Papilio demoleus is a common and widespread swallowtail butterfly. The butterfly is also known as the lime butterfly,[1][2]lemon butterfly, lime swallowtail, and chequered swallowtail.[2] 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. When the adult stage is taken into consideration, the lime swallowtail is the shortest-lived butterfly, with male adults dying after four days and females after a week.[3] The butterfly is native to Asia and Australia, and can be considered an invasive pest in other parts of the world.[1][2] The butterfly has spread to Hispaniola island (Dominican Republic) in the Western Hemisphere,[4] and to Mahé, Seychelles.[5]
The butterfly is tailless and has a wingspan 80–100mm.[6]: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 caterpillar ages, its hunger for leaf tissue continues to grow.[7]
"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."[8]
"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.[8]
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."[8][9]
Collecting nectar from chinarose at ABS Academy Campus, Durgapur, West Bengal, India
The Southeast Asian subspecies Papilio demoleus malayanus recently established an abundant non-native population on Mahé in Seychelles[5] This species was probably accidentally introduced to Mahé a few years ago (first records in November 2016). Further dispersal events of Papilio demoleus within Seychelles to other granitic islands of the archipelago, e.g. Praslin and La Digue, are expected.[5]
Formerly absent from Borneo, it is now one of the commonest papilionids in Sabah and Sarawak in Malaysian Borneo, Kalimantan (Indonesian Borneo), and Brunei.[5][11]
In recent years, the butterfly has spread to Hispaniola island (Dominican Republic) in the Western Hemisphere,[4] and subsequently to Jamaica,[12] and Puerto Rico.[13] The Dominican population originated from Southeast Asia but how the butterfly reached there is not known.[14]
The widespread range of P. demoleus indicates the butterfly's tolerance and adaptation to diverse habitats. It is found in savannahs, fallow lands, gardens, evergreen and semi-evergreen forests, and shows a preference for streams and riverbeds.[15] In India, it is mostly found in the plains, but can be found on the hills of peninsular India and up to 7,000 feet (2,100m) in the Himalayas. It is common in urban gardens and may also be encountered in wooded country.[7] 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 food plants.[12][16]
Taxonomy
Five related butterflies form the group of lime butterflies in the genus Papilio of which P. demoleusLinnaeus, 1758 is the flagship species, which gives the name to the group. The other morphologically related butterflies are:[17]
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.[17]
Six subspecies are recognised in P. demoleus:[12][17]
P. d. libaniusFruhstorfer, 1908 — Taiwan, Philippines, Sula, Talaud
P. d. malayanusWallace, 1865 — Sumatra and the Malaysian peninsula, and now expanded its range to Southern Europe (one record in Portugal), Greater Sunda Islands (Sumatra, Java, Borneo, and Bali), Wallacea (Sulawesi, Sumbawa, Lombok, Timor, Leti, Flores, Wetar, and Alor), New Guinea, Bismarck Archipelago, Christmas Island, Torres Strait Islands (Dauan Island), Greater Antilles Islands (Hispaniola, Jamaica, Puerto Rico, Cuba, and Nueva Gerona), and Seychelles (Mahé)[5]
This butterfly is an avid mud-puddler and visitor of flowers. It basks with its wings held wide open on tufts of grass and herbs, and generally keeps within a metre of the ground, even on cloudy days. It relies on its quick flight for escape.[15] 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.[15] 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 food plants.[15]
Research on freshly emerged imagines of P. 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.[18]
P. demoleus basking
While resting, the butterfly closes its wing over its back and draws the forewings between the hindwings.[15]
The number of generations of P. demoleus is dependent upon temperature[13] – near the equator, nine generations have been recorded,[12] while in warm temperate China, five generations have been recorded.[13] In the ideal conditions of a laboratory, a generation has been recorded to take place in just over 30 days.[13] The typical time for one generation of P. demoleus to mature in the field ranges from 26 to 59 days.[12] In cold climates, the lime butterfly is known to pass the winter as pupae.[12] Typically, the butterfly undergoes five instars as a caterpillar.[12]
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 1.5mm in height.[12][15][19] Fertile eggs develop a small red mark at the apex.[20]
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 head and two rows of subdorsal fleshy spines. The second, third, and fourth instars are dark, with a glossy, dark-brown head, and white markings on the eighth and ninth 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.[12][15]
As the instars progress, this resemblance is lost. From the fifth instar onwards, the caterpillars now turn cylindrical in shape, tapered towards the rear, and uniformly pale green in colour with a white subspiracular band. An additional black band is developed on the fourth and fifth segments with two black and two bluish spots on them. The eighth and ninth 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.[12][15]
The pupa, which is rugose (wrinkled), stout, and 30mm 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.[12][15]
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.[12][15][21]
The adults fly in every month, but are particularly abundant during and after the monsoons.[12]
Captive breeding of P. demoleus in Riyadh has revealed these data about the lifespan of various stages at that locality:[22]
Number of generations per year: 8
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
Early-instar "birds-dropping" caterpillar
early fifth-instar caterpillar
late fifth-instar caterpillar
Pupa in rearing cage
Parasitism and predation
Cocoons of a parasitoid wasp (Apanteles species), next to a perforated P. demoleus caterpillar
Despite their two-stage camouflage scheme, some caterpillars of P. demoleus are found by parasitic wasps, which lay dozens of eggs in them. The parasitic wasplarvae 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.[15]
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 preyed upon by two unidentified species of spiders which were abundant on citrus trees.[22]
In China, species of fungi in the genus Ophiocordyceps are known to parasitize many kinds of caterpillars, including P. demoleus.[citation needed] The spores were spread out on the parents, and infect the young caterpillar, then when the caterpillars become pupae, they will fail to develop into adults; instead, the fungi kill and eat the caterpillar flesh from within, and grow a spore bud out of the dead caterpillar corpse. The fungi known as dōng chóng xià cǎo are thought to have medicinal properties in China, and are known in English as caterpillar fungus.
In India, these braconid wasp parasitoids are known to parasitize P. demoleus larvae:[12]
In Jamaica, an encyrtid egg parasitoid and a chalcidoid parasitoid have been reported.[12]
Food plants
Key lime, a species of the cultivated lime, the principal food plant of P. demoleus
The larval food plants of P. demoleus in Asia are from the family Rutaceae, while in Australia and Papua New Guinea, the butterfly also feeds on host plants of family Fabaceae.[12]
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.[13] 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.[12]
Hand-picking of caterpillars and spraying with endosulfan 35 EC (2ml/10litres of water) were the recommended means of pest control by Indian government agencies and agricultural colleges,[24] however, endosulfan has since been banned by the Supreme Court of India.[25][26]
Papilio glaucus, the eastern tiger swallowtail, is a species of butterfly native to eastern North America. It is one of the most familiar butterflies in the eastern United States, ranging north to southern Ontario, Canada, and is common in many different habitats. It flies from spring until fall, during which it produces two to three broods. Adults feed on the nectar of many species of flowers, mostly from those of the families Apocynaceae, Asteraceae, and Fabaceae. P. glaucus has a wingspan measuring 7.9 to 14 cm. The male is yellow with four black "tiger stripes" on each forewing. Females may be either yellow or black, making them dimorphic. The yellow morph is similar to the male, but with a conspicuous band of blue spots along the hindwing, while the dark morph is almost completely black.
Papilio helenus, the red Helen, is a large swallowtail butterfly found in forests of southern India and parts of southeast Asia.
Graphium agamemnon, the tailed jay, is a predominantly green and black tropical butterfly that belongs to the swallowtail family. The butterfly is also called the green-spotted triangle, tailed green jay, or green triangle. It is a common, non-threatened species native to Nepal, India, Sri Lanka through Southeast Asia and Australia. Several geographic races are recognized. The species was first described by Carl Linnaeus in his 1758 10th edition of Systema Naturae.
Papilio polytes, the common Mormon, is a common species of swallowtail butterfly widely distributed across Asia.
Troides minos, the southern birdwing, also called Sahyadri birdwing, is a large and striking swallowtail butterfly endemic to South India. With a wingspan of 140–190 mm, it is the second largest butterfly of India. It is listed as Least Concern in the IUCN Red List.
Pachliopta aristolochiae, the common rose, is a swallowtail butterfly belonging to the genus Pachliopta, the roses, or red-bodied swallowtails. It is a common butterfly which is extensively distributed across south and southeast Asia.
Papilio liomedon, the Malabar banded swallowtail, is a member of the swallowtail butterfly family found in southern India. Earlier considered a subspecies of the banded swallowtail of southeast Asia, it is now considered a distinct species.
Papilio crino, the common banded peacock, is a species of swallowtail (Papilionidae) butterfly found in parts of the Indian subcontinent, including India, Nepal, Bhutan and Sri Lanka.
Papilio polymnestor, the blue Mormon, is a large swallowtail butterfly found in south India and Sri Lanka. It is the "state butterfly" of the Indian state of Maharashtra. With a wingspan of 120–150 mm, it is the fourth largest butterfly of India.
Papilio bootes, the tailed redbreast, is a swallowtail butterfly found in Asia. Within their wide distribution about four population variants have been named as subspecies. They have been placed within the Menelaides clade by a 2015 phylogenetics study.
Papilio krishna, the Krishna peacock, is a large swallowtail butterfly found in forests in China, Nepal, north east India, Myanmar and Vietnam.
Papilio protenor, the spangle, is a butterfly found in India belonging to the swallowtail family.
Papilio (Chilasa) agestor, the tawny mime, is a swallowtail butterfly, native to Indian subcontinent and widely found across Asia. The butterfly belongs to the mime subgenus, Chilasa, of the genus Papilio or the black-bodied swallowtails.
The giant swallowtail is the largest butterfly in North America. It is abundant through many parts of eastern North America; populations from western North America and down into Panama are now considered to belong to a different species, Papilio rumiko. Though it is often valued in gardens for its striking appearance, its larval stage can be a serious pest to citrus farms, which has earned its caterpillars the names orange dog or orange puppy. The giant swallowtail caterpillars possess remarkable camouflage from predators by closely resembling bird droppings. They use this, along with their osmeteria, to defend against predators such as wasps, flies, and vertebrates.
Papilio demodocus, the citrus swallowtail or Christmas butterfly, is a swallowtail butterfly which commonly occurs over the entirety of sub-Saharan Africa, including Madagascar, besides the southern Arabian Peninsula. The caterpillars feed on various native plants of especially the family Rutaceae, but have also taken to the leaves of cultivated citrus trees.
Papilio aegeus, the orchard swallowtail butterfly or large citrus butterfly is a species of butterfly from the family Papilionidae, that is found in eastern Australia and Papua New Guinea.
Papilio demolion, the banded swallowtail, is a species of swallowtail butterfly belonging to the family Papilionidae.
Papilio xuthus, the Asian swallowtail, Chinese yellow swallowtail, Japanese Swallowtail, or Xuthus swallowtail, is a yellow-colored, medium to large sized swallowtail butterfly found in northeast Asia, northern Myanmar, southern China, Taiwan, the Korean Peninsula, Japan, Siberia and the Hawaiian Islands. The butterfly has been observed once in Dunedin, New Zealand in 1996 emerging from a chrysalis in a car yard specializing in Japanese used cars; it is thought the chrysalis arrived through one of the cars. It was also recorded in the state of Arunachal Pradesh, India, in 2014.
Papilio anactus, the dainty swallowtail, dingy swallowtail or small citrus butterfly is a medium-sized butterfly from the family Papilionidae, that is endemic to Australia.
Papilio rumiko is a species of swallowtail butterfly. It is found from the south-western United States, through Mexico, El Salvador, Honduras, Costa Rica to Panama. The northernmost record is north-eastern Colorado.
References
1 2 Varshney, R.K.; Smetacek, Peter (2015). A Synoptic Catalogue of the Butterflies of India. New Delhi: Butterfly Research Centre, Bhimtal & Indinov Publishing, New Delhi. p.5. doi:10.13140/RG.2.1.3966.2164. ISBN978-81-929826-4-9.
↑ Evans, W.H. (1932). Identification of Indian Butterflies(Free full text download (first edition)) (2ed.). Mumbai: Bombay Natural History Society. pp.454 (with 32 plates). Retrieved 14 November 2010.
1 2 Wynter-Blyth, M.A. (1957). Butterflies of the Indian Region (Reprint of 2009 by Today & Tomorrow's Publishers, New Delhied.). Mumbai, India: Bombay Natural History Society. p.395. ISBN978-81-7019-232-9. Retrieved 22 October 2010.
↑ Ilse, Dora; Vaidya, Vidyadhar G. (1955). "Spontaneous feeding response to colours in Papilio demoleus L". Proceedings of the Indian Academy of Sciences, Section B (abstract). 43 (1): 23–31. doi:10.1007/BF03050215. S2CID82455044.
1 2 O, Akhtar. "Insect pests". Plant diseases identification. Retrieved 29 May 2020.
↑ 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. Bibcode:1978RSPSB.200..295S. doi:10.1098/rspb.1978.0021. JSTOR77392. S2CID84365809.
1 2 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). 91 (1–5). Blackwell Publishing Ltd: 286–292. doi:10.1111/j.1439-0418.1981.tb04481.x.
↑ Unattributed (2008). "Acid Lime (Citrus aurantifolia (Christm) Swingle)". TNAU Agritech Portal – Horticulture:Fruit Crops:Acid Lime. Tamil Nadu Agricultural University. Archived from the original on 19 October 2011. Retrieved 20 March 2012. .
Chattopadhyay, Jagannath. (2007), "Swallowtail Butterflies, Biology and Ecology of a Few Indian Species". Desh Prakashan, Kolkata, West Bengal, India. ISBN978-81-905719-1-3.
F. Martin Brown and Bernard Heineman, Jamaica and its Butterflies (E. W. Classey, London 1972), plate VIII
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