen Alexandra's birdwing, is the largest species of butterfly in the world, with females reaching wingspans slightly in excess of 25 cm to 28 cm. This birdwing is restricted to the forests of the Oro Province in eastern Papua New Guinea.
Ornithoptera goliath, the Goliath birdwing, is a birdwing butterfly found in New Guinea. It is the second largest butterfly in the world, after the Queen Alexandra's birdwing.
Trogonoptera brookiana, Rajah Brooke's birdwing, is a birdwing butterfly from the rainforests of the Thai-Malay Peninsula, Borneo, Natuna, Sumatra, and various small islands west of Sumatra. The butterfly was named by the naturalist Alfred Russel Wallace in 1855, after James Brooke, the Rajah of Sarawak. The larval host plants are Aristolochia acuminata and A. foveolata. Adults sip flower nectar from plants such as Bauhinia. Rajah Brooke's birdwing is a protected species, listed under Appendix II of CITES, meaning that international export is restricted to those who have been granted a permit. It is the national butterfly of Malaysia.
Rothschild's birdwing is a large birdwing butterfly, endemic to the Arfak Mountains in Western New Guinea.
Ornithoptera chimaera, the chimaera birdwing, is a birdwing butterfly of the family Papilionidae. It is found in mountain areas of New Guinea, 1000 meters above sea level.
Ornithoptera tithonus, the Tithonus birdwing, is a species of birdwing butterfly found on New Guinea and other neighbouring islands.
Ornithoptera paradisea, the paradise birdwing, is a species of birdwing butterfly found in New Guinea.
Troides vandepolli, the van de Poll's birdwing, is a montane birdwing butterfly occurring on Java and Sumatra. It is endemic for Indonesia and is protected by the Convention on International Trade in Endangered Species (CITES).
Troides plato, the silver birdwing, is a birdwing butterfly endemic to Timor.
Troides magellanus, the Magellan birdwing, is a large and striking species of birdwing butterfly found in the Philippines and on Taiwan's Orchid Island.
Troides dohertyi, the Talaud black birdwing, is a birdwing butterfly endemic to the Talaud and Sangihe islands.
Troides haliphron, the haliphron birdwing, is a birdwing butterfly confined to Sulawesi and the lesser Sunda Islands.
Ornithoptera priamus, the common green birdwing, Cape York birdwing, Priam's birdwing, northern birdwing or New Guinea birdwing is a widespread species of birdwing butterfly found in the central and south Moluccas, New Guinea, Bismarck Archipelago, Solomon Islands, and north-east Australia.
Ornithoptera victoriae, the Queen Victoria's birdwing, is a birdwing butterfly of the family Papilionidae, found in the Solomon Islands and Papua New Guinea.
Ornithoptera richmondia, the Richmond birdwing, is a species of birdwing butterfly that is endemic to Australia. It is the second smallest of the birdwing species, the smallest being Ornithoptera meridionalis.
Troides darsius, the Sri Lankan birdwing, is a species of birdwing butterfly found in Sri Lanka. It is the largest butterfly on the island and is also the national butterfly of Sri Lanka.
Ornithoptera aesacus, the Obi Island birdwing, is an extremely rare species of birdwing butterfly, endemic to the Island of Obira, Indonesia. While being a very rare species, they are commercially bred and traded.
Troides andromache, the Borneo birdwing or Kinabalu Birdwing, is a species of butterfly in the family Papilionidae. It is found only in Borneo. In 2023, it was selected as the official state butterfly of the state of Sabah, Malaysian Borneo.
Ornithoptera allotei is the name given to a birdwing butterfly that is a natural hybrid between Ornithoptera victoriae and Ornithoptera priamus urvillianus. Despite the fact that hybrids do not warrant a binomial name, the name Ornithoptera allotei persists from the original description of the butterfly as a species.
Troides cuneifera, the Mountain Birdwing, is a large butterfly belonging to the swallowtail family, Papilionidae, found in the Thai-Malay Peninsula, Sumatra, Java and Borneo.
This page is based on this Wikipedia article Text is available under the CC BY-SA 4.0 license; additional terms may apply. Images, videos and audio are available under their respective licenses.
