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Indian peafowl

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"Common peacock" redirects here. For the butterfly with this common name, see Papilio polyctor.

Indian peafowl
Peacock in Nagarhole National Park
Peahen in Bandipur National Park
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Aves
Order: Galliformes
Family: Phasianidae
Genus: Pavo
Species:
P. cristatus
Binomial name
Pavo cristatus
Map showing native range

The Indian peafowl (Pavo cristatus), also known as the common peafowl or blue peafowl, is a peafowl species native to the Indian subcontinent. While it originated in the Indian subcontinent, it has since been introduced to many other parts of the world. Male peafowl are referred to as peacocks, and female peafowl are referred to as peahens, although both sexes are often referred to colloquially as a "peacock".

The Indian peafowl displays a marked form of sexual dimorphism. The brightly coloured male has a blue coloured head with a fan-shaped crest and is best known for their long train. The train is made up of elongated upper-tail covert feathers with colourful eyespots. These stiff feathers are raised into a fan and quivered in a display during courtship. The peahen is predominantly brown in colour, with a white face and iridescent green lower neck, and lack the elaborate train. There are several colour mutations of the Indian peafowl including the leucistic white peafowl.

Despite the length and size of the covert feathers, the peacock is still capable of flight. The peafowl lives mainly on the ground in open forests or on cultivable lands where it forages for berries and grains, and also preys on snakes, lizards and small rodents. It makes loud calls, which makes it easier to detect, and are often used to indicate the presence of a predator in the forest areas. It forages on the ground in small groups and usually escapes on foot through undergrowth and avoids flying, though it flies into tall trees to roost.

The function of the Indian peacock's elaborate train has been debated for more than a century. In the 19th century, Charles Darwin found it a puzzle, hard to explain through ordinary natural selection. His later explanation, sexual selection, is widely but not universally accepted. In the 20th century, Amotz Zahavi argued that the train was a handicap, and that males were honestly signalling their fitness in proportion to the splendour of their trains. Despite extensive study, opinions remain divided on the mechanisms involved.

The Indian peafowl is listed as Least Concern on the IUCN Red List. It is the national bird of India and venerated in Hindu and Greek mythology.

Taxonomy and etymology

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Carl Linnaeus assigned the technical name of Pavo cristatus (means "crested peafowl" in classical Latin) in his work Systema Naturae in 1758.[2] The Latin word originated from the Greek word taos derived from Persian tavus, which is said to have come from the Tamil word tokei.[3] The Ancient Hebrew word tuki might have been derived from Tamil tokei or the Egyptian tekh.[4]

The earliest usage of the word in written English was from the 14th century where Geoffrey Chaucer (1343–1400) used the word in a simile "proud a pekok" in his epic poem Troilus and Criseyde.[5][6] Various spelling variants included peacock, pacok, pecok, pekok, pokok, and pocok among others.[3][6]

The Sanskrit word Mayura and the later Pali word Mora was probably derived from the Moriya region, which gave rise to Chandragupta Maurya of the Maurya Empire.[7]

Description

[edit]
Head of a male Indian peafowl, showing its fan shaped crest.

The Indian peafowl's size, color and shape of the crest make them easily identifiable within their native distribution range.[8] It displays a marked form of sexual dimorphism.[9][10] A male peafowl or peacock is a larger sized bird with an average bill to tail length of 100–120 cm (40–46 in) and as much as 200–230 cm (78–90 in) to the end of a fully grown train. It weighs 4.1–5.2 kg (9–11.5 lb) and is amongst the heaviest birds in Phasianidae. The male has a metallic blue crown with short and curled, blue-greenish head feathers. It has a fan-shaped crest with bare black shafts and tipped with bluish-green webbing. A white stripe above the eye and a crescent shaped white patch below the eye are formed by bare white skin. The lore, chin and throat are covered with greenish feathers. It has a long blue neck with scaly bronze-green feathers with black and copper markings in the back. The scapular region and wings are made of chestnut colored primary feathers with black secondaries. The tail is dark brown with glossy green chest, buff thighs, and blackish-brown abdomen and tail coverts.[11][12]

The male is best known for its elongated train, which extend from the rump. The train is made up of elongated upper tail coverts, which are bronze-green train with the outermost and longer feathers ending up with an elaborate eye-spot. The eye-spots consist of a purplish-black, heart-shaped nucleus, enclosed by blue and an outer copper rim, which is surrounded by alternating green and bronze. A few of the outer feathers lack the spot and end in a crescent shaped black tip.[11][12] The feathers of the train does not have colored pigments and the colorization is a result of the micro-structure of the feathers and the optical phenomena involved.[13] The male has a spur on the leg above the hind toe. The train feathers and the tarsal spur of the male starts developing only in the second year of its life. The trains are not fully developed until the age of four.[14] The train feathers of the male Indian peafowl are also moulted every year, usually starting at the end of the monsoon in August or September and are fully developed by February to March.[11][14] The moult of the flight feathers may be spread out across the year.[15]

The females or peahens, are smaller at around 15 in (38 cm) in length and weigh 2.7–4.1 kg (6–9 lb). The peahen has a rufous-brown head with a crest, whose tips are chestnut colored and edged with green. The upper body is brownish with pale mottling and the primaries, secondaries and tail are dark brown. The lower neck is metallic green with dark brown breast feathers glossed with green and whitish underparts. Both the sexes have dark brown eyes, brown colored beak and legs. Young males also resemble the females with chestnut colored primaries.[11].[12]

Mutations and hybrids

[edit]
A melanistic black-shouldered Indian peafowl specimen from Naturalis Biodiversity Center, Leiden.
A displaying leucistic white Indian peafowl from Jardin des Plantes, Paris.

There are several colour mutations of Indian peafowl. These very rarely occur in the wild, but selective breeding has made them common in captivity. The black-shouldered mutation was initially considered as a subspecies or even a separate species of the Indian peafowl (P. nigripennis).[16] English naturalist Charles Darwin (1809–1882) presented firm evidence for it being a variety under domestication, which is now well established and accepted. It was important for Darwin to prove that it was a colour variation rather than a wild species as it was contrary to his theory of slow modification by natural selection in the wild.[17] In this genetic variation, the adult male is melanistic with black wings. The young birds are creamy white with fulvous-tipped wings. The gene which produces melanism in the male, causes s a dilution of colour in females, which have creamy white and brown markings.[11][18] Other forms of mutations include the pied and white mutations, which are the result of allelic variation at specific loci.[19][20]

Crosses between a male green peafowl (Pavo muticus) and a female Indian peafowl (P. cristatus) produce a stable hybrid called a "Spalding", named after Keith Spalding, a bird fancier from California.[21] There can be problems if birds of unknown pedigree are released into the wild, as the viability of such hybrids and their offspring is often reduced as per Haldane's rule.[22][23]

Distribution and habitat

[edit]
The Indian peafowl is found across most of India and Sri Lanka, in forests, bushy lands, and hills. Pictured are a displaying male peafowl (left) and a female with chicks.

The Indian peafowl is a resident breeder in the Indian subcontinent and is found across most of India and Sri Lanka. In India, it is found across the country from the Indus valley in the north-west to Assam in the north-east, and from Himalayas in the north to the southern tip, except for the marshlands of Sunderbans in East India. In India, it is found up to elevations of 5,000 ft (1,500 m) in the north and upto 6,000 ft (1,800 m) in the mountains of the south. In Sri Lanka, it largely inhabits the drier lowland areas. It is generally found in forests, small hills, and bushy areas near water sources. It also occupies cultivable lands and live in a semi-domesticated state in human habitations.[24][12][25] The peafowl has since been introduced in many other parts of the world and has become feral in some areas.[26][27] It was supposedly introduced into Europe by Alexander the Great, while the bird might have been introduced earlier and had reached Athens by 450 BCE.[28][29]

Genome sequencing studies of the Indian peafowl have indicated that the population suffered at least two bottlenecks (four mya and 450,000 years ago), which resulted in a severe reduction in its effective population size.[30] Population studies of birds at roosting sites have indicated a higher proportion of males in Northern India (170–210 for 100 females) but a lower proportion in Southern India (47 males for 100 females).[31]

Behaviour and ecology

[edit]

