Primate
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{{otheruses|Primate (disambiguation)}}
{{Taxobox
| name = Primates<ref name=MSW3>{{MSW3 Groves|pages=111-184|id=12100001}}</ref>
| fossil_range = [[Late Cretaceous]] - Recent
| image = Olive baboon.jpg
| image_width = 250px
| image_caption = [[Olive Baboon]], an [[Old World monkey]]
| regnum = [[Animal]]ia
| phylum = [[Chordate|Chordata]]
| classis = [[Mammal]]ia
| infraclassis = [[Eutheria]]
| superordo = [[Euarchontoglires]]
| ordo = '''Primates'''
| ordo_authority = [[Carolus Linnaeus|Linnaeus]], 1758
| subdivision_ranks = Families
| subdivision = * 15, See [[Primate#Classification|classification]]
| range_map = Non-human primate range.png
| range_map_width = 240px
| range_map_caption = Range of the non-human primates
}}
A '''primate''' is any member of the biological [[order (biology)|order]] '''Primates''' ([[Latin]]: "prime, first rank"<ref>From [[Old French]] or [[French language|French]] ''primat'', from a noun use of Latin ''primat-'', from ''primus'' (“‘prime, first rank’”). The English singular ''primate'' was derived via [[back-formation]] from the Latin inflected form. [[Carl Linnaeus|Linnaeus]] thought this the "highest" order of mammals)</ref>), the group that contains all the species commonly related to the [[lemur]]s, [[monkey]]s, and [[ape]]s, with the last category including [[human]]s.<ref name=Goodman2>{{cite journal | journal = Journal of Molecular Evolution | year = 1990 | volume = 30 | pages = 260–266 | url = http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&list_uids=2109087&dopt=Abstract | title = Primate evolution at the DNA level and a classification of hominoids | author = M. Goodman, D. A. Tagle, D. H. Fitch, W. Bailey, J. Czelusniak, B. F. Koop, P. Benson, J. L. Slightom | doi = 10.1007/BF02099995}}</ref> With the exception of humans, which now inhabit every continent on Earth, most primates live in tropical or subtropical regions of [[the Americas]], [[Africa]] and [[Asia]]. A few species exist as far north in the Americas as southern [[Mexico]], and as far north in Asia as northern [[Japan]].
The Primates order is divided informally into three main groupings: [[prosimian]]s, monkeys of the [[New World monkey|New World]], and monkeys and apes of the [[Catarrhini|Old World]]. The prosimians are species whose bodies most closely resemble that of the early proto-primates. The most well known of the prosimians, the [[lemur]]s, are located on the [[Madagascar (island)|island of Madagascar]] and to a lesser extent on the [[Comoros Islands]], isolated from the rest of the world. The New World monkeys include the familiar [[capuchin monkey|capuchin]], [[howler monkey|howler]], and [[squirrel monkey]]s. They live exclusively in the Americas. Discounting humans, the rest of the [[simian]]s, the [[Old World monkey]]s and the apes, inhabit Africa and southern and central Asia, although fossil evidence shows many species existed in [[Europe]] as well.
According to fossil evidence, primitive ancestors of primates already existed in the late [[Cretaceous]]. [[Molecular clock]] studies suggest that the primate branch is even more ancient (originating at least in the mid-Cretaceous). They are now thought to be most closely related to [[flying lemur]]s and, more distantly, to [[treeshrew]]s. They probably have descended from [[Plesiadapiformes]].
==Distinguishing features==
Primates have radiated in [[arboreal]] habitats and although they have primitive (unspecialized) [[body plan]], retain many characteristics that are adaptations to this environment. They are distinguished by the retention of the [[clavicle]] in the pectoral girdle, [[Glenohumeral joint|shoulder joint]]s which allow high degrees of movement in all directions, five digits on the fore and hind limbs with [[Opposable thumb|opposable]] thumbs and big toes, fingernails, sensitive tactile pads on the ends of the digits, a trend towards reduced snout and [[Olfaction|olfactory]] senses, a reduced number of teeth compared to primitive mammals, a complex visual system with high [[visual acuity]] and [[color vision]], a large brain in comparison to body size, enlarged [[cerebral cortex]], two [[mammary gland]]s, 1-2 young per pregnancy, a long [[gestation]] and developmental period and a trend toward holding the torso upright leading to [[bipedalism]].<ref>{{cite book |author=Pough, F. W., Janis, C. M. & Heiser, J. B. |title=Vertebrate Life |chapter=Characteristics of Primates |year=2005 |origyear=1979 |edition=7th edition |publisher= Pearson |pages=630 |isbn=0-13-127836-3}}</ref> Primates are frequently highly [[social]], with flexible dominance hierarchies.<ref name=palaeos>{{cite web |author=White, T. & Kazlev, A. |url=http://www.palaeos.com/Vertebrates/Units/480Archonta/480.400.html#Primates | title=Archonta: Primates |publisher=[[Palaeos]]| accessdate=2008-06-03 | date=2006-01-08}}</ref> New World species form [[monogamous]] [[pair bond]]s, and show substantial [[Paternal|paternal care]] of young unlike most Old World monkeys.
==Anatomy and morphology==
===Skull and brain===
[[Image:Primate skull series with legend.png|thumb|right|Primate skull size and weight comparison]]
Primates have two forward-facing eyes on the front on the [[skull]], unlike many other mammals which have eyes on the side of the skull. The skull has a large domed [[cranium]] which is particularly prominent in [[anthropoid]]s. The cranium protects the large brain, a distinguishing characteristic of this group. The [[human skull]] has a three-fold greater [[endocranium|endocranial volume]] (inner surface of skull) than the volume of the greatest non-human primate, reflecting a larger brain size. The mean human endocranial volume is 1201 [[cubic centimetre]]s, it is 469 cm³ in gorillas, 400 cm³ in chimpanzees and 397 cm³ in orangutans.<ref>{{cite book | author= Aiello, L. & Dean, C. | year=1990 | title=An Introduction to Human Evolutionary Anatomy | publisher=Academic Press | pages=193 | isbn=0120455900}}</ref> There is a tendency within primates towards reduction of the olfactory region of the brain and expansion of the [[cerebrum]], especially the [[cerebral cortex]]. This is correlated with an increasing reliance on vision and increasingly complex social behavior.<ref name=adw>{{cite web | url=http://animaldiversity.ummz.umich.edu/site/accounts/information/Primates.html | title="Primates" (On-line) |publisher=Animal Diversity Web | author=Myers, P. | date=1999 | accessdate=2008-06-03}}</ref> The primary evolutionary trend of primates has been the elaboration of the brain, in particular the [[neocortex]]; a layer of the cerebral hemispheres which is involved with [[sense|sensory perception]], generation of [[motor cortex|motor commands]], spatial reasoning, [[consciousness|conscious thought]] and, in humans, [[language]]. While other mammals rely heavily on olfactory stimuli the arboreal habit of primates has led to a tactile, visually dominant sensory system.
===Eyes===
Primates have forward-facing [[Primates#Color vision|color]] [[binocular vision]]. This allows overlapping fields of view, which give primates [[Stereoscopy|stereoscopic vision]] and accurate distance perception, which was useful for the brachiating ancestors of humans. [[Evolution of color vision in primates|Color vision]] is important for finding and collecting food. All primates have a [[supraorbital ridge|bony ridge]] located above the eye sockets which reinforces weaker bones in the face which are put under strain during chewing. [[Strepsirrhine]]s have a [[postorbital bar]], a bone which runs around the eyesocket, to protect their eyes, which is in contrast to the higher primates, [[haplorrhine]]s, which have evolved fully enclosed sockets.<ref name=Campbell>{{cite book | author = Campbell, Bernard G., Loy, James D. | year = 2000 | title = Humankind Emerging (8th edition) | publisher = Allyn & Bacon | pages = 85 |isbn=0673523640}}</ref>
===Hands and feet===
[[Image:PrimateFeet.jpg|thumb|upright|An 1893 drawing of the hands and feet of primates]]
All primates have five digits on each limb ([[Dactyly#pentadactyly|pentadactyly]]), with keratin nails on the anterior ends. The ventral sides of the hands and feet have [[tactile pad]]s on the [[distal phalanges]]. They have [[thumb|opposable thumbs]], which are a characteristic primate feature, but are not limited to this order; [[opossum]]s, for example, also have opposing thumbs. These thumbs allow some species to use [[Tool use by animals|tools]] to perform some tasks. In primates, the combination of opposing thumbs, short fingernails (rather than claws) and long, inward-closing [[finger]]s is a relic of the ancestral practice of gripping branches, and has, in part, allowed some species to develop [[brachiation]] as a significant means of transportation. Prosimians have clawlike nails on the second toes of their feet.
===Limbs and vertebral column===
The primate [[clavicle]] is retained as prominent element of the [[pectoral girdle]], this allows the [[shoulder joint]] broad mobility. Apes have more mobile shoulder joints and arms due to the dorsal position of the [[scapula]], broad ribcages that are flatter front-to-back, and a shorter, less mobile spine compared to Old World monkeys (with [[caudal]] [[vertebrae]] greatly reduced, resulting in tail loss in some species). Old World monkeys are unlike apes in that most have tails, and unlike the New World monkeys in that their tails are never [[Prehensile tail|prehensile]]. Only the New World [[Atelidae]] family have prehensile tails.
===Nose, mouth and teeth===
Primates show a trend towards a reduced [[snout]].<ref name=palaeos /> Technically, the distinction of Old World monkeys from New World monkeys depends on the structure of the nose, and the distinction of Old World monkeys from apes depends on [[dentition]]. In New World monkeys the nostrils face sideways, whilst in Old World monkeys, they face downwards. There is a considerably varied [[dentition|dental pattern]] in primates and although some have lost most of their [[incisor]]s; all retain at least one lower incisor. In most strepsirhines, the lower incisors form a [[toothcomb]] which is thought to be used in grooming and possibly foraging. Old World monkeys also have eight [[premolar]]s, compared with twelve in New World monkeys. The Old World species are divided into apes and monkeys depending on the number of [[cusp]]s on their [[molar]]s (apes have five—the "Y-5" molar pattern, Old World monkeys have only four in a [[bilophodont]] pattern). A [[hypocone]] evolved in early primate history, while the [[paraconid]] was lost, leaving both upper and lower teeth with a basically [[quadrate]] pattern. Primitively, primate molars were [[brachydont]] and [[tuberculosectorial]], but they have become bunodont and quadrate in a number of modern forms.<ref name=adw/> Prosimians are distinguished by their immobilized upper lips, moist tip to their nose and forward-facing lower front teeth.
==Physiology==
===Sexual dimorphism===
[[Image:Hamadryas Baboon.jpg|thumb|right|Distinct sexual size dimorphism can be seen between the male [[Hamadryas Baboon]]s (grey) and the female (brown)]]
{{main|Sexual dimorphism in non-human primates}}
[[Sexual dimorphism]] is often exhibited in [[simian]]s; though to a greater degree in Old World species (apes and some monkeys) than New World species. Recent studies have mainly used the technique of [[comparative analysis]] to examine both the variation in the expression of the dimorphism among primates and the fundamental causes of sexual dimorphism. Primates usually have dimorphism in [[body mass]]<ref>{{cite journal |author=Ralls, K |year=1976 |title=Mammals in Which Females are Larger Than Males |journal=The Quarterly Review of Biology |volume=51 |issue=2 |doi=10.1086/409310 | pages=245}}</ref><ref>{{cite journal | author=Lindstedtand & Boyce | year=1985 | journal=The American Naturalist | volume=125 |pages=873 |doi = 10.1086/284385 |title = Seasonality, Fasting Endurance, and Body Size in Mammals}}</ref> and canine tooth size<ref>{{cite journal |author=Frisch, J. E. |year=1963 |title=Sex-differences in the canines of the gibbon (''Hylobates lar'') |journal = Primates |volume = 4 |issue = 2 |doi=10.1007/BF01659148 |pages=1}}</ref><ref>{{cite journal |author=Kay, R. F. |year=1975 |title=The functional adaptations of primate molar teeth |journal=American Journal of Physical Anthropology |volume=43 |issue=2 |pages=195–215 |doi=10.1002/ajpa.1330430207}}</ref> along with pelage and skin color.<ref>{{cite book |author=Crook, J. H. |editor=Campbell, B. G. |year=1972 |chapter=Sexual selection, dimorphism, and social organization in the primates |title=Sexual selection and the descent of man |publisher=Aldine Transaction |pages=246 |isbn=978-0202020051}}</ref> The dimorphism can been attributed to and affected by different factors; [[mating system]],<ref name="cheverud">{{cite journal |author=Cheverud, J.M., Dow, M. M. & Leutenegger, W. |year=1985 |title=The quantitative assessment of phylogenetic constraints in comparative analyses: Sexual dimorphism in body weight among primates |journal=Evolution |volume=39 |issue=6 |pages=1335–1351 |doi=10.2307/2408790}}</ref> size,<ref name="cheverud" /> habitat and diet.<ref name="leutenegger">{{cite journal | author=Leutenegger,W. & Cheverud, J. M. | year=1982 | title=Correlates of sexual dimorphism in primates: Ecological and size variables | journal = International Journal of Primatology | volume = 3 | issue = 4 | doi=10.1007/BF02693740 | pages=387}}</ref>
Comparative analyses have substantiated the [[sexual selection]] hypotheses, and have generated a more complete understanding of the relationship between sexual selection, [[natural selection]], and mating systems in primates. Studies are helping to find the relative contribution of the various selective and non-selective mechanisms in sexual dimorphism evolution and expression.<ref>{{cite journal |author=Plavcan, J.M. |year=2001 |title=Sexual dimorphism in primate evolution |journal=American Journal of Physical Anthropology |volume=33 |pages=25–53 |doi=10.1002/ajpa.10011}}</ref> These studies have shown that dimorphism is the product of changes in both male and female traits. [[Ontogenic scaling]], where relative extension of a common growth trajectory occurs, may show some insight into the relationship between sexual dimorphism and growth patterns.<ref name="ohiggins">{{cite journal |author=O'Higgins, P. & Collard, M. |year=2002 |title=Sexual dimorphism and facial growth in papionin monkeys |journal=Journal of Zoology |volume=257 |issue=2 |pages=255–272 |doi=10.1017/S0952836902000857}}</ref> There is some evidence from the fossil record that suggests that there was [[convergent evolution]] of dimorphism, and some extinct [[hominid]]s probably exceeded dimorphism of any living primate.
===Color vision===
{{main|Evolution of color vision in primates}}
[[Color vision]] in primates is unique in the evolution of most [[eutherian]] [[mammals]]. While our remote [[vertebrate]] ancestors possessed [[trichromatic vision|trichromacy]], our [[nocturnal]], [[warm-blooded]], mammalian ancestors lost one of three cones in the [[retina]] at the time of [[dinosaur]]s. [[Fish]], [[reptile]]s and [[bird]]s are therefore trichromatic while all mammals, with the exception of some primates and [[marsupial]]s,<ref>{{cite journal | author=Arrese, C. A., ''et al'' | year=2005 | title=Cone topography and spectral sensitivity in two potentially trichromatic marsupials, the quokka (Setonix brachyurus) and quenda (Isoodon obesulus) | journal=Proceedings of Biological Science | volume=272 | issue=1565 | doi=10.1098/rspb.2004.3009 | pages=791–796}}</ref> are strictly dichromats.
Primates achieve trichromacy through color receptors ([[cone cell]]s), with [[visible spectrum|spectral]] peaks in the [[Violet (color)|violet]] (short wave, [[Photopsin#Types|S]]), [[green]] (middle wave, M), and [[yellow]]-green (long wave, L) wavelengths. All primates, however, are not capable of trichromacy. The catarrhines are routinely trichromatic, meaning that both males and females possess three [[opsin]]s (pigments) sensitive to 430 nanometre, 530 nm, and 560 nm wavelengths.<ref name=Bowmaker1991>{{cite journal | author = Bowmaker, J. K., and S. Astell | year = 1991 | title = Photosensitive and photostable pigments in the retinae of Old World monkeys | journal = Journal of Experimental Biology | volume = 156 | pages = 1–19 |issn=0022-0949 |url=http://jeb.biologists.org/cgi/reprint/156/1/1.pdf |format=pdf |accessdate=2008-06-16}}</ref> Platyrrhines, on the other hand are non-routinely trichromatic; only a small population of platyrrhines are trichromats.<ref name=Surridge2003>{{cite journal | author = Surridge, A. K., and D. Osorio | year = 2003 | title = Evolution and selection of trichromatic vision in primates | journal = Trends in Ecology and Evolution | volume = 18 | pages = 198–205 | doi = 10.1016/S0169-5347(03)00012-0}}</ref>
Howler monkeys have reinvented routine trichromatism through a recent [[gene duplication]] of the red-green opsin gene.<ref name=Lucas2003>{{cite journal | author = Lucas, P. W., and N. J. Dominy | year = 2003 | title = Evolution and function of routine trichromatic vision in primates | journal = Evolution | volume = 57 |pages = 2636–2643 | doi = 10.1554/03-168}}</ref> This has allowed trichromacy for both sexes, the X chromosