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Molecular Biology and Genetics
Barcode data: Papilio demoleus
Below is a sequence of the barcode region Cytochrome oxidase subunit 1 (COI or COX1) from a member of the species.
See the BOLD taxonomy browser for more complete information about this specimen and other sequences.
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Statistics of barcoding coverage: Papilio demoleus
Public Records: 15
Specimens with Barcodes: 20
Species With Barcodes: 1
Papilio demoleus is a common and widespread swallowtail butterfly. The butterfly is also known as the common lime butterfly, lemon butterfly, lime swallowtail, small citrus butterfly, chequered swallowtail, dingy swallowtail and citrus 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. The butterfly has also been referred to as the Butterfly of Death, a name it shares with a morphologically similar species, Papilio demodocus, which flies in Africa. The butterfly is a pest and invasive species from the Old World which has spread to the Caribbean and Central America.
The butterfly is tailless and has a wingspan 80–100 mm.:52 Above, the background colour is black. A broad, irregular yellow band is found on the wings above, which is broken in the case of the forewing. Besides this the butterfly has a large number of irregular spots on the wing. The upper hindwing has a red tornal spot with blue edging around it.
As the butterfly caterpillar ages, its hunger for leaf tissue continues to grow.
Detailed description as given in Bingham (1905):
Upper side of wings has the ground colour black. The fore wing has the base below cell and basal half of latter so irrorated with yellow scales as to form more or less complete transverse dotted lines, two outwardly oblique yellow spots in cell and a curved spot at its upper apex; a spot at base and another beyond it in interspace 8; a discal transverse series of cream-yellow spots irregular in arrangement and size extends from interspace la to 8; the series interrupted in interspace 5 and the spot in interspace 7 double; this is followed by a sinuous postdiscal series of spots and an admarginal terminal series of smaller spots. In many specimens between the discal and postdiscal series the black ground-colour is irrorated with yellowish scales. Hind wing: base and an edging that decreases in width along the dorsal margin irrorated with yellow scales; followed by a broad medial yellow irregular band, a sinuous postdiscal series of outwardly emarginate yellow spots and a terminal series of smaller similarly coloured spots as on the fore wing. The inner margin of the medial band is curved inwards, the outer margin is very irregular and uneven; in the cell the band does not reach the apex, but beyond the cell there are one or more cream-yellow spots, and the black groundcolour is irrorated with yellowish scales; finally at the tornal angle there is an oval ochraceous-red spot emarginate on its inner side in the female and in both sexes surmounted by a blue lunule; while in interspace 7 between the medial band and the postdiscal spot there is a large ocellus-like spot of the black ground-colour more or less irrorated with blue scales.
The underside has the ground-colour similar, the cream-coloured markings paler and conspicuously larger. The markings differ from those on the upperside in that the forewing has the basal half of cell and base of wing below it with cream-coloured streaks that coalesce at base; irregular ochraceous spots in interspaces 5 to 8 and the discal series of spots complete not interrupted in interspace 5. On the underside of the hind wing, the black at base of wing and along the dorsal margin centred largely with pale cream-colour; the ocellus in interspace 7, the apex of the cell and the black groundcolour between the medial band and postdiscal markings in interspaces 2-6 centred with ochraceous, margined with blue.
Antennae dark reddish brown, touched with ochraceous on the innerside towards the club; head, thorax and abdomen dusky black, the head and thorax anteriorly streaked with cream-vellow: beneath: the palpi, thorax and abdomen cream-yellow with lateral longitudinal black lines on the last.
Status, range and habitat
Papilio demoleus is an aggressive and very common butterfly. It is perhaps the most widely distributed swallowtail in the world. The butterfly can be found in:
Oman, UAE, Saudi Arabia, Kuwait, Bahrain, Qatar, Iran, western and possibly eastern Afghanistan, and western Pakistan, Sri Lanka, India (including the Andamans), Nepal, Burma, Thailand, the Philippines, Kampuchea, southern China (including Hainan, Guangdong province), Taiwan, Japan (rare strays), Malaysia, Singapore, Indonesia (Kalimantan, Sumatra, Sula, Talaud, Flores, Alor and Sumba), Papua New Guinea, Australia (including Lord Howe's island), apparently Hawaii and possibly other Pacific Ocean islands.
In recent years the butterfly has spread to Hispaniola island (Dominican Republic) in the Western Hemisphere, and subsequently to Jamaica, and Puerto Rico. The Dominican population originated from Southeast Asia but how the butterfly reached there is not known.
The widespread range of Papilio demoleus indicates the butterfly's tolerance and adaptation to diverse habitats. It is to be found in savannahs, fallow lands, gardens, evergreen and semi-evergreen forests and shows a preference for stream and riverbeds. In India it is mostly found in the plains but can be found on the hills of peninsular India and up to 7000 feet in the Himalayas. It is common in urban gardens and may also be encountered in wooded country. The butterfly is also a very successful invader, its spread appearing to be due to its strong flight, increase in urbanisation and agricultural land use that opens up new areas for dispersal, and greater availability of foodplants.
Five related butterflies form the group of lime butterflies in the genus Papilio of which Papilio demoleus Linnaeus, 1758 is the flagship species which gives the name to the group. The other morphologically related butterflies are:
- Papilio demodocus (Esper, 1798)
- Papilio erithonioides Grose-Smith, 1891
- Papilio grosesmithi Rothschild, 1926
- Papilio morondavana Grose-Smith, 1891
Research into the biogeography, phylogeny, and analysis of vicariance relationships dating back to the Cretaceous, of the "lime butterfly" or "demoleus" group, suggest that the group of lime swallowtails diversified in Madagascar in the middle Miocene.
- P. d. demoleus Linnaeus, 1758 — Across Asia from China to the Arabian peninsula, and now in some islands in the West Indies, including Jamaica
- P. d. libanius Fruhstorfer, 1908 — Taiwan, Philippines, Sula, Talaud.
- P. d. malayanus Wallace, 1865 — Sumatra and the Malaysian peninsula.
- P. d. novoguineensis Rothschild, 1908 — Papua New Guinea.
- P. d. sthenelus Macleay, 1826 — Australia.
- P. d. stenelinus Rothschild, 1895 — Sumba, Flores and Alor.
This butterfly is an avid mud-puddler and visitor of flowers. It basks with its wings held wide open on tufts of grass, herbs and generally keeps within a metre above the ground, even on cloudy days. It relies on its quick flight for escape. It is an interesting butterfly in that it has a number of modes of flight. In the cool of the morning, the flight is slow considering that it is an edible and unprotected swallowtail. As the day progresses, it flies fast, straight and low. In the hotter part of the day, it may be found settling on damp patches where it will remain motionless, except for an occasional flutter of wings, if not disturbed.
It is also a frequent visitor of flowers in gardens, where it shows a preference for flowers of smaller herbs rather than larger plants such as the ubiquitous Lantana with its plentiful blooms. It can be found swarming in the groves of its foodplants.
Research on freshly emerged imagines of Papilio demoleus showed that they have an inborn or spontaneous preference while feeding for blue and purple colours while the yellow, yellowish-green, green and blue-green colours are completely neglected.
While resting, the butterfly closes its wing over its back and draws the forewings between the hindwings.
Lime butterflies mating in Narsapur, Medak district, India.
The number of generations of Papilio demoleus is dependent upon temperature - near the equator, nine generations have been recorded, while in warm temperate China, five generations have been recorded. In the ideal conditions of a laboratory, a generation has been recorded to take place in just over 30 days. The average time for one generation of Papilio demoleus to mature in the field ranges from 26 to 59 days. In cold climates, the lime butterfly is known to pass the winter as pupae. Typically, the butterfly undergoes five instars as a caterpillar.
The female butterfly goes from plant to plant, laying a single egg at a time on top of a leaf which it holds onto with its legs, and flies off as soon as the egg is laid. The egg is round, light yellowish in colour, flattened at the base, smooth-surfaced and about 1.5 mm in height. Fertile eggs develop a small red mark at the apex.
The newly hatched caterpillar stays in the middle of the upperside of the leaf. The first instar of the caterpillar is black, with a black heads and two rows of sub-dorsal fleshy spines. The second, third and fourth instars are dark, with glossy, dark-brown head, and white markings on the 8th and 9th segments of the caterpillar which resemble a white patch of uric acid deposited in a bird's droppings, helping them escape predation while remaining in moderately open places.
As the instars progress, this resemblance is lost. From the fifth instar onwards, the caterpillars now turn cylindrical in shape, taperered towards the rear, and uniformly pale green in colour with a white sub-spiracular band. An additional black band is developed on the 4th and 5th segments with two black and two bluish spots on them. The 8th and 9th segments, which earlier provided the camouflage markings now develop a brown and white band. At this stage, the caterpillars are forced to inhabit secluded places.
The pupa, which is rugose, stout and 30 mm in length, has two projections to the front on its head and also one on its thorax and resembles that of the common Mormon Papilio polytes, the difference being that the common Mormon pupa has a deeper cut between the projections and its abdomen is more protruded on the sides, having a small point.
The pupa is dimorphic with regards to colour, with the colour developing according to the prevalent colour and texture in the background. The green morph, which is found amongst green vegetation and smoother textures, is light green and unmarked or with yellow dorsal markings. When situated among brown or dry objects, the pupa tend to turn light grey-brown to pink-brown and develop cryptic dark brown and black striation.
Captive breeding of Papilio demoleus in Riyadh have revealed the following data about the life-span of various stages at that locality:
- No of generations per year: Eight.
- Duration of egg stage: 3.1 to 6.1 days.
- Duration of larva stage: 12.9 and 22.7 days.
- Duration of pupa stage: 8.0 to 22.4 days.
- Duration of adult stage: 4 to 6 days with average of 5.1 days.
Parasitism and predation
Despite their two stage camouflage scheme, some caterpillars of Papilio demoleus are found by parasitic wasps which lay dozens of eggs in them. The parasitic wasp larva eat the caterpillar from the inside. Initially the vital organs are avoided, but by the time the caterpillar is ready to pupate even the vital organs are consumed. Shortly before, or soon after the caterpillar pupates, the parasitoids emerge from their host thus killing it.
In Saudi Arabia, the highest mortality rate was found to be in larvae and pupae in cultivated populations due to a bacterium of the genus Bacillus. In addition, eggs and larvae were heavily predated upon by two unidentified species of spiders which were abundant on citrus trees.
In India, the following braconid wasp parasitoids are known to parasitize Papilio demoleus larvae:
In Thailand, a number of organisms have been recorded attacking immature stages of Papilio demoleus:
- Egg parasites -
- Larval stage -
- Pupal parasites -
In Jamaica, an encyrtid egg parasitoid and a chalcidoid parasitoid have been reported.
- Cultivated lime, orange and lemon.
- Atalanta racemosa
- Glycosmis pentaphylla, Glycosmis arborea
- Ruta graveolens
- Bael (Aegle marmelos)
- Murraya koenigii
- Chloroxylon swietenia
- Ber (Ziziphus mauritiana)
- Acronychia pedunculata
- Microcitrus australis (Australian round-lime, Australian lime).
They have been observed on:
- Many species of Cullen : Cullen australasicum, C. badocanum, C. balsamicum, C. cinereum, C. patens (spreading scurf-pea, native verbine), C. pustulatum and C. tenax (tough scurf-pea, emu-foot, emu grass), and C. leucanthum.
- Psoralea pinnata (fountain bush).
The lime butterfly is an economic pest on many cultivated citrus species in India, Pakistan, Iraq and the Middle East. Due to its history of successful dispersal and range extension, the lime butterfly is likely to spread from its original point of introduction in Hispaniola in the Caribbean to neighbouring Florida, Central America and South America. Due to its capability for rapid population growth under favourable circumstances and its having been recorded to have five generations in a year in temperate regions of China, it is considered a serious potential threat. The caterpillars can completely defoliate young citrus trees (below 2 feet) and devastate citrus nurseries. In mature trees, caterpillars may prefer young leaves and leaf flush.
Hand-picking of caterpillars and spraying with Endosulfan 35 EC (2 ml/10 litres of water) were the recommended means of pest control by Indian government agencies and agricultural colleges, however, Endosulfan has since been banned by the Supreme Court of India.
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- Bingham, C.T. (1905). The Fauna of British India including Ceylon and Burma - Butterflies (Vol 1). London: Taylor and Francis. p. 519. Retrieved 7 November 2010.
- Collins, N. Mark; Collins, Michael G. (1985). Threatened Swallowtails of the World:the IUCN red data book. IUCN Protected Area Programme Series. Gland, Switzerland and Cambridge, U.K.: IUCN. pp. 401 & 8 plates. ISBN 978-2-88032-603-6. Retrieved 22 October 2010.
- Guerrero, Kelvin A.; Veloz, Denia; Boyce, Sarah Lyn & Farrell, Brian D. (2004). "First New World Documentation of an Old World Citrus Pest, the Lime Swallowtail Papilio demoleus (Lepidoptera: Papilionidae), in the Dominican Republic (Hispaniola)". American Entomologist (Entomological Society of America) 50 (4): 227–229. Retrieved 15 November 2010.
- Lewis, Delano S. (January 2009). "Lime Swallowtail, Chequered Swallowtail, Citrus Swallowtail Papilio demoleus Linnaeus (Insecta: Lipidoptera: Papilionidae)". University of Florida (IFAS Extension). Retrieved 26 November 2010.
- Homziak, Nicholas T.; Homziak, Jurij (2006). "Papilio demoleus (Lepidoptera: Papilionidae): A new record for the United States, Commonwealth of Puerto Rico" (full free download). Florida Entomologist 89 (4): 485–488. doi:10.1653/0015-4040(2006)89[485:PDLPAN]2.0.CO;2. Retrieved 11 November 2010.
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- Kunte, Krushnamegh (2000). Butterflies of Peninsular India. India, a lifescape (reprint 2006 ed.). Hyderabad: Universities Press (India) Ltd. p. 254. ISBN 978-81-7371-354-5. Retrieved 27 November 2010.
- Heppner John B. (8 December 2006). "Pest Alert: Lime Swallowtail in the Caribbean and possible impacts for Florida citrus". Florida Department of Agriculture and Consumer Services Division of Plant Industries. Retrieved 27 November 2010.
- Zakharov, E. V.; Smith, C. R., Lees, D. C., Cameron, A., Vane-Wright, R. I. and Sperling, F. A. H. (2004). "Independent gene phylogenies and morphology demonstrate a Malagasy origin for a wide-ranging group of swallowtail butterflies" (full free download). Evolution 58: 2763–2782. doi:10.1111/j.0014-3820.2004.tb01628.x. Retrieved 15 November 2010.
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- Grund, R. (9 December 1999). "Papilio demoleus sthenelus W.S. Macleay (Chequered Swallowtail)". South Australian Butterflies. Retrieved 23 August 2010.
- Smith, A. G. (1978). "Environmental Factors Influencing Pupal Colour Determination in Lepidoptera. I. Experiments with Papilio polytes, Papilio demoleus and Papilio polyxenes". Proceedings of the Royal Society of London. Series B, Biological Sciences (abstract) 200 (1140): 295–329. doi:10.1098/rspb.1978.0021. Retrieved 29 November 2010.
- Badawi, Ali (1981). "Studies on some aspects of the biology and ecology of the citrus butterfly Papilio demoleus L. in Saudi Arabia (Papilionidae, Lepidoptera)". Zeitschrift für Angewandte Entomologie (Abstract) (Blackwell Publishing Ltd) 91 (1-5): 286–292. doi:10.1111/j.1439-0418.1981.tb04481.x.
- Wynter-Blyth, M.A. (1957) pg 500.
- Unattributed (2008). "Acid Lime (Citrus aurantifolia (Christm) Swingle)". TNAU Agritech Portal - Horticulture:Fruit Crops:Acid Lime. Tamil Nadu Agricultural University. Retrieved 20 March 2012. Alt webcitation url.
- "Supreme Court bans Endosulfan for eight weeks - Economic Times". Articles.economictimes.indiatimes.com. 13 May 2011. Retrieved 20 March 2012.. Alt webcitation url.
- "Supreme Court refuses to lift ban on endosulfan - Times Of India". Articles.timesofindia.indiatimes.com. 5 August 2011. Retrieved 20 March 2012. Alt webcitation url.
- Chattopadhyay, Jagannath. (2007),"Swallowtail Butterflies, Biology and Ecology of a few Indian Species." Desh Prakashan, Kolkata, West Bengal, India. ISBN 978-81-905719-1-3.
- F. Martin Brown and Bernard Heineman, Jamaica and its Butterflies (E. W. Classey, London 1972), plate VIII
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