Peafowl are best known for the male's extravagant display feathers which, despite actually growing from their back, are thought of as a tail. The "train" is in reality made up of the enormously elongated upper tail coverts. The tail itself is brown and short as in the peahen. The colours result not from any green or blue pigments but from the micro-structure of the feathers and the resulting optical phenomena.[32] The long train feathers (and tarsal spurs) of the male develop only after the second year of life. Fully developed trains are found in birds older than four years. In northern India, these begin to develop each February and are moulted at the end of August.[33] The moult of the flight feathers may be spread out across the year.[34]

Peafowl forage on the ground in small groups, known as musters, that usually have a cock and 3 to 5 hens. After the breeding season, the flocks tend to be made up only of females and young. They are found in the open early in the mornings and tend to stay in cover during the heat of the day. They are fond of dust-bathing and at dusk, groups walk in single file to a favourite waterhole to drink. When disturbed, they usually escape by running and rarely take to flight.[26]

Peafowl produce loud calls especially in the breeding season. They may call at night when alarmed and neighbouring birds may call in a relay like series. Nearly seven different call variants have been identified in the peacocks apart from six alarm calls that are commonly produced by both sexes.[35]

Peafowl roost in groups during the night on tall trees but may sometimes make use of rocks, buildings or pylons. In the Gir forest, they chose tall trees in steep river banks.[36][37] Birds arrive at dusk and call frequently before taking their position on the roost trees.[38] Due to this habit of congregating at the roost, many population studies are made at these sites. The population structure is not well understood. In a study in northern India (Jodhpur), the number of males was 170–210 for 100 females but a study involving evening counts at the roost site in southern India suggested a ratio of 47 males for 100 females.[31]

Sexual selection

[edit]
See also: Fisherian runaway
Thayer in his "Peacock in the Woods" (1907) suggested that the function of the ornate tail was camouflage

The colours of the peacock and the contrast with the much duller peahen were a puzzle to early thinkers. Charles Darwin wrote to Asa Gray that the "sight of a feather in a peacock's tail, whenever I gaze at it, makes me sick!" as he failed to see an adaptive advantage for the extravagant tail which seemed only to be an encumbrance. Darwin developed a second principle of sexual selection to resolve the problem, though in the prevailing intellectual trends of Victorian Britain, the theory failed to gain widespread attention.[39]

The American artist Abbott Handerson Thayer tried to show, from his own imagination, the value of the eyespots as disruptive camouflage in a 1907 painting.[40] He used the painting in his 1909 book Concealing-Coloration in the Animal Kingdom, denying the possibility of sexual selection and arguing that essentially all forms of animal colouration had evolved as camouflage.[41] He was roundly criticised in a lengthy paper by Theodore Roosevelt, who wrote that Thayer had only managed to paint the peacock's plumage as camouflage by sleight of hand, "with the blue sky showing through the leaves in just sufficient quantity here and there to warrant the author-artists explaining that the wonderful blue hues of the peacock's neck are obliterative because they make it fade into the sky."[42]

In the 1970s a possible resolution to the apparent contradiction between natural selection and sexual selection was proposed. Amotz Zahavi argued that peacocks honestly signalled the handicap of having a large and costly train. However, the mechanism may be less straightforward than it seems – the cost could arise from depression of the immune system by the hormones that enhance feather development.[43][44]

Male courting female

The ornate train is believed to be the result of sexual selection by the females. Males use their ornate trains in a courtship display: they raise the feathers into a fan and quiver them. However, recent studies have failed to find a relation between the number of displayed eyespots and mating success.[45] Marion Petrie tested whether or not these displays signaled a male's genetic quality by studying a feral population of peafowl in Whipsnade Wildlife Park in southern England. She showed that the number of eyespots in the train predicted a male's mating success, and this success could be manipulated by cutting the eyespots off some of the male's ornate feathers.[46]

