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Orangutan

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(Redirected fromPongo (genus))
Genus of Asian apes
This article is about the group of primates. For other uses, seeOrangutan (disambiguation).
A request that this article title be changed toOrangutangOrangutang isunder discussion. Pleasedo not move this article until the discussion is closed.

Orangutans
Temporal range:Early Pleistocene – Recent
Orang Utan, Semenggok Forest Reserve, Sarawak, Borneo, Malaysia.JPG
Bornean orangutan
(Pongo pygmaeus)
Scientific classificationEdit this classification
Kingdom:Animalia
Phylum:Chordata
Class:Mammalia
Order:Primates
Suborder:Haplorhini
Infraorder:Simiiformes
Family:Hominidae
Subfamily:Ponginae
Genus:Pongo
Lacépède, 1799
Type species
Pongo borneo
Lacépède, 1799
(=Simia pygmaeusLinnaeus, 1760)[1]
Species

Pongo pygmaeus
Pongo abelii
Pongo tapanuliensis
Pongo hooijeri
Pongo weidenreichi

Range of the threeextant species
Synonyms

FaunusOken, 1816
LophotusFischer, 1813
MacrobatesBillberg, 1828
SatyrusLesson, 1840

Orangutans aregreat apes native to therainforests ofIndonesia andMalaysia. They are now found only in parts ofBorneo andSumatra, but during thePleistocene they ranged throughoutSoutheast Asia andSouth China. Classified in thegenusPongo, orangutans were originally considered to be one species. In 1996, they were divided into two species: theBornean orangutan (P. pygmaeus, with three subspecies) and theSumatran orangutan (P. abelii); a third species, theTapanuli orangutan (P. tapanuliensis), was identified definitively in 2017. The orangutans are the only surviving members of the subfamilyPonginae, which diverged genetically from the other hominids (gorillas,chimpanzees, andhumans) between 19.3 and 15.7 million years ago.

The mostarboreal of the great apes, orangutans spend most of their time in trees. They have proportionally long arms and short legs, and have reddish-brown hair covering their bodies. Adult males weigh about 75 kg (165 lb), while females weigh about 37 kg (82 lb).Dominant adult males develop distinctive cheek pads or flanges and make long calls that attract females and intimidate rivals; younger subordinate males do not and more resemble adult females. Orangutans are the most solitary of the great apes: social bonds occur primarily between mothers and their dependent offspring. Fruit is the most important component of an orangutan's diet, but they will also eat vegetation,bark,honey, insects and bird eggs. They can live over 30 years, both in the wild and in captivity.

Orangutans are among the most intelligentprimates. Theyuse a variety of sophisticated tools and construct elaborate sleeping nests each night from branches and foliage. The apes' learning abilities have been studied extensively. There may be distinctive cultures within populations. Orangutans have been featured in literature and art since at least the 18th century, particularly in works that comment on human society. Field studies of the apes were pioneered by primatologistBirutė Galdikas and they have been kept in captive facilities around the world since at least the early 19th century.

All three orangutan species are consideredcritically endangered. Human activities have caused severe declines in populations and ranges. Threats to wild orangutan populations includepoaching (forbushmeat and retaliation for consumingcrops),habitat destruction anddeforestation (forpalm oil cultivation andlogging), and the illegalpet trade. Severalconservation andrehabilitation organisations are dedicated to the survival of orangutans in the wild.

Etymology

Most Western sources attribute the name "orangutan" (also written orang-utan, orang utan, orangutang, and ourang-outang[2]) to theMalay wordsorang, meaning 'person', andhutan, meaning 'forest'.[3][4] The Malay used the term to indicate forest-dwelling humans; the first recorded Malay use of "orang-utan" referring to the ape identifies it as a Western term.[5]: 12  There is, however, some evidence to suggest that the term may have been used in regard to apes in premodernOld Malay.[3]

In Western sources, the first printed attestation of the word for the apes is in Dutch physicianJacobus Bontius's 1631Historiae naturalis et medicae Indiae orientalis. He reported that Malays claimed that the ape could talk but preferred not to "lest he be compelled to labour".[6] The word appeared in several German-language descriptions of Indonesian zoology in the 17th century. It has been argued that the word comes specifically from theBanjarese variety of Malay,[7] but the age of the Old Javanese sources mentioned above make Old Malay a more likely origin for the term. Cribb and colleagues (2014) suggest that Bontius's account referred not to apes (as this description was from Java where the apes were not known to be from) but to humans suffering some serious medical condition (most likelycretinism) and that his use of the word was misunderstood byNicolaes Tulp, who was the first to use the term in a publication a decade later.[5]: 10–18 

