Birdwing

Birdwing
One male and female representative of the genera; Trogonoptera, Troides and Ornithoptera. Species from top to bottom, left to right: Trogonoptera brookiana (♂), Trogonoptera brookiana (♀), Troides dohertyi (♂), Troides dohertyi (♀), Ornithoptera aesacus (♂), Ornithoptera goliath (♀).
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Lepidoptera
Family:	Papilionidae
Subfamily:	Papilioninae
Tribe:	Troidini
Genus:	Trogonoptera Troides Ornithoptera
Genera & Species
3 Genera, 36 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,^[1] thereby limiting (and in the case of O. alexandrae completely banning) international trade.

Troides darsius, popularly known as the Sri Lanka Birdwing, is the National Butterfly of Sri Lanka.

Adult physical description

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.

The male (above, dorsal surface) and female (below, ventral surface) Ornithoptera goliath are completely different, as is typical of species in the genus Ornithoptera.

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. rhadamathus) 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 lycaenid 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 hybrid specimens 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.

Life history

A mating pair of Ornithoptera euphorion (female above, male below).

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. 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. Some species have tubercles of contrasting colours, 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.

Status and protection

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.^[2] Most species of all three genera have now been reared in captivity, though with significant differences in the quantities reared of each species.^[2] 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.^[3]

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.

Taxonomy and systematics

Genus: Trogonoptera

• Trogonoptera brookiana
• Trogonoptera trojana

Genus: Troides

Troides helena (left) and a male Trogonoptera brookiana (right) in a Kuala Lumpur butterfly conservatory. Each represent a typical species from their genus.

subgenus: Ripponia

• Troides hypolitus

subgenus: Troides

• Troides aeacus
• Troides amphrysus
• Troides andromache
• Troides criton
• Troides cuneifera
• Troides darsius
• Troides dohertyi
• Troides haliphron
• Troides helena
• Troides magellanus
• Troides minos
• Troides miranda
• Troides oblongomaculatus
• Troides plateni
• Troides plato
• Troides prattorum - Commercially bred, but supplies of this butterfly are sporadic, so it is still very rare in collections.
• Troides rhadamantus
• Troides riedeli
• Troides staudingeri
• Troides vandepolli

Genus: Ornithoptera

Males of the green O. p. poseidon and the blue O. p. urvillianus, showing some of the distinct variations in this species. The latter has sometimes been regarded as a separate species, but hybrids between these subspecies are fully viable.^[4]

subgenus: Aetheoptera

• Ornithoptera victoriae

subgenus: Ornithoptera

• Ornithoptera aesacus - Twenty years ago this was (by far) the world's rarest birdwing species. Now commercially bred.
• Ornithoptera croesus
• Ornithoptera priamus - Occurs as a number of subspecies, some of which are often regarded as full species e.g. O. (priamus) urvilleanus.

subgenus: Schoenbergia

• Ornithoptera chimaera
• Ornithoptera goliath - A mosaic gynandromorphic specimen of this species sold by a Taiwanese dealer for US$28,000 in July 2006, which is possibly sets the world record for the highest price paid for a butterfly.
• Ornithoptera meridionalis
• Ornithoptera paradisea
• Ornithoptera rothschildi
• Ornithoptera tithonus

subgenus: Straatmana

• Ornithoptera alexandrae- The world's largest butterfly. Listed on CITES Appendix I.

subgenus unassigned:

• Ornithoptera euphorion - There is a spectacular and rare genetic mutation of this butterfly (less than 40 known examples - all from a single aberrant female) where the males are gold instead of green [1].
• Ornithoptera richmondia - The smallest Ornithoptera species. Occasionally (and wrongly^{[citation needed]}) regarded as a subspecies of O. priamus.

Hybrids

There are two Ornithoptera species, now regarded as hybrids:

• O. rothschildi x O. priamus poseidon - Originally described as species Ornithoptera akakeae. Known from a single female specimen.
• O. victoriae x O. priamus urvilleanus - Originally described as species Ornithoptera allotei. This butterfly is, because of its rarity, one of the World's most valuable, with male specimens typically selling for more than £4,000.00 (US$7,000.00). It would be an ideal candidate for commercial exploitation because its parents are not rare on Bougainville Island and can (apparently) be easily induced to mate with one another.^[5]

Cited references

1. ^ ^{a b c} CITES (2011). Appendices I, II and III. Version 27 April 2011.
2. ^ ^{a b} United Nations Environment Programme World Conservation Monitoring Centre (2007). Review of trade in ranched birdwing butterflies. European Commission, 2008.
3. ^ CITES (2006). Earth Negotiations Bulletin. Summary of the 22nd Meeting of the CITES Animals Committee.
4. ^ Nagypal, T. (2000-2008). Ornithoptera priamus. Version July 29, 2010.
5. ^ Nagypal, T. The World of Birdwing Butterflies

Other references

• 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.
• Monteith, G. (2000). Queensland Museum: Birdwing butterflies. Retrieved June 28, 2005.
• 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, J. & Low, A.M. (1982). A Monograph of the Birdwing Butterflies. The systematics of Ornithoptera, Troides and related genera. Vol. 2, Part 1. Trogonoptera and Ripponia. Klampenborg, Denmark : Scandinavian Science Press pp. 9–104
• Haugum, J. & Low, A.M. (1983). [1978-1983] A Monograph of the Birdwing Butterflies. The systematics of Ornithoptera, Troides and related genera. Vols 1 & 2. Klampenborg : Scandinavian Science Press 356 pp.
• Haugum, J. & Low, A.M. (1983). A Monograph of the Birdwing Butterflies. The systematics of Ornithoptera, Troides and related genera. Vol. 2, Part 2. Troides; amphrysus and haliphron groups. Klampenborg, Denmark : Scandinavian Science Press pp. 105–204
• Haugum, J. & Low, A.M. (1985). A Monograph of the Birdwing Butterflies. The systematics of Ornithoptera, Troides and related genera. Vol. 2, Part 3. Troides; helena and aeacus groups. Klampenborg, Denmark : Scandinavian Science Press pp. [1-8], 241-356

External links

• Butterflycorner.net (English/German)
• Genus Troides at Lepidoptera.pro
• Birdwings on posatge stamps
• Pteron Birdwing Gallery. In Japanese but with binomial names.