Although the removal of eyespots makes males less successful in mating,[46] eyespot removal substantially changes the appearance of male peafowls. It is likely that females mistake these males for sub-adults, or perceive that the males are physically damaged. Moreover, in a feral peafowl population, there is little variation in the number of eyespots in adult males. It is rare for adult males to lose a significant number of eyespots. Therefore, females' selection might depend on other sexual traits of males' trains. The quality of train is an honest signal of the condition of males; peahens do select males on the basis of their plumage. A recent study on a natural population of Indian peafowls in the Shivalik area of India has proposed a "high maintenance handicap" theory. It states that only the fittest males can afford the time and energy to maintain a long tail. Therefore, the long train is an indicator of good body condition, which results in greater mating success.[47] While train length seems to correlate positively with MHC diversity in males, females do not appear to use train length to choose males.[48] A study in Japan also suggests that peahens do not choose peacocks based on their ornamental plumage, including train length, number of eyespots and train symmetry.[49] Another study in France brings up two possible explanations for the conflicting results that exist. The first explanation is that there might be a genetic variation of the trait of interest under different geographical areas due to a founder effect and/or a genetic drift. The second explanation suggests that "the cost of trait expression may vary with environmental conditions," so that a trait that is indicative of a particular quality may not work in another environment.[46]

Fisher's runaway model proposes positive feedback between female preference for elaborate trains and the elaborate train itself. This model assumes that the male train is a relatively recent evolutionary adaptation. However, a molecular phylogeny study on peacock-pheasants shows the opposite; the most recently evolved species is actually the least ornamented one.[50] This finding suggests a chase-away sexual selection, in which "females evolve resistance to male ploys".[51] A study in Japan goes on to conclude that the "peacocks' train is an obsolete signal for which female preference has already been lost or weakened".[49]

However, some disagreement has arisen in recent years concerning whether or not female peafowl do indeed select males with more ornamented trains. In contrast to Petrie's findings, a seven-year Japanese study of free-ranging peafowl came to the conclusion that female peafowl do not select mates solely on the basis of their trains. Mariko Takahashi found no evidence that peahens expressed any preference for peacocks with more elaborate trains (such as trains having more ocelli), a more symmetrical arrangement, or a greater length.[49] Takahashi determined that the peacock's train was not the universal target of female mate choice, showed little variance across male populations, and, based on physiological data collected from this group of peafowl, do not correlate to male physical conditions. Adeline Loyau and her colleagues responded to Takahashi's study by voicing concern that alternative explanations for these results had been overlooked, and that these might be essential for the understanding of the complexity of mate choice.[46] They concluded that female choice might indeed vary in different ecological conditions.

A 2013 study that tracked the eye movements of peahens responding to male displays found that they looked in the direction of the upper train of feathers only when at long distances and that they looked only at the lower feathers when males displayed close to them. The rattling of the tail and the shaking of the wings helped in keeping the attention of females.[52]

Breeding

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Egg, collection Museum Wiesbaden
Peahen with three chicks near Hodal in Faridabad District of Haryana, India

Peacocks are polygamous, and the breeding season is spread out but appears to be dependent on the rains. Peafowls usually reach sexual maturity at the age of 2 to 3 years old.[53] Several males may congregate at a lek site and these males are often closely related.[54] Males at leks appear to maintain small territories next to each other and they allow females to visit them and make no attempt to guard harems. Females do not appear to favour specific males.[55] The males display in courtship by raising the upper-tail coverts into an arched fan. The wings are held half open and drooped and it periodically vibrates the long feathers, producing a ruffling sound. The cock faces the hen initially and struts and prances around and sometimes turns around to display the tail.[26] Males may also freeze over food to invite a female in a form of courtship feeding.[56] Males may display even in the absence of females. When a male is displaying, females do not appear to show any interest and usually continue their foraging.[31]

The peak season in southern India is April to May, January to March in Sri Lanka and June in northern India. The nest is a shallow scrape in the ground lined with leaves, sticks and other debris. Nests are sometimes placed on buildings[57] and, in earlier times, have been recorded using the disused nest platforms of the white-rumped vultures. The clutch consists of 4–8 fawn to buff white eggs which are incubated only by the female. The eggs take about 28 days to hatch. The chicks are nidifugous and follow the mother around after hatching.[12] Downy young may sometimes climb on their mothers' back and the female may carry them in flight to a safe tree branch.[58] An unusual instance of a male incubating a clutch of eggs has been reported.[26][59]

Feeding

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Peafowl are omnivorous and eat seeds, insects (including termites), worms,[60] fruits, small mammals, frogs,[60] and reptiles (such as lizards).[60] They feed on small snakes but keep their distance from larger ones.[61] In the Gir forest of Gujarat, a large percentage of their food is made up of the fallen berries of Zizyphus.[62] They also feed on tree and flower buds, petals, grain, and grass and bamboo shoots.[60] Around cultivated areas, peafowl feed on a wide range of crops such as groundnut, tomato, paddy, chili and even bananas.[31] Around human habitations, they feed on a variety of food scraps and even human excreta.[26] In the countryside, it is particularly partial to crops and garden plants.

Mortality factors

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Large animals such as leopards, dholes, golden jackals, lions, and tigers can ambush adult peafowls.[37][63] However, only leopards regularly prey upon peafowls as adult peafowls are difficult to catch since they can usually escape ground predators by flying into trees.[64][65][66] They are also sometimes hunted by large birds of prey such as the changeable hawk-eagle and rock eagle-owl.[67][68] Chicks are somewhat more prone to predation than adult birds. Adults living near human habitations are sometimes hunted by domestic dogs or by humans in some areas (southern Tamil Nadu) for folk remedies involving the use of "peacock oil".[31]

Foraging in groups provides some safety as there are more eyes to look out for predators.[69] They also roost on high tree tops to avoid terrestrial predators, especially leopards.[37]

In captivity, birds have been known to live for 23 years but it is estimated that they live for only about 15 years in the wild.[70]

Conservation and status

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A white peafowl in Indira Gandhi Zoological Park, Visakhapatnam

Indian peafowl are widely distributed in the wild across South Asia and protected both culturally in many areas and by law in India. Conservative estimates of the population put them at more than 100,000.[71] Illegal poaching for meat, however, continues and declines have been noted in parts of India.[72] Peafowl breed readily in captivity and as free-ranging ornamental fowl. Zoos, parks, bird-fanciers and dealers across the world maintain breeding populations that do not need to be augmented by the capture of wild birds.

Poaching of peacocks for their meat and feathers and accidental poisoning by feeding on pesticide treated seeds are known threats to wild birds.[73] Methods to identify if feathers have been plucked or have been shed naturally have been developed, as Indian law allows only the collection of feathers that have been shed.[74]

In parts of India, the birds can be a nuisance to agriculture as they damage crops.[26] Its adverse effects on crops, however, seem to be offset by the beneficial role it plays by consuming prodigious quantities of pests such as grasshoppers. They can also be a problem in gardens and homes where they damage plants, attack their reflections (thereby breaking glass and mirrors), perch and scratch cars or leave their droppings. Many cities where they have been introduced and gone feral have peafowl management programmes. These include educating citizens on how to prevent the birds from causing damage while treating the birds humanely.[75][76][77]

In culture

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Kartikeya with his consorts riding a peacock, painting by Raja Ravi Varma

Prominent in many cultures, the peacock has been used in numerous iconic representations, including being designated the national bird of India in 1963.[26] The peacock, known as mayura in Sanskrit, has enjoyed a fabled place in India since and is frequently depicted in temple art, mythology, poetry, folk music and traditions.[78] A Sanskrit derivation of mayura is from the root mi for kill and said to mean "killer of snakes".[79] It is also likely that the Sanskrit term is a borrowing from Proto-Dravidian *mayVr (whence the Tamil word for peacock மயில் (mayil)) or a regional Wanderwort.[80][81] Many Hindu deities are associated with the bird, Krishna is often depicted with a feather in his headband, while worshippers of Shiva associate the bird as the steed of the God of war, Kartikeya (also known as Skanda or Murugan). A story in the Uttara Ramayana describes the head of the Devas, Indra, who unable to defeat Ravana, sheltered under the wing of peacock and later blessed it with a "thousand eyes" and fearlessness from serpents.[79] Another story has Indra who after being cursed with a thousand ulcers was transformed into a peacock with a thousand eyes and this curse was removed by Vishnu.[82]

In Buddhist philosophy, the peacock represents wisdom.[83] Peacock feathers are used in many rituals and ornamentation. Peacock motifs are widespread in Indian temple architecture, old coinage, textiles and continue to be used in many modern items of art and utility.[29] A folk belief found in many parts of India is that the peacock does not copulate with the peahen but that she is impregnated by other means. The stories vary and include the idea that the peacock looks at its ugly feet and cries whereupon the tears are fed on by the peahen causing it to be orally impregnated while other variants incorporate sperm transfer from beak to beak.[84] Similar ideas have also been ascribed to Indian crow species.[85] In Greek mythology the origin of the peacock's plumage is explained in the tale of Hera and Argus.[21] The main figure of the Yazidi religion Yezidism, Melek Taus, is most commonly depicted as a peacock.[86][87] Peacock motifs are widely used even today such as in the logos of the US NBC and the PTV television networks and the Sri Lankan Airlines.

A peacock or in his pride, on a field azure, on the arms of Saint-Paul, Savoie

These birds were often kept in menageries and as ornaments in large gardens and estates. In medieval times, knights in Europe took a "Vow of the Peacock" and decorated their helmets with its plumes. In several Robin Hood stories, the titular archer uses arrows fletched with peacock feathers. Feathers were buried with Viking warriors[88] and the flesh of the bird was said to cure snake venom and many other maladies. Numerous uses in Ayurveda have been documented. Peafowl were said to keep an area free of snakes.[89] In 1526, the legal issue as to whether peacocks were wild or domestic fowl was thought sufficiently important for Cardinal Wolsey to summon all the English judges to give their opinion, which was that they are domestic fowl.[90]

In Anglo-Indian usage of the 1850s, to peacock meant making visits to ladies and gentlemen in the morning. In the 1890s, the term "peacocking" in Australia referred to the practice of buying up the best pieces of land ("picking the eyes") so as to render the surrounding lands valueless.[91] The English word "peacock" has come to be used to describe a man who is very proud or gives a lot of attention to his clothing.[92]

A golden peacock (in Yiddish, Di Goldene Pave) is considered by some as a symbol of Ashkenazi Jewish culture, and is the subject of several folktales and songs in Yiddish.[93][94] Peacocks are frequently used in European heraldry. Heraldic peacocks are most often depicted as facing the viewer and with their tails displayed. In this pose, the peacock is referred to as being "in his pride". Peacock tails, in isolation from the rest of the bird, are rare in British heraldry, but see frequent use in German systems.[95]

The American television network NBC uses a stylized peacock as a legacy of its early introduction of color television, alluding to the brilliant color of a peacock, and continues to promote the bird as a trademark of its broadcasting and streaming services.

References

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Further reading

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  • Galusha, JG; Hill, LM (1996) A study of the behaviour of Indian Peacocks Pavo cristatus on Protection Island, Jefferson County, Washington, USA. Pavo 34(1&2):23–31.
  • Ganguli, U (1965) A Peahen nests on a roof. Newsletter for Birdwatchers . 5(4):4–6.
  • Prakash, M (1968) Mating of Peacocks Pavo cristatus. Newsletter for Birdwatchers . 8(6), 4–5.
  • Rao, MS; Zaki, S; Ganesh, T (1981). "Colibacillosis in a Peacock". Current Science. 50 (12): 550–551.
  • Sharma, IK (1969). "Habitat et comportment du Pavon (Pavo cristatus)". Alauda. 37 (3): 219–223.
  • Sharma, IK (1970). "Analyse ecologique des parades du paon (Pavo cristatus)". Alauda. 38 (4): 290–294.
  • Sharma, IK (1972). "Etude ecologique de la reproduction de la paon (Pavo cristatus)". Alauda. 40 (4): 378–384.
  • Sharma, IK (1973). "Ecological studies of biomass of the Peafowl (Pavo cristatus)". Tori. 22 (93–94): 25–29. doi:10.3838/jjo1915.22.25.
  • Sharma, IK (1974). "Notes ecologique sur le paon bleu, Pavo cristatus". Les Carnets de Zoologie. 34: 41–45.
  • Sharma, IK (1981). "Adaptations and commensality of the Peafowl (Pavo cristatus) in the Indian Thar Desert". Annals Arid Zone. 20 (2): 71–75.
  • Shrivastava AB, Nair NR, Awadhiya RP, Katiyar AK (1992). "Traumatic ventriculitis in Peacock (Pavo cristatus)". Indian Vet. J. 69 (8): 755.
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