The word was first attested in English in 1693 by physicianJohn Bulwer in the formOrang-Outang,[2][8] and variants ending with-ng are found in many languages. This spelling (and pronunciation) has remained in use in English up to the present but has come to be regarded asincorrect.[9][10][11] The loss of "h" inhutan and the shift from -ng to -n has been taken to suggest the term entered English throughPortuguese.[7] In Malay, the term was first attested in 1840, not as an indigenous name but referring to how the English called the animal.[12] The word 'orangutan' in modern Malay and Indonesian was borrowed from English orDutch in the 20th century—explaining the missing initial 'h' of 'hutan'.[7]

The name of the genus,Pongo, comes from a 16th-century account byAndrew Battel, an English sailor held prisoner by the Portuguese inAngola, which describes two anthropoid "monsters" named Pongo and Engeco. He is now believed to have been describinggorillas, but in the 18th century, the terms orangutan and pongo were used for allgreat apes. French naturalistBernard Germain de Lacépède used the termPongo for the genus in 1799.[13][5]: 24–25  Battel's "Pongo", in turn, is from theKongo wordmpongi[14][15] or othercognates from the region:Lumbupungu,Vilimpungu, orYombiyimpungu.[16]

Taxonomy and phylogeny

See also:Orangutan–human last common ancestor

The orangutan was firstdescribed scientifically in 1758 in theSystema Naturae ofCarl Linnaeus asHomo troglodytes.[5]: 20  It was renamedSimiapygmaeus in 1760 by his studentChristian Emmanuel Hopp and given the namePongo by Lacépède in 1799.[5]: 24–25  The populations on the two islands were suggested to be separate species whenP. abelii was described by French naturalistRené Lesson in 1827.[17] In 2001,P. abelii was confirmed as a full species based on molecular evidence published in 1996,[18][19]: 53 [20] and three distinct populations onBorneo were elevated to subspecies (P. p. pygmaeus,P. p. morio andP. p. wurmbii).[21] The description in 2017 of a third species,P. tapanuliensis, from Sumatra south ofLake Toba, came with a surprising twist: it is more closely related to the Bornean species,P. pygmaeus than to its fellow Sumatran species,P. abelii.[20]

Head shots of male Bornean, Sumatran and Tapanuli orangutans
Flanged maleBornean,Sumatran andTapanuli orangutans

The Sumatran orangutan genome was sequenced in January 2011.[22][23] Following humans andchimpanzees, the Sumatran orangutan became the third species ofgreat ape whose genome was sequenced. Subsequently, the Bornean species's genome was sequenced. Bornean orangutans (P. pygmaeus) exhibit lessgenetic diversity than Sumatran orangutans (P. abelii), despite the latter's population being six to seven times greater. The researchers hope these genetic data may help conservationists preserve the endangered ape, as well as learn more about humangenetic diseases.[23] Similarly to gorillas and chimpanzees, orangutans have 48 diploid chromosomes, in contrast to humans,which have 46.[24]: 30 

According tomolecular evidence, within apes (superfamily Hominoidea), thegibbons diverged during the earlyMiocene between 24.1 and 19.7 million years ago (mya), and the orangutans diverged from the African great ape lineage between 19.3 and 15.7 mya. Israfil and colleagues (2011) estimated based onmitochondrial,Y-linked, andX-linkedloci that the Sumatran and Bornean species diverged 4.9 to 2.9 mya.[25]: Fig. 4  By contrast, the 2011 genome study suggested that these two species diverged as recently as circa 400,000 years ago. The study also found that orangutans evolved at a slower pace than both chimpanzees and humans.[23] A 2017 genome study found that the Bornean and Tapanuli orangutans diverged from Sumatran orangutans about 3.4 mya, and from each other around 2.4 mya. Millions of years ago, orangutans travelled from mainland Asia to Sumatra and then Borneo as the islands were connected byland bridges during therecent glacial periods when sea levels were much lower. The present range of Tapanuli orangutans is thought to be close to where ancestral orangutans first entered what is now Indonesia from mainland Asia.[20][26]

Taxonomy of genusPongo[1]Phylogeny of superfamily Hominoidea[25]: Fig. 4 
GenusPongo
  • Bornean orangutan (Pongo pygmaeus)
    • Pongo pygmaeus pygmaeus – northwest populations
    • Pongo pygmaeus morio – east populations
    • Pongo pygmaeus wurmbii – southwest populations
  • Sumatran orangutan (Pongo abelii – Sumatra northwest ofLake Toba)
  • Tapanuli orangutan (Pongo tapanuliensis – Sumatra south of Lake Toba)
Hominoidea

gibbons (family Hylobatidae)

orangutans (genusPongo)

gorillas (genusGorilla)

chimpanzees (genusPan)

humans (genusHomo)

Partial fossil skull of ape
Fossil skull ofSivapithecus sivalensis, an extinct relative of orangutan

Fossil record

The three orangutan species are the only extant members of the subfamilyPonginae. Thissubfamily also includes extinctapes such asLufengpithecus, which occurred 8–2 mya in southern China and Thailand;[19]: 50 Indopithecus, which lived in India from 9.2 to 8.6 mya; andSivapithecus, which lived in India and Pakistan from 12.5 mya until 8.5 mya.[27] These animals likely lived in drier and cooler environments than orangutans do today.Khoratpithecus piriyai, which lived 5–7 mya in Thailand, is believed to be the closest known relative of the living orangutans and inhabited similar environments.[19]: 50  The largest known primate,Gigantopithecus, was also a member of Ponginae and lived in China, from 2 mya to 300,000 years ago.[28][19]: 50 

The oldest known record ofPongo is from theEarly Pleistocene ofChongzuo, consisting of teeth ascribed to extinct speciesP. weidenreichi.[29][30]Pongo is found as part of the faunal complex in the Pleistocene cave assemblage in Vietnam, alongsideGiganopithecus, though it is known only from teeth. Somefossils described under the nameP. hooijeri have been found in Vietnam, and multiple fossil subspecies have been described from several parts of southeastern Asia. It is unclear if these belong toP. pygmaeus orP. abelii or, in fact, represent distinct species.[31] During the Pleistocene,Pongo had a far more extensive range than at present, extending throughoutSundaland and mainland Southeast Asia and South China. Teeth of orangutans are known fromPeninsular Malaysia that date to 60,000 years ago.[32] The youngest remains from South China, which are teeth assigned toP. weidenreichi, date to between 66 and 57,000 years ago.[33] The range of orangutans had contracted significantly by the end of the Pleistocene, most likely because of the reduction of forest habitat during theLast Glacial Maximum. They may have nevertheless survived into the Holocene in Cambodia and Vietnam.[29][32]

Characteristics

Head and shoulder shots of an adult male and female orangutan
Adult male (left) and female Tapanuli orangutans

Orangutans display significantsexual dimorphism; females typically stand 115 cm (45 in) tall and weigh around 37 kg (82 lb), while adult males stand 137 cm (54 in) tall and weigh 75 kg (165 lb). The tallest orangutan recorded was a 180 cm (71 in). Compared to humans, they have proportionally long arms, a male orangutan having anarm span of about 2 m (6 ft 7 in), and short legs. They are covered in long reddish hair that starts out bright orange and darkens tomaroon orchocolate with age, while the skin is grey-black. Though largely hairless, males' faces can develop some hair, giving them a beard.[34][35][19]: 13–15 

Orangutans have small ears and noses; the ears are unlobed.[34] The meanendocranial volume is 397 cm3.[36] The cranium is elevated relative to the face, which is incurved andprognathous.[34] Compared to chimpanzees and gorillas, thebrow ridge of an orangutan is underdeveloped.[37] Females and juveniles have relatively circular skulls and thin faces while mature males have a prominentsagittal crest, large cheek pads or flanges,[34] extensive throat pouches and longcanines.[34][19]: 14  The cheek pads are made mostly of fatty tissue and are supported by the musculature of the face.[38] The throat pouches act asresonance chambers for making long calls.[39]

An orangutan skeleton
Skeleton of subadult Bornean orangutan

Orangutan hands have four long fingers but a dramatically shorteropposable thumb for a strong grip on branches as they travel high in the trees. The resting configuration of the fingers is curved, creating a suspensory hook grip. With the thumb out of the way, the fingers (and hands) can grip securely around objects with a small diameter by resting the tops of the fingers against the inside of the palm, thus creating a double-locked grip.[40] Their feet have four long toes and an opposable big toe, giving them hand-like dexterity. The hip joints also allow for their legs to rotate similarly to their arms and shoulders.[19]: 15 

Orangutans move through the trees by both vertical climbing andsuspension. Compared to other great apes, they infrequently descend to the ground where they are more cumbersome. Unlike gorillas and chimpanzees, orangutans are not trueknuckle-walkers, instead bending their digits and walking on the sides of their hands and feet.[41][42]

Compared to their relatives in Borneo, Sumatran orangutans are more slender with paler and longer hair and a longer face.[35] Tapanuli orangutans resemble Sumatran orangutans more than Bornean orangutans in body build and hair colour.[20] They have shaggier hair, smaller skulls, and flatter faces than the other two species.[43]

Ecology and behaviour

Wild orangutan in the Danum Valley (Sabah, Malaysia, Borneo island)

Orangutans are mainlyarboreal and inhabittropical rainforest, particularly lowlanddipterocarp and oldsecondary forest.[35][44] Populations are more concentrated near riverside habitats, such asfreshwater andpeat swamp forest, while drier forests away from the flooded areas have fewer apes. Population density also decreases at higher elevations.[24]: 92  Orangutans occasionally enter grasslands, cultivated fields, gardens, youngsecondary forest, and shallow lakes.[44]

Most of the day is spent feeding, resting, and travelling.[45] They start the day feeding for two to three hours in the morning. They rest during midday, then travel in the late afternoon. When evening arrives, they prepare their nests for the night.[44] Potential predators of orangutans includetigers,clouded leopards andwild dogs.[24]: 91  The most common orangutan parasites arenematodes of the genusStrongyloides and theciliateBalantidium coli. AmongStrongyloides, the speciesS. fuelleborni andS. stercoralis are reported in young individuals.[46] Orangutans also use the plant speciesDracaena cantleyi as an anti-inflammatory balm.[47] Captive animals may suffer an upper respiratory tract disease.[48]

Diet and feeding

Orangutan on a branch eating some leaves
Although orangutans may consume leaves, shoots, and bird eggs, fruit is the most important part of their diet.

Orangutans are primarilyfruit-eaters, which can take up 57–80% of their foraging time. Even during times of scarcity, fruit is 16% of their feeding time. Fruits with soft pulp,arils or seed-walls are consumed the most, particularlyfigs but alsodrupes and berries.[24]: 65  Orangutans are thought to be the sole fruit disperser for some plant species including thevine speciesStrychnos ignatii which contains the toxicalkaloidstrychnine.[49]

Orangutans also include leaves in their diet, which take up 25% of their average foraging time. Leaves are eaten more when fruit is less available, but even during times of fruit abundance, orangutans will eat leaves 11–20% of the time. They appear to depend on the leaf and stem material ofBorassodendron borneensis during times of low fruit abundance. Other food items consumed by the apes includebark,honey, bird eggs, insects and small vertebrates includingslow lorises.[44][24]: 65–66 

In some areas, orangutans may practisegeophagy, which involves consuming soil and other earth substances. They will uproot soil from the ground as well as eatshelter tubes from tree trunks. Orangutans also visit the sides of cliffs or earth depressions for theirmineral licks. Orangutans may eat soils for their anti-toxickaolin minerals, since their diet contains toxictannins andphenolic acids.[24]: 67 

Social life

Two orangutans swinging on tree branches
Orangutans are the least social of the great apes.

The social structure of the orangutan can be best described assolitary but social; they live a more solitary lifestyle than the other great apes.[50] Bornean orangutans are generally more solitary than Sumatran orangutans.[35] Most social bonds occur between adult females and their dependent and weaned offspring. Resident females live with their offspring in defined home ranges that overlap with those of other adult females, which may be their immediate relatives. One to several resident female home ranges are encompassed within the home range of a resident male, who is their main mating partner.[50][51] Interactions between adult females range from friendly to avoidance to antagonistic.[52] Flanged males are often hostile to both other flanged males and unflanged males, while unflanged males are more peaceful towards each other.[53]

Orangutansdisperse and establish their home ranges by age 11. Females tend to live near their birth range, while males disperse farther but may still visit their birth range within their larger home range.[51][54] They enter a transient phase, which lasts until a male can challenge and displace a dominant, resident male from his home range.[55] Both resident and transient orangutans aggregate on large fruiting trees to feed. The fruits tend to be abundant, so competition is low and individuals may engage in social interactions.[56][57][58] Orangutans will also form travelling groups with members moving between different food sources.[55] They are often consortships between an adult male and a female.[56]Social grooming is uncommon among orangutans.[59]

Communication


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Orangutans communicate with various vocals and sounds. Males will make long calls, both to attract females and to advertise themselves to other males.[39] These calls have three components; they begin with grumbles, peak with pulses and end with bubbles. Both sexes will try to intimidate conspecifics with a series of low frequency noises known collectively as the "rolling call". When uncomfortable, an orangutan will produce a "kiss squeak", which involves sucking in air through pursed lips. Mothers produce throatscrapes to keep in contact with their offspring. Infants make soft hoots when distressed. When building a nest, orangutans will produce smacks orblow raspberries.[60] Orangutan calls display consonant- and vowel-like components and they maintain their meaning over great distances.[61] They also displayrecursion via three layers of rhythmic sounds.[62]

Mother orangutans and offspring also use several different gestures and expressions such as beckoning, stomping, lower lip pushing, object shaking and "presenting" a body part. These communicate goals such as "acquire object", "climb on me", "climb on you", "climb over", "move away", "play change: decrease intensity", "resume play" and "stop that".[63]

Reproduction and development

Flanged male orangutan
Unflanged male orangutan

Males become sexually mature at around age 15. They may exhibitarrested development by not developing the distinctive cheek pads, pronounced throat pouches, long fur, or long calls until a resident dominant male is absent. The transformation from unflanged to flanged can occur quickly. Flanged males attract females inoestrous with their characteristic long calls, which may also suppress development in younger males.[39][19]: 100 

Unflanged males wander widely in search of oestrous females and upon finding one, mayforce copulation on her; forced copulation by orangutans is unusually high among mammals. Females prefer to mate with the fitter flanged males, forming pairs with them and benefiting from their protection.[64][55][65] Non-ovulating females do not usually resist copulation with unflanged males, as the chance of conception is low.[65]Homosexual behaviour has been recorded in the context of both affiliative and aggressive interactions.[66]

A mother orangutan with her offspring
Mother orangutan with young

Unlike females of other non-human great ape species, orangutans do not exhibitsexual swellings to signal fertility.[65] A female first gives birth around 15 years of age and they have a six- to nine-year interbirth interval, the longest among the great apes.[67] Gestation is around nine months long and infants are born at a weight of 1.5–2 kg (3.3–4.4 lb).[19]: 99  Usually only a single infant is born; twins are a rare occurrence.[68] Unlike many other primates, male orangutans do not seem to practiseinfanticide. This may be because they cannot ensure they will sire a female's next offspring, because she does not immediately begin ovulating again after her infant dies.[69] There is evidence that females with offspring under six years old generally avoid adult males.[70]

Females do most of the caring of the young. The mother will carry the infant while travelling, suckle it and sleep with it.[19]: 100  During its first four months, the infant is almost never without physical contact and clings to its mother's belly. In the following months, the amount of physical contact the infant has with its mother declines. When an orangutan reaches the age of one-and-a-half years, its climbing skills improve and it will travel through the canopy holding hands with other orangutans, a behaviour known as "buddy travel".[59] After two years of age, juvenile orangutans will begin to move away from their mothers temporarily. They reach adolescence at six or seven years of age and are able to live alone but retain some connections with their mothers.[19]: 100  Females may nurse their offspring for up to eight years, which is more than any other mammal.[71] Typically, orangutans live over 30 years both in the wild and in captivity.[19]: 15 

Nesting

Orangutan lying on its back in a nest
An orangutan lying in its nest

Orangutansbuild nests specialised for either day or night use. These are carefully constructed; young orangutans learn from observing their mother's nest-building behaviour. In fact, nest-building allows young orangutans to become less dependent on their mother. From six months of age onwards, orangutans practise nest-building and gain proficiency by the time they are three years old.[72]

Construction of a night nest is done by following a sequence of steps. Initially, a suitable tree is located. Orangutans are choosy about sites, though nests can be found in many tree species. To establish a foundation, the ape grabs the large branches under it and bends them so they join. The orangutan then does the same to smaller, leafier branches to create a "mattress". After this, the ape stands and braids the tips of branches into the mattress. Doing this increases the stability of the nest. Orangutans make their nests more comfortable by creating "pillows", "blankets", "roofs" and "bunk-beds".[72]

Intelligence

Further information:Primate cognition
See also:Great ape language
An orangutan imitating human speech[73]

Orangutans are among the most intelligent non-human primates. Experiments suggest they cantrack the displacement of objects both visible and hidden.[74][75]Zoo Atlanta has a touch-screen computer on which their two Sumatran orangutans play games.[76] A 2008 study of two orangutans at theLeipzig Zoo showed orangutans may practise "calculatedreciprocity", which involves an individual aiding another with the expectation of being paid back. Orangutans are the first nonhuman species documented to do so.[77]

In a 1997 study, two captive adult orangutans were tested with thecooperative pulling paradigm. Without any training, the orangutans succeeded in pulling off an object to get food in the first session. Over the course of 30 sessions, the apes succeeded more quickly, having learned to coordinate.[78] An adult orangutan has been documented to pass themirror test, indicatingself-awareness,[79] while another test with a 2-year-old failed to reveal self-recognition.[80]

Studies in the wild indicate that flanged male orangutans plan their movements in advance and signal them to other individuals.[81] Experiments have also suggested that orangutans cancommunicate about things that are not present: mother orangutans remain silent in the presence of a perceived threat but when it passes, the mother produces an alarm call to their offspring to teach them about the danger.[82] Orangutans and other great apes showlaughter-like vocalisations in response to physical contact such as wrestling, play chasing or tickling. This suggests that laughter derived from a common origin among primate species and therefore evolved before the origin of humans.[83] Orangutans can learn to mimic new sounds by purposely controlling the vibrations of their vocal folds, a trait that led to speech in humans.[73][84]Bonnie, an orangutan at theUS National Zoo, was recorded spontaneously whistling after hearing a caretaker. She appears to whistle without expecting a food reward.[85]

Tool use and culture

An orangutan using a stick to pick at a hole in a rock with a cup of orange-juice concentrate.
An orangutan at theSan Diego Zoo using a tool to extract orange-juice concentrate

Tool use in orangutans was observed by primatologistBirutė Galdikas in ex-captive populations.[86] Orangutans in Suaq Balimbing were recorded to develop a tool kit for use in foraging which consisted of both insect-extraction sticks for use in the hollows of trees and seed-extraction sticks for harvesting seeds from hard-husked fruit. The orangutans adjusted their tools according to the task at hand, and preference was given to oral tool use.[87][88] This preference was also found in an experimental study of captive orangutans.[89] Orangutans have been observed to use sticks to poke at catfish, causing them to leap out of the water so the orangutan can grab them.[90][91] Orangutan have also been documented to keep tools for later.[92] When building a nest, orangutans appear to be able to determine which branches would better support their body weight.[93]

PrimatologistCarel P. van Schaik and biological anthropologist Cheryl D. Knott further investigated tool use in different wild orangutan populations. They compared geographic variations in tool use related to the processing ofNeesia fruit. The orangutans of Suaq Balimbing were found to be avid users of insect and seed-extraction tools when compared to other wild orangutans.[94][95] The scientists suggested these differences are cultural as they do not correlate with habitat. The orangutans at Suaq Balimbing are closely spaced and relatively tolerant of each other; this creates favourable conditions for the spreading of new behaviours.[94] Further evidence that highly social orangutans are more likely to exhibit cultural behaviours came from a study of leaf-carrying behaviours of formerly captive orangutans that were being rehabilitated on the island of Kaja in Borneo.[96]

Wild orangutans in Tuanan, Borneo, were reported to use tools in acoustic communication. They use leaves to amplify the kiss squeak sounds they produce. The apes may employ this method of amplification todeceive the listener into believing they are larger animals.[97] In 2003, researchers from six different orangutan field sites who used the same behavioural coding scheme compared the behaviours of the animals from each site. They found each orangutan population used different tools. The evidence suggested the differences were cultural: first, the extent of the differences increased with distance, suggesting cultural diffusion was occurring, and second, the size of the orangutans' cultural repertoire increased according to the amount of social contact present within the group. Social contact facilitates cultural transmission.[98]

During afield observation in 2022, a male Sumatran orangutan, known to researchers as Rakus, chewedFibraurea tinctoria vine leaves andapplied the mashed plant material to an open wound on his face.[99] According toprimatologists who had been observing Rakus at a nature preserve, "Five days later the facial wound was closed, while within a few weeks it had healed, leaving only a small scar".[99][100]

Personhood

Main articles:Great ape personhood,Great Ape Project, andGreat ape research ban

In June 2008, Spain would become the first country to recognise the rights of some non-human great apes, based on the guidelines of theGreat Ape Project, which are that chimpanzees,bonobos, orangutans, and gorillas are not to be used for animal experiments.[101][102] In December 2014, a court in Argentina ruled that an orangutan named Sandra at theBuenos Aires Zoo must be moved to a sanctuary in Brazil to provide her "partial or controlled freedom". Sandra has since been relocated toThe Center for Great Apes in theUnited States, as it is the only accredited orangutansanctuary in theAmericas.[103] Animal rights groups like Great Ape Project Argentina argued the ruling should apply to all species in captivity, and legal specialists from the Argentina's Federal Chamber ofCriminal Cassatio considered the ruling applicable only to non-human hominids.[104]

Orangutans and humans

Birutė Galdikas shown speaking into a microphone
Orangutan researcherBirutė Galdikas presenting her book about the apes

Orangutans were known to the native people of Sumatra and Borneo for millennia. The apes are known asmaias inSarawak andmawas in other parts of Borneo and in Sumatra.[12] While some communities hunted them for food and decoration, others placed taboos on such practices. In central Borneo, some traditional folk beliefs consider it bad luck to look an orangutan in the face. Some folk tales involve orangutans mating with and kidnapping humans. There are even stories of hunters being captured by female orangutans.[19]: 66–71 

Europeans became aware of the existence of the orangutan in the 17th century.[19]: 60  Explorers in Borneo hunted them extensively during the 19th century. In 1779, Dutch anatomistPetrus Camper, who observed the animals and dissected some specimens, gave the first scientific description of the orangutan.[19]: 64–65  Camper mistakenly thought that flanged and unflanged male orangutans were different species, a misconception corrected after his death.[105]

Little was known about orangutan behaviour until the field studies of Birutė Galdikas,[106] who became a leading authority on the apes. When she arrived in Borneo in 1971, Galdikas settled into a primitive bark-and-thatch hut at a site she dubbed Camp Leakey, inTanjung Puting. She studied orangutans for the next four years and developed herPhD thesis forUCLA.[107] Galdikas became an outspoken advocate for orangutans and the preservation of their rainforest habitat, which is rapidly being devastated byloggers,palm oil plantations,gold miners, and unnaturalforest fires.[108] Along withJane Goodall andDian Fossey, Galdikas is considered to be one ofLeakey's Angels, named after anthropologistLouis Leakey.[109]

In fiction

Main article:Orangutans in popular culture
Illustration of an orangutan attacking a woman from The Murders in the Rue Morgue by Daniel Vierge
1870 illustration for "The Murders in the Rue Morgue" byDaniel Vierge

Orangutans first appeared in Western fiction in the 18th century and have been used to comment on human society. Written by the pseudonymous A. Ardra,Tintinnabulum naturae (The Bell of Nature, 1772) is told from the point of view of a human-orangutan hybrid who calls himself the "metaphysician of the woods". Around 50 years later, the anonymously written workThe Orang Outang is narrated by a pure orangutan in captivity in the US, writing a letter critiquing Boston society to her friend in Java.[5]: 108–09 

Thomas Love Peacock's 1817 novelMelincourt features Sir Oran Haut Ton, an orangutan who lives among English people and becomes a candidate forMember of Parliament. The novel satirises the class and political system of Britain. Oran's purity and status as a "natural man" stands in contrast to the immorality and corruption of the "civilised" humans.[5]: 110–11  InFrank Challice Constable'sThe Curse of Intellect (1895), the protagonist Reuben Power travels to Borneo and captures an orangutan to train it to speak so he can "know what a beast like that might think of us".[5]: 114–15  Orangutans are featured prominently in the 1963 science fiction novelPlanet of the Apes byPierre Boulle and themedia franchise derived from it. They are typically portrayed asbureaucrats likeDr. Zaius, the science minister.[5]: 118–19, 175–76 

Orangutans are sometimes portrayed as antagonists, notably in the 1832Walter Scott novelCount Robert of Paris and the 1841Edgar Allan Poe short storyThe Murders in the Rue Morgue.[5]: 145 Disney's 1967animated musical adaptation ofThe Jungle Book added a jazzy orangutan namedKing Louie, who tries to getMowgli to teach him how to make fire.[5]: 266  The 1986 horror filmLink features an intelligent orangutan which serves a university professor but has sinister motives; he plots against humanity and stalks a female student assistant.[5]: 174–75  Other stories have portrayed orangutans helping humans, such asThe Librarian inTerry Pratchett's fantasy novelsDiscworld and inDale Smith's 2004 novelWhat the Orangutan Told Alice.[5]: 123  More comical portrayals of the orangutan include the 1996 filmDunston Checks In.[5]: 181 

In captivity

Sketch of the female orangutan known as Jenny sitting in a chair
The Female Orang – Utan (Jenny sitting in a chair)[110]c. 1830s

By the early 19th century, orangutans were being kept in captivity. In 1817, an orangutan joined several other animals in London'sExeter Exchange. He rejected the company of other animals, aside from a dog, and preferred to be with humans. He was occasionally taken on coach rides clothed in asmock-frock and hat and even given drinks at an inn where he behaved politely for the hosts.[5]: 64–65  TheLondon Zoo housed a female orangutan namedJenny who was dressed in human clothing and learned to drink tea. She is remembered for her meeting withCharles Darwin who compared her reactions to those of a human child.[110][111]

Zoos and circuses in the Western world would continue to use orangutans and othersimians as sources for entertainment, training them to behave like humans attea parties and to perform tricks. Notable orangutan "character actors" include: Jacob and Rosa of theTierpark Hagenbeck in Hamburg, Germany, in the early 20th century;Joe Martin ofUniversal City Zoo in the 1910s and 1920s; andJiggs of theSan Diego Zoo in the 1930s and 1940s.[5]: 187–89, 193–94 [112]Animal rights groups have urged a stop to such acts, considering them abusive.[113] Starting in the 1960s, zoos became more concerned with education and orangutans' exhibits were designed to mimic their natural environment and let them display their natural behaviours.[5]: 185, 206 Ken Allen, an orangutan of the San Diego Zoo, became world famous in the 1980s for multiple escapes from his enclosures. He was nicknamed "thehairy Houdini" and was the subject of a fan club, T-shirts, bumper stickers and a song titledThe Ballad of Ken Allen.[114]

Galdikas reported that her cook was sexually assaulted by a captive male orangutan.[115] The ape may have suffered from a skewed species identity and forced copulation is a standard mating strategy for low-ranking male orangutans.[116] American animal traffickerFrank Buck claimed to have seen human mothers acting aswet nurses to orphaned orangutan babies in hopes of keeping them alive long enough to sell to a trader, which would be an instance ofhuman–animal breastfeeding.[117]

Conservation

Status and threats

Main article:Endangerment of orangutans

All three species are critically endangered according to theIUCN Red List of mammals.[118][119][120] They are legally protected from capture, harm or killing in both Malaysia and Indonesia,[121] and are listed underAppendix I byCITES, which prohibits their unlicensed trade under international law.[122] The Bornean orangutan range has become more fragmented, with few or no apes documented in the southeast.[119] The largest remaining population is found in the forest around theSabangau River, but this environment is at risk.[123] The Sumatran orangutan is found only in the northern part of Sumatra, most of the population inhabiting theLeuser Ecosystem.[118] The Tapanuli orangutan is found only in theBatang Toru forest of Sumatra.[120]

A view of deforested land
Deforestation forpalm oil production in Indonesia

Birutė Galdikas wrote that orangutans were already threatened bypoaching anddeforestation when she began studying them in 1971.[124] By the 2000s, orangutan habitats decreased rapidly because of logging, mining andfragmentation by roads. A major factor has been the conversion of vast areas oftropical forest topalm oil plantations in response to international demand. Hunting is also a major problem, as is the illegalpet trade.[118][119]

Orangutans may be killed for thebushmeat trade[125] and bones are secretly sold in souvenir shops in several cities in Indonesian Borneo.[126] Conflicts between locals and orangutans also pose a threat. Orangutans that have lost their homes often raid agricultural areas and end up being killed by villagers.[127] Locals may also be motivated to kill orangutans for food or because of their perceived danger.[128] Mother orangutans are killed so their infants can be sold as pets. Between 2012 and 2017, the Indonesian authorities, with the aid of the Orangutan Information Center, seized 114 orangutans, 39 of which were pets.[129]

Estimates in the 2000s found that around 6,500 Sumatran orangutans and around 54,000 Bornean orangutans remain in the wild.[130] A 2016 study estimates a population of 14,613 Sumatran orangutans in the wild, twice that of previous population estimates,[131] while 2016 estimates suggest 104,700 Bornean orangutans exist in the wild.[119] A 2018 study found that Bornean orangutans declined by 148,500 individuals from 1999 to 2015.[132] Fewer than 800 Tapanuli orangutans are estimated to still exist, which puts the species among the most endangered of the great apes.[133][43]

Conservation centres and organisations

Peter Pratje with an orangutan
Zoologische Gesellschaft Frankfurt Programm Director Peter Pratje works with orangutans inBukit Tigapuluh, Indonesia.

Several organisations are working for the rescue, rehabilitation and reintroduction of orangutans. The largest of these is theBorneo Orangutan Survival (BOS) Foundation, founded by conservationistWillie Smits and which operates projects such as the Nyaru Menteng Rehabilitation Program founded by conservationistLone Drøscher Nielsen.[134][135][136]

A female orangutan was rescued from a village brothel in Kareng Pangi village, Central Kalimantan, in 2003. The orangutan was shaved and chained for sexual purposes. Since being freed, the orangutan, named Pony, has been living with the BOS. She has been re-socialised to live with other orangutans.[137] In May 2017, the BOS rescued an albino orangutan from captivity in a remote village in Kapuas Hulu, on the island of Kalimantan in Indonesian Borneo. According to volunteers at BOS, albino orangutans are extremely rare (one in ten thousand). This is the first albino orangutan the organisation has seen in 25 years of activity.[138]

Other major conservation centres in Indonesia include those atTanjung Puting National Park,Sebangau National Park,Gunung Palung National Park andBukit Baka Bukit Raya National Park in Borneo and theGunung Leuser National Park andBukit Lawang in Sumatra. In Malaysia, conservation areas include Semenggoh Wildlife Centre and Matang Wildlife Centre also in Sarawak, and theSepilok Orang Utan Sanctuary in Sabah.[139] Major conservation centres headquartered outside the orangutans' home countries includeFrankfurt Zoological Society,[140]Orangutan Foundation International, which was founded by Galdikas,[141] and theAustralian Orangutan Project.[142] Conservation organisations such as theOrangutan Land Trust work with the palm oil industry to improvesustainability and encourages the industry to establishconservation areas for orangutans.[143][144]

See also

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Look uporangutan in Wiktionary, the free dictionary.
Extant species of familyHominidae(great apes)
Hominidae
Ponginae
Pongo
(Orangutans)
Homininae
Gorilla
(Gorillas)
Hominini
Pan
Homo
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Extant
ape species
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Notable non-humanapes
Bonobos
Chimpanzees
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Orangutans
Microchoerinae
"Anaptomorphinae"
"Omomyinae"
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Eosimiidae
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Eosimias sinensis
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Pliopithecoidea
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Aegyptopithecus zeuxis
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