The orangutans (Malay "orang" (man) and "(h)utan" (forest); hence, "man of the forest") are the two exclusively Asian species of extant great apes. Native to Indonesia and Malaysia, orangutans are currently found only in rainforests on the islands of Borneo and Sumatra. Classified in the genus Pongo, there are only two surviving species: the Bornean orangutan (P. pygmaeus) and the Sumatran orangutan (P. abelii). They are also the only surviving species of the subfamily Ponginae, which also included several other species like Gigantopithecus.
The largest living arboreal animals, they have proportionally longer arms than the other, more terrestrial, great apes. Their hair is typically reddish-brown, instead of the brown or black hair typical of their relatives. Dominant adult males have distinctive cheek pads. Orangutans are the more solitary of the great apes, with most social bonds occurring between mothers and their dependent offspring. Fruit is the most important component of a orangutans diet, however, the apes will also eat vegetation, bark, honey, insects and even bird eggs. They are among the most intelligent primates and use a variety of sophisticated tools, also making sleeping nests each night from branches and foliage. The apes have been extensively studied for their and learning abilities. Their may even be distinctive cultures within populations.
Both orangutan species are considered to be Endangered with the Sumatran orangutan being Critically Endangered. Threats to orangutan populations include poaching, habitat destruction and the illegal pet trade.
Taxonomy and phylogeny
The two orangutan species are the only extant members of the subfamily Ponginae. This subfamily also included the extinct genera Lufengpithecus, which lived in southern China and Thailand 2–8 million years ago (mya), and Sivapithecus, which lived India and Pakistan 12.5–8.5 mya.:50 These apes likely lived in drier and cooler environments then orangutans do today. It is Khoratpithecus piriyai, which lived in Thailand 7–5 mya, that is believed to have been the closest known relative of the orangutans.:50 Another notable member of Ponginae was Gigantopithecus, which lived in China, India and Vietnam 5 million to 100,000 years ago.:50 It was the largest known primate. It is likely that great ape evolution was centered in Africa and the lineage that lead to the Asian apes split off around 20 million years ago (mya), and orangutans arose in that lineage around 11–14 mya.:50
- Genus Pongo
The populations on the two islands were classified as subspecies until recently, when they were elevated to full specific level, and the three distinct populations on Borneo were elevated to subspecies. The population currently listed as P. p. wurmbii may be closer to the Sumatran orangutan than the other Bornean orangutan subspecies. If confirmed, abelii would be a subspecies of P. wurmbii (Tiedeman, 1808). Regardless, the type locality of pygmaeus has not been established beyond doubts, and may be from the population currently listed as wurmbii (in which case wurmbii would be a junior synonym of pygmaeus, while one of the names currently considered a junior synonym of pygmaeus would take precedence for the northwest Bornean taxon). To further confuse, the name morio, as well as various junior synonyms that have been suggested, have been considered likely to all be junior synonyms of the population listed as pygmaeus in the above, thus leaving the east Bornean populations unnamed.
In addition, some fossils described under the name P. 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 to P. pygmaeus or P. abelii or, in fact, represent distinct species.
Orangutans have 48 diploid chromosomes, and its genome was sequenced in January 2011. Following humans and chimpanzees, the Sumatran orangutan has become the third species of hominid to have its genome sequenced. The draft of the genome sequence is based on a captive female named Susie.
The researchers also published less complete copies from ten wild orangutans, five from Borneo and five from Sumatra. It was found that genetic diversity was lower in Bornean orangutans (Pongo pygmaeus) than in Sumatran ones (Pongo abelii), despite the fact that Borneo is home to six or seven times as many orangutans as Sumatra. The comparison has shown that these two species diverged around 400,000 years ago, more recently than was previously thought. It was also found that the orangutan genome has evolved much more slowly than chimpanzee and human DNA.
Anatomy and physiology
An orangutan has a large, bulky body, a thick neck, very long, strong arms, short, bowed legs, and no tail. It is mostly covered with long reddish-brown hair and grey-black skin. Sumatran orangutans have the more sparse and lighter coloured coat. Though largely hairless, the face of orangutan can develop some hair in males, giving them a moustache. There is significant sexual dimorphism: females can grow to around 4 ft 2 in or 127 cm and weigh around 100 lb (45 kg) while flanged adult males can reach 5 ft 9 in or 175 cm in height and weigh over 260 lb (118 kg).
Orangutan hands are similar to humans' hands; they have four long fingers and an opposable thumb. However, the joint and tendon arrangement in the orangutans' hands produces two adaptations that are significant in their success as arboreal species. The resting configuration of the fingers is curved, creating a suspensory hook grip.:301 Additionally, when gripping tightly, an orangutans fingers and hands can, without the use of the thumb, wrap completely around an object with a small enough diameter, with the tops of the fingers resting parallel against the inside of the palm, creating a double-locked grip.:301
Their feet have four long toes and an opposable big toe.:15 Orangutans can grasp things with both their hands and their feet. A male orangutans has an arm span of about 2 m (6.6 ft).:14 Their fingers and toes are curved, allowing them to better grip onto branches. Orangutans have less restriction in the movements of their legs than humans since their hips joint have the same flexibility as their shoulder and arm joints.:15 Unlike gorillas and chimpanzees, orangutans are not true knuckle-walkers, and are instead fist-walkers.
The orangutan has a large head with a prominent mouth area. Adult males have large cheek flaps:14 that show their dominance to other males. The cheek flaps are made mostly of fat and are supported but the musculature of the face. The age of maturity for females is approximately 12 years. Typically, orangutans live over 30 years in both the wild and captivity.:14 Mature males have throat pouches that allows them to make loud calls.:14
Ecology and behaviour
Orangutans live in primary and old secondary forests, particularly dipterocarp forests and peat swamp forests. Both species can be found in both mountainous and lowland swampy areas. Sumatran orangutans live in elevations as high as 1500 m (4921 ft), while Bornean orangutans live no higher than 1000 m (3281 ft). Other habitats used by orangutans include grasslands, cultivated fields, gardens, young secondary forest, and shallow lakes. Orangutans are the most arboreal of the great apes, spending nearly all of their time in the trees. Most of the day is spent feeding, resting, and travelling. They start the day feeding for 2–3 hours in the morning. They rest during midday then travel in the late afternoon. When evening arrives, they begin to prepare their nest for the night. Orangutans do not swim, although they have been recorded wading in water.
Tigers are likely to be the biggest predatory threat to orangutans. Other potential predators include clouded leopards, wild dogs and crocodiles. The absence of tigers on Borneo may explain why Bornean orangutans can be found on the ground more often than their Sumatran relatives. Orangutans communicate with various vocalizations. Male will make long calls, both to attract females and advertise themselves to other males. Both sexes will try to intimidate conspecifics with a series of low guttural noises known collectively as the "rolling call". When annoyed, an orangutan will suck in air through pursed lips, which makes a kissing sound and is hence known as the "kiss squeak". Infants make soft hoots when distressed. Orangutans are also known to blow a raspberry.
Orangutans are opportunistic foragers, and their diets vary markedly from month to month. Fruit makes up 65–90 percent of the orangutan diet, and those with sugary or fatty pulp are favoured. Ficus fruits are commonly eaten, because they are easy to harvest and digest. Lowland dipterocarp forests are preferred by orangutans because of their plentiful fruit. Bornean orangutans consume at least 317 different food items that include young leaves, shoots, bark, insects, honey and bird eggs. Bark is eaten as a last resort in times of food scarcity; fruits are always more popular.
A decade-long study of urine and faecal samples at the Gunung Palung Orangutan Conservation Project in West Kalimantan has shown that orangutans give birth during and after the high fruit season (though not every year), during which they consume various abundant fruits, totaling up to 11,000 calories per day. In the low fruit season they eat whatever fruit is available in addition to tree bark and leaves, with daily intake at only 2,000 calories. Together with a long lactation period, orangutans also have a long birth interval.
Orangutans are thought to be the sole fruit disperser for some plant species including the climber species Strychnos ignatii which contains the toxic alkaloid strychnine. It does not appear to have any effect on orangutans except for excessive saliva production.
Geophagy, the practice of eating soil or rock, has been observed in orangutans. There are three main reasons for this dietary behaviour; for the addition of mineral nutrients to their diet; for the ingestion of clay minerals that can absorb toxic substances; or to treat a disorder such as diarrhoea.
Orangutans live a more solitary lifestyle than the other great apes. Most social bonds occur between adult females and their dependent and weaned offspring. Adult males and independent adolescents of both sexes tend to live alone. The society of the orangutan is made up of resident and transient individuals of both sexes. Resident females live with their offspring in defined home ranges that overlap with those of other adult females, who may be their relatives like mothers and sisters. One to several resident female home ranges are encompassed within the home range of a resident male, who is their main mating partner. Transient males and females move widely. Orangutans usually travel alone, but they may travel in small groups in their sub-adult years. However this behaviour ends at adulthood. The social structure of the orangutan can be best described as solitary but social. Interactions between adult females range from friendly, to avoidance to antagonistic. Resident males may have overlapping ranges and interactions between them tend to be hostile.
During dispersal, females tend to settle in home ranges that overlap with their mothers. However, they do not seem to have any special social bonds with them. Males disperse much farther from their mothers and enter into a transient phase. This phase lasts until a male can challenge and displace a dominant, resident male from his home range. Adult males dominate sub-adult males. 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 intereactions. Orangutans will also form traveling groups with members moving between different food sources. These groups tend to be made of only a few individuals. They also tend to be consortships between an adult male and female.
Reproduction and parenting
Male orangutans exhibit arrested development. They mature at around 15 years of age by which time they have fully descended testicles and can reproduce. However they do not develop the distinctive cheek pads, pronounced throat pouches, long fur or long-calls until they are between 15 and 20 years old. The development of thse characteristics depends largely on the absence of a resident male. Males without them are known as unflanged males in contrast to the more developed flanged males. The transformation from unflanged to flanged can occur very quickly. Unflanged and flanged males have two different mating strategies. Flanged males attract oestrous females with their characteristic long calls. Those calls may also suppress development in younger males. Unflanged males wander widely in search of estrous females and upon finding one, will force copulation on her. Both strategies are successful, however females prefer to mate with flanged males and seek their company for protection against unflanged males. Resident males may form consortships with females that can last days, weeks or months after copulation.
Female orangutans experience their first ovulatory cycle around 5.8–11.1 years. These occur earlier in females with more body fat. Like other great apes, female orangutans enter a period of infertility during adolescence which may last for 1–4 years. Female orangutans also have a 22–30 day menstrual cycle. Gestation lasts for nine months with females giving birth to their first offspring between 14 and 15 years old. Female orangutans have eight year intervals between births; the longest interbirth intervals among the great apes. Unlike many other primates, male orangutans do not seem to practice infanticide. This may be because they cannot ensure that they will sire a female's next offspring because she does not immediately begin ovulating again after her infant dies.
Male orangutans play almost no role in raising the young. Females do most of the caring and socializing of the young. A female often has an older offspring with her to help in socializing the infant. Infant orangutans are completely dependent on their mothers for the first two years of their lives. The mother will carry the infant during traveling, as well as feed it and sleep with it in the same night nest. For the first four months, the infant is carried on its belly and never relieves physical contact. In the following months, the time a infant spends with its mother decreases. When an orangutan reaches the age of two, its climbing skills improve and will travel through the canopy holding hands with other orangutans, a behaviour known as "buddy travel". Orangutans are juveniles for about two to five years of age and will start to temporarily move away from their mothers. Juveniles are usually weaned at about four years of age. Adolescent orangutans will socialize with their peers while still having contact with their mothers.
Tool use and culture
Evidence of sophisticated tool manufacture and use in the wild was reported from a population of orangutans in Suaq Balimbing (Pongo abelii) in 1996. These orangutans developed a tool kit for use in foraging that consisted of insect-extraction tools for use in the hollows of trees, and seed-extraction tools which were used in harvesting seeds from hard-husked fruit. The orangutans adjusted their tools according to the nature of the task at hand and preference was given to oral tool use. This preference was also found in an experimental study of captive orangutans (P. pygmaeus).
Carel P. van Schaik from the University of Zurich and Cheryl D. Knott from Harvard University further investigated tool use in different wild orangutan populations. They compared geographic variations in tool use related to the processing of Neesia fruit. The orangutans of Suaq Balimbing (P. abelii) were found to be avid users of insect and seed-extraction tools when compared to other wild orangutans. The scientists suggested that these differences are cultural. The orangutans at Suaq Balimbing live in dense groups and are socially tolerant; this creates good conditions for social transmission. Further evidence that highly social orangutans are more likely to exhibit cultural behaviours came from a study of leaf-carrying behaviours of ex-captive orangutans that were being rehabilitated on the island of Kaja in Borneo.
In 2003, researchers from six different orangutan field sites who used the same behavioural coding scheme compared the behaviours of the animals from the different sites. They found that the different orangutan populations behaved differently. The evidence suggested that the differences in behaviour were cultural: first, because the extent of the differences increased with distance, suggesting that cultural diffusion was occurring, and second, because the size of the orangutans’ cultural repertoire increased according to the amount of social contact present within the group. Social contact facilitates cultural transmission.
Orangutans build day as well as night nests. These are carefully constructed; young orangutans learn from observing their mother's nest-building behaviour. In fact, nest-building is a leading cause in young orangutans leaving their mother for the first time. From six months of age onwards, orangutans practice nest building and gain proficiency by the time they are three years old.
Construction of a night nest is done by following a sequence of steps. Initially a suitable tree is located, orangutans being selective about sites even though many tree species are utilised. The foundation is then built by pulling together branches under them and joining them at a point. After the foundation has been built, the orangutan bends smaller, leafy branches onto the foundation; this serves the purpose of and is termed as the "mattress". After this orangutans stand and braid the tips of branches into the mattress. This increases the stability of the nest and forms the final act of nest building. In addition, orangutans may add additional features such as "pillows", "blankets", "roofs" and "bunk-beds" to their nest.
Orangutans do not limit their tool use to foraging, displaying or nest-building activities. Wild orangutans (P. pygmaeus wurmbii) in Tuanan, Borneo, were reported to use tools in acoustic communication. They use leaves to amplify the kiss squeak sounds that they produce. Some have suggested that the apes employ this method of amplification in order to deceive the listener into believing that they are larger animals.
A two year study of orangutan symbolic capability was conducted from 1973-1975 by Gary L. Shapiro with Aazk, a juvenile female orangutan at the Fresno City Zoo (now Chaffee Zoo) in Fresno, California. The study employed the techniques of David Premack who used plastic tokens to teach the chimpanzee, Sarah, linguistic skills. Shapiro continued to examine the linguistic and learning abilities of ex-captive orangutans in Tanjung Puting National Park, in Indonesian Borneo, between 1978 and 1980. During that time, Shapiro instructed ex-captive orangutans in the acquisition and use of signs following the techniques of R. Allen and Beatrix Gardner who taught the chimpanzee, Washoe, in the late-1960s. In the only signing study ever conducted in a great ape's natural environment, Shapiro home-reared Princess, a juvenile female who learned nearly 40 signs (according to the criteria of sign acquisition used by Francine Patterson with Koko, the gorilla) and trained Rinnie, a free-ranging adult female orangutan who learned nearly 30 signs over a two year period. For his dissertation study, Shapiro examined the factors influencing sign learning by four juvenile orangutans over a 15-month period.
The first orangutan language study program, directed by Dr. Francine Neago, was listed by Encyclopædia Britannica in 1988. The Orangutan language project at the Smithsonian National Zoo in Washington, D.C., uses a computer system originally developed at UCLA by Neago in conjunction with IBM.
Zoo Atlanta has a touch screen computer where their two Sumatran orangutans play games. Scientists hope that the data they collect from this will help researchers learn about socializing patterns, such as whether they mimic others or learn behaviour from trial and error, and hope the data can point to new conservation strategies.
A 2008 study of two orangutans at the Leipzig Zoo showed that orangutans are the first non-human species documented to use 'calculated reciprocity' which involves weighing the costs and benefits of gift exchanges and keeping track of these over time.
Orangutans and humans
Orangutans were known to the native people of Sumatra and Borneo for millennia. While some communities hunted them for food and decoration, others placed taboos on such practices.:66 In central Borneo, some traditional folk beliefs consider it bad luck to look in the face of an orangutan. Some folk tales involve orangutans mating with and kidnapping humans.:68 There are even stories of hunters being seduced by female orangutans.:71 Europeans became aware of the existence of the orangutan possibly as early as the 17 century.:68 The first accurate description of orangutans was given by Dutch anatomist Petrus Camper, who observed the animals and dissected some specimens.:64 However, little was known about orangutans until the field studies of Birutė Galdikas.
Male orangutans have been reported to display sexual attraction to human women to the point of rape. The cook of noted primatologist Birutė Galdikas was raped by an orangutan. According to reporter Carole Jahme, rape "was almost the fate of Julia Roberts" when she was seized by a male orangutan before being freed by the film crew.
The Sumatran species is critically endangered and the Bornean species of orangutans is endangered according to the IUCN Red List of mammals, and both are listed on Appendix I of CITES. The total number of Bornean orangutans is estimated to be less than 14% of what it was in the recent past (from around 10,000 years ago until the middle of the twentieth century) and this sharp decline has occurred mostly over the past few decades due to human activities and development. Species distribution is now highly patchy throughout Borneo: it is apparently absent or uncommon in the south-east of the island, as well as in the forests between the Rejang River in central Sarawak and the Padas River in western Sabah (including the Sultanate of Brunei). The largest remaining population is found in the forest around the Sabangau River, but this environment is at risk. A similar development have been observed for the Sumatran orangutans.
A 2007 study by the Government of Indonesia noted in 2004 it was estimated that there was a total wild population of 61,234 orangutans, 54,567 of which were found on the island of Borneo. The table below shows a breakdown of the species and subspecies and their estimated populations from the report:
|Pongo abelii||Sumatran orangutan||Sumatra||6,667|
|Pongo pygmaeus||Bornean orangutan||Borneo|
|P. p. morio||Northeast Bornean orangutan||Sabah||11,017|
|P. p. morio||Northeast Bornean orangutan||East Kalimantan||4,825|
|P. p. wurmbii||Central Bornean orangutan||Central Kalimantan||>31,300|
|P. p. pygmaeus||Northwest Bornean orangutan||West Kalimantan and Sarawak||7,425|
This indicates a decline from some estimates between 2000 and 2003 which found 7,300 Sumatran orangutan individuals in the wild and between 45,000 and 69,000 Bornean orangutans. Thousands of orangutans don't reach adulthood due to human disruption. Orangutans are killed for food while others are killed because of disruption in people's property. Mother orangutans are killed so their infants can be sold as pets. Many of the infants die without the help of their mother. Since recent trends are steeply down in most places due to logging and burning, it is forecast that the current numbers are below these figures.
Orangutan habitat destruction due to logging, mining and forest fires, as well as fragmentation by roads, has been increasing rapidly in the last decade. A major factor in that period of time has been the conversion of vast areas of tropical forest to oil palm plantations in response to international demand (the palm oil is used for cooking, cosmetics, mechanics, and more recently as source of biodiesel). Some UN scientists believe that these plantations could lead to irreparable damage to orangutan habitat by the year 2012. Some of this activity is illegal, occurring in national parks that are officially off limits to loggers, miners and plantation development. There is also a major problem with hunting and illegal pet trade. In early 2004 about 100 individuals of Bornean origin were confiscated in Thailand and 50 of them were returned to Kalimantan in 2006. Several hundred Bornean orangutan orphans who were confiscated by local authorities have been entrusted to different orphanages in both Malaysia and Indonesia. They are in the process of being rehabilitated into the wild. Besides preserved animal and bones of Orangutan, Orangutan skulls have also trade silently in souvenir shops in several cities in Kalimantan, Indonesia with price between Rp.500,000 ($58.80) to Rp.2 million ($235) per skull.
Conservation centres and organisations
A number of organisations are working for the rescue, rehabilitation and reintroduction of orangutans. The largest of these is the Borneo Orangutan Survival Foundation, founded by Dr. Willie Smits. Funded in part by the Australian Orangutan Project, it employs between six hundred and a thousand people at a hundred sites. It operates a number of large projects, including the Samboja Lestari Forest Rehabilitation Program and the Nyaru Menteng Rehabilitation Program managed by Lone Drøscher Nielsen. Other major conservation centres in Indonesia include those at Tanjung Puting National Park and Sebangau National Park in Central Kalimantan, Kutai in East Kalimantan, Gunung Palung National Park in West Kalimantan, and Bukit Lawang in the Gunung Leuser National Park on the border of Aceh and North Sumatra. In Malaysia, conservation areas include Semenggoh Wildlife Centre in Sarawak and Matang Wildlife Centre also in Sarawak, and the Sepilok Orang Utan Sanctuary near Sandakan in Sabah. This last is partially documented in the film Among The Great Apes With Michelle Yeoh.
- ^ The Orangutan was originally named by Linnaeus as Simia pygmaeus. In placing the species in this new genus, Lacépède renamed it Pongo borneo and this is the name that applies to the type species; however, due to the Principle of Priority, the correct name to use when referring to it outside of this context is Pongo pygmaeus.
- ^ a b c d e f g h i j k l m n o Payne, J; Prundente, C (2008). Orangutans: Behavior, Ecology and Conservation. New Holland Publishers. ISBN 0-262-16253-9.
- ^ a b Groves, C. (2005). Wilson, D. E., & Reeder, D. M, eds. ed. Mammal Species of the World (3rd ed.). Baltimore: Johns Hopkins University Press. pp. 183–184. OCLC 62265494. ISBN 0-801-88221-4. http://www.bucknell.edu/msw3/browse.asp?id=12100803.
- ^ a b c Bradon-Jones, D.; Eudey, A. A.; Geissmann, T.; Groves, C. P.; Melnick, D. J.; Morales, J. C.; Shekelle, M.; Stewart, C. B. (2004). "Asian primate classification" (PDF). International Journal of Primatology 25: 97–164. doi:10.1023/B:IJOP.0000014647.18720.32. http://www.gibbons.de/main/papers/pdf_files/2004asianprimates.pdf.
- ^ Schwartz, J.H.; Vu The Long; Nguyen Lan Cuong, Le Trung Kha; Tattersall, I. 1995: A review of the Pleistocene hominoid fauna of the Socialist Republic of Vietnam (excluding Hylobatidae). Anthropological papers of the American Museum of Natural History, (76): 1-24. Abstract & pdf
- ^ Sharshov, Alexander. "New Page 1". SB RAS Novobrisk. Institute of Cytology and Genetics. http://www.bionet.nsc.ru/labs/chromosomes/maps/orangutan.htm. Retrieved 28 January 2011.
- ^ Locke, D. P.; Hillier, L. W.; Warren, W. C.; Worley, K. C.; Nazareth, L. V.; Muzny, D. M.; Yang, S. P.; Wang, Z. et al (2011). "Comparative and demographic analysis of orang-utan genomes". Nature 469 (7331): 529–533. doi:10.1038/nature09687. PMC 3060778. PMID 21270892. http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=3060778.
- ^ a b c d Singh, Ranjeet (2011-01-04). "Orang-utans join the genome gang". Nature. doi:10.1038/news.2011.50. http://www.nature.com/news/2011/110126/full/news.2011.50.html. Retrieved 2011-01-27.
- ^ "Orangutan Anatomy Page". Red-ape.co.uk. http://www.red-ape.co.uk/orangutan/orang_anatomy.html. Retrieved 2009-07-03.
- ^ a b c d e f g Cawthon Lang KA (2005-06-13). "Primate Factsheets: Orangutan (Pongo) Taxonomy, Morphology, & Ecology". http://pin.primate.wisc.edu/factsheets/entry/orangutan. Retrieved 2007-10-12.
- ^ "Sumatran Orangutan Society". Archived from the original on 2006-11-14. http://web.archive.org/web/20061114233340/http://www.orangutans-sos.org/faq.php. Retrieved 2007-04-01.
- ^ a b Schwartz, Jeffrey (1988). Orang-utan Biology. USA: Oxford University Press. ISBN 0-19-504371-5.
- ^ Schwartz, Jeffrey (1987). The Red Ape: Orangutans and Human Origins. Cambridge, MA: Westview Press. p. 286. ISBN 0-8133-4064-0.
- ^ Winkler, L. A. (1989). "Morphology and relationships of the orangutan fatty cheek pads". American Journal of Primatology 17 (4): 305–19. doi:10.1002/ajp.1350170405.
- ^ Galdikas, BMF (1988). "Orangutan diet, range, and activity at Tanjung Puting, Central Borneo". J Mammal 9 (1): 1–35. doi:10.1007/BF02740195.
- ^ Kuliukas, A. (2001). Bipedal Wading in Hominoidae past and present (Master of Science thesis). University College London. http://www.riverapes.com/Me/Work/Thesis.PDF.
- ^ van Schaik, C.; MacKinnon, J. (2001). "Orangutans". In MacDonald, D. The Encyclopedia of Mammals (2nd ed.). Oxford University Press. pp. 420–23. ISBN 0-87196-871-1.
- ^ a b c d Utami, SS; Goossens, B; Bruford, MW; de Ruiter, JR; van Hooff, JARAM (2002). "Male bimaturism and reproductive success in Sumatran orangutans". Behav Ecol 13 (5): 643–52. doi:10.1093/beheco/13.5.643.
- ^ a b "Vocal Repertoire". Orang Utan Republik Foundation. http://www.orangutanrepublik.org/become-aware/orangutan-specifics/vocal-repertoire#. Retrieved March 17, 2012.
- ^ a b Galdikas, Birute M.F. (1988). "Orangutan Diet, Range, and Activity at Tanjung Puting, Central Borneo". International Journal of Primatology 9 (1): 1. doi:10.1007/BF02740195.
- ^ Asrianti, T. (2011). For orangutans, less food means lowered fertility. The Jakarta Post, July 4, 2011.
- ^ Rijksen, H. D. (December 1978). "A field study on Sumatran orang utans (Pongo pygmaeus abelii, Lesson 1827): Ecology, Behaviour and Conservation". The Quarterly Review of Biology 53 (4): 493. doi:10.1086/410942. JSTOR 2826733.
- ^ Highfield, Roger (1 April 2008). "Science: Monkeys were the first doctors (Telegraph.co.uk)". The Daily Telegraph (London). http://www.telegraph.co.uk/science/science-news/3338047/Science-monkeys-were-the-first-doctors.html. Retrieved 2010-05-20.
- ^ Matt Walker Wild orangutans treat pain with natural anti-inflammatory New Scientist 28 July 2008.
- ^ a b Teboekhorst, I; Schurmann, C; Sugardjito, J (1990). "Residential status and seasonal movements of wild orang-utans in the Gunung Leuser Reserve (Sumatera, Indonesia)". Animal Behaviour 39 (6): 1098–1109. doi:10.1016/S0003-3472(05)80782-1.
- ^ a b c d e f g h i j k l Cawthon Lang KA (2005-06-13). "Primate Factsheets: Orangutan (Pongo) Taxonomy, Morphology, & Ecology". http://pin.primate.wisc.edu/factsheets/entry/orangutan. Retrieved 2012-02-19.
- ^ Singleton, I; van Schaik, CP. (2002). "The social organisation of a population of Sumatran orangutans". Folia Primatol 73 (1): 1–20. doi:10.1159/000060415. PMID 12065937.
- ^ a b c Delgado, RA Jr.; van Schaik, CP. (2000). "The behavioral ecology and conservation of the orangutan (Pongo pygmaeus): a tale of two islands". Evol Anthro 9 (1): 201–18. doi:10.1002/1520-6505(2000)9:5<201::AID-EVAN2>3.0.CO;2-Y.
- ^ a b Fox, EA. (2002). "Female tactics to reduce sexual harassment in the Sumatran orangutan (Pongo pygmaeus abelii)". Behav Ecol Sociobiol 52 (2): 93–101. doi:10.1007/s00265-002-0495-x.
- ^ Schürmann, CL; van Hooff, JARAM (1986). "Reproductive strategies of the orangutan: new data and a reconsideration of existing sociosexual models". Int J Primatol 7 (3): 265–87. doi:10.1007/BF02736392.
- ^ Beaudrot LH, Kahlenberg SM, Marshall AJ. (2009) "Why male orangutans do not kill infants," Behav Ecol Sociobiol 63(11):1549-1562. PMID 19701484 Full text at PMC: 2728907
- ^ a b c Munn C, Fernandez M. (1997) Infant development. In: Orangutan species survival plan husbandry manual, Sodara C,( ed) Chicago (IL): Orangutan Species Survival Plan. p 59-66.
- ^ Deaner, RO; van Schaik, CP; Johnson, V. (2006). "Do some taxa have better domain-general cognition than others? A meta-analysis of nonhuman primate studies" (PDF). Evol Psych 4: 149–196. http://www.epjournal.net/filestore/ep04149196.pdf.
- ^ Galdikas, BMF (1982). "Orang-Utan tool use at Tanjung Putting Reserve, Central Indonesian Borneo (Kalimantan Tengah)". J Hum Evol 10: 19–33. doi:10.1016/S0047-2484(82)80028-6.
- ^ Van Schaik, CP; Fox, EA; Sitompul, AF. (1996). "Manufacture and use of tools in wild Sumatran orangutans – implications or human evolution". Naturwissenschaften 83 (4): 186–188. doi:10.1007/s001140050271. PMID 8643126.
- ^ Fox EA, Sitompul AF, Van Schaik CP. 1999. Intelligent tool use in wild Sumatran orangutans. In: Parker S, Mitchell RW and Miles HL, editors. The Mentality of Gorillas and Orangutans. Cambridge, UK : Cambridge University Press. p: 99-116
- ^ O'Malley, RC; McGrew, WC. (2000). "Oral tool use by captive orangutans (Pongo pygmaeus)". Folia Primatol 71 (5): 334–341. doi:10.1159/000021756. PMID 11093037.
- ^ a b Van Schaik, Carel P.; Knott, Cheryl D. (2001). "Geographic variation in tool use onNeesia fruits in orangutans". American Journal of Physical Anthropology 114 (4): 331–342. doi:10.1002/ajpa.1045. PMID 11275962.
- ^ Van Schaik, CP; van Noordwijk, MA; Wich, SA. (2006). "Innovation in wild Bornean orangutans (Pongo pygmaeus wurmbii)". Behaviour 143 (7): 839–876. doi:10.1163/156853906778017944.
- ^ Russon, AE; Handayani, DP; Kuncoro, P; Ferisa, A. (2007). "Orangutan leaf-carrying for nest-building: toward unraveling cultural processes". Animal Cognition 10 (2): 189–202. doi:10.1007/s10071-006-0058-z. PMID 17160669.
- ^ a b Van Schaik, CP; Ancrenaz, M; Borgen, G; Galdikas, B; Knott, CD; Singleton, I; Suzuki, A; Utami, SS et al (2003). "Orangutan cultures and the evolution of material culture". Science 299 (5603): 102–105. doi:10.1126/science.1078004. PMID 12511649.
- ^ a b Didik, Prasetyo; Ancrenaz, Marc; Morrogh-Bernard, Helen C.; Atmoko, S. Suci Utami; Wich, Serge A. & van Schaik, Carel P. (2009). "Nest building in orangutans". In Wich, Serge A.; Atmoko, S. Suci Utami; Setia, Tatang Mitra. Orangutans: geographic variation in behavioral ecology and conservation. Oxford University Press. pp. 270–275. ISBN 978-0-19-921327-6. http://books.google.co.in/books?id=ZHsNfZC3DfYC&lpg=PA270&dq=nest-building%20by%20primates&pg=PA269#v=onepage&q&f=false. Retrieved 5 July 2011.
- ^ a b Hardus, ME; Lameira, AR; van Schaik, CP; Wich, SA. (2009). "Tool use in wild orang-utans modifies sound production: a functionally deceptive innovation?". Proceedings of the Royal Society B: Biological Sciences 276 (1673): 3689–3694. doi:10.1098/rspb.2009.1027.
- ^ Shapiro, G. L. (1975). Teaching linguistics concepts to a juvenile orangutan. California State University. Unpublished Masters Thesis.
- ^ Shapiro, G. L. (1982). "Sign acquisition in a home-reared/free-ranging orangutan: Comparisons with other signing apes". American Journal of Primatology 3 (1–4): 121–29. doi:10.1002/ajp.1350030111.
- ^ Shapiro, G. L. (1985). Factors influencing the variance in sign learning performance by four juvenile orangutans (Thesis). University of Oklahoma. http://ourf.org/images/documents/glsdissertation.pdf.
- ^ "Orangutan Language Project". Think Tank Research Projects. Smithsonian National Zoological Park. http://nationalzoo.si.edu/Animals/ThinkTank/ResearchProjects/OLP/default.cfm. Retrieved 2006-12-20.
- ^ Turner, Dorie (2007-04-12). "Orangutans play video games (for research) at Georgia zoo". http://www.usatoday.com/tech/gaming/2007-04-12-orangutans-games_N.htm. Retrieved 2007-04-12.
- ^ Dufour, V. Pelé, M.; Neumann, M.; Thierry, B.;Call, J. (2008). "Calculated reciprocity after all: computation behind token transfers in orang-utans". Biol. Lett 5 (2): 172–75. doi:10.1098/rsbl.2008.0644.
- ^ de Waal, Frans (January 1995). "The Loneliest of Apes". The New York Times. http://www.nytimes.com/1995/01/29/books/the-loneliest-of-apes.html?scp=3&sq=galdikas%20fossey%20goodall&st=cse. Retrieved 2012-2-26.
- ^ Wrangham, Richard W; Peterson, Dale (1996). Demonic males : apes and the origins of human violence. Boston: Houghton Mifflin. p. 137. ISBN 978-0-395-69001-7.
- ^ ""Beauty and the Beasts" by Carole Jahme". Salon.com. 2001-08-23. http://www.salon.com/2001/08/23/primates/singleton/. Retrieved 2011-01-27.
- ^ a b c d e f g h Singleton, I., Wich, S.A. & Griffiths, M. (2008). "Pongo abelii". IUCN Red List of Threatened Species. Version 2010.4. International Union for Conservation of Nature. http://www.iucnredlist.org/apps/redlist/details/39780. Retrieved 28 Jan. 2011.
- ^ a b c d e f g h i j k Ancrenaz, M., Marshall, A., Goossens, B., van Schaik, C., Sugardjito, J., Gumal, M. & Wich, S. (2008). "Pongo pygmaeus". IUCN Red List of Threatened Species. Version 2010.4. International Union for Conservation of Nature. http://www.iucnredlist.org/apps/redlist/details/17975. Retrieved 28 Jan. 2011.
- ^ Cheyne, S. M.; Thompson, C. J.; Phillips, A. C.; Hill, R. M.; Limin, S. H. (2007). "Density and population estimate of gibbons (Hylobates albibarbis) in the Sabangau catchment, Central Kalimantan, Indonesia". Primates 49 (1): 50–56. doi:10.1007/s10329-007-0063-0. PMID 17899314.
- ^ "Orangutan Action Plan 2007-2017" (in Indonesian) (PDF). Government of Indonesia. 2007. p. 5. http://www.yorku.ca/arusson/Papers/GoI%20OU%20action%20plan%2007-17.pdf. Retrieved 1 May 2010.
- ^ Patricia L. Miller-Schroeder (2004). Orangutans. The Untamed World. p. 64. ISBN 1-55388-049-8. http://books.google.com/?id=EOb5Ca5DjTQC&lpg=PP1&dq=Orangutans&pg=PP1#v=onepage&q=. Retrieved 2009-07-07.
- ^ Smith, David (2007-03-25). "Five years to save the orang utan". World (London: The Observer). http://observer.guardian.co.uk/world/story/0,,2042243,00.html. Retrieved 2010-05-20.
- ^ "The Last Stand of the Orangutan". United Nations Environment Programme. February, 2007. http://www.unep-wcmc.org/resources/publications/LastStand.htm. Retrieved 2007-12-03.
- ^ "Stop orangutan skull trade". antaranews.com. September 7, 2011. http://www.antaranews.com/en/news/75425/stop-orangutan-skull-trade. Retrieved 2012-01-16.
- ^ Thompson, Shawn (2010). The Intimate Ape: Orangutans and the Secret Life of a Vanishing Species. Citadel Press. p. 180. ISBN 978-0-8065-3133-5.
- ^ "Sepilok Orang Utan Sanctuary". Tourism Malaysia. http://www.tourism.gov.my/destinations/detail.php?theme=AD&map_code=sepilok&state=sabah. Retrieved 2011-12-30.