||This article includes a list of references, but its sources remain unclear because it has insufficient inline citations. (December 2012)|
|Male (left) and female (right) representative of each birdwing genus: Trogonoptera (top), Troides (middle), and Ornithoptera (bottom)|
|Genera & species|
Birdwings are papilionid butterflies native to the Indian Subcontinent, mainland and archipelagic Southeast Asia and Australasia, and are usually regarded as belonging to three genera: Ornithoptera, Trogonoptera, and Troides. Some authorities include additional genera. The exact number of species is debated, but most recent authorities recognize between 30 and 40. Birdwings are named for their exceptional size, angular wings, and birdlike flight.
Included among the birdwings are some of the largest butterflies in the world: the largest, Queen Alexandra's Birdwing (Ornithoptera alexandrae); the second largest, the Goliath Birdwing (O. goliath); and the largest butterfly endemic to Australia, the Cairns Birdwing (O. euphorion). Another well-known species is Rajah Brooke's Birdwing (Trogonoptera brookiana), a particularly attractive species named after Sir James Brooke, the first White Rajah of 19th century Sarawak.
Due to their size and bright colours, they are popular among collectors of butterflies, but all birdwings are now listed by CITES, thereby limiting (and in the case of O. alexandrae completely banning) international trade.
Taxonomy and systematics
- Ornithoptera rothschildi x Ornithoptera priamus poseidon — Ornithoptera akakeae
- Ornithoptera victoriae x Ornithoptera priamus urvillianus — Ornithoptera allotei
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.
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.
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 ca. four months to get from egg to adult. Barring predation, this species can also survive up to three months as an adult.
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 20–50 cm above her.
Birdwings are typified by large size (up to a maximum body length of 7.6 cm or 3 inches and a wingspan of 28 cm or 11 inches in O. alexandrae), showy coloration (in contrasting shades of green, yellow, black, white, and sometimes blue or orange), and slender, lanceolate forewings. With few exceptions (i.e., the New Guinean O. 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 riodinid Ancyluris 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.
Ornithoptera may be found in Indonesia (east of Weber's line: the Moluccas and West Papua), Papua New Guinea (including Bougainville Island), the Solomon Islands, and Australia (north Queensland and at far east Australia).
Status and protection
With the exception of Queen Alexandra's Birdwing (O. alexandrae), all birdwings are listed in Appendix II of CITES, 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. Most species of all three genera have now been reared in captivity, though with significant differences in the quantities reared of each species. O. alexandrae is listed on Appendix I 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.
Richmond Birdwing (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.
- CITES (2011). Appendices I, II and III. Version 27 April 2011.
- The World of Birdwing Butterflies. Trogonoptera brookiana. Accessed 20 May 2012
- ARKive Rajah Brooke’s birdwing. Accessed 20 May 2012
- Collins, N. M. & M. G. Morris. (1985) Threatened Swallowtail Butterflies of the World. IUCN. ISBN 2-88032-603-6
- United Nations Environment Programme World Conservation Monitoring Centre (2007). Review of trade in ranched birdwing butterflies. European Commission, 2008.
- CITES (2006). Earth Negotiations Bulletin. Summary of the 22nd Meeting of the CITES Animals Committee.
- IUCN redlist, entries for Troides and Ornithoptera.
- d'Abrera, Bernard. (1975). Birdwing Butterflies of the World Hill House Publishers ISBN 0947352422
- American Museum of Natural History. BioBulletin: Birdwing butterflies Retrieved June 28, 2005 from
- Campbell, A.L., Naik, R.R., Sowards, L., and Stone, M.O. (2002). Biological infrared imaging and sensing. Micron 33, 211–225.
- Collins, N.M., Morris, M.G., IUCN, (1985). Threatened Swallowtail Butterflies of the World: the IUCN Red Data Book (1985) IUCN pdf
- 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
- Reed, R.D., and Sperling, F.A.H. (2001). Tree of Life: Papilionidae Retrieved June 28, 2005
- Savela, M. (2005). Troides. Retrieved June 28, 2005 from
- Vukusic, P., Sambles, J. R., and Ghiradella, H. (2000). Optical classification of microstructure in butterfly wing-scales. Photonics Science News, 6, 66–66.
- Nagypal, Tony. The World of Birdwing Butterflies.
- 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
- 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. ISBN 9783931374839 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.
- Images representing Ornithoptera, at EoL
- Butterflycorner.net (English/German)
- Genus Troides at Lepidoptera.pro
- Birdwings on posatge stamps
- Pteron Birdwing Gallery. In Japanese but with binomial names.
Wikimedia sister projects
- Wikimedia Commons has media related to Ornithoptera and Troides
- Wikispecies has information related to Ornithoptera, Trogonoptera, and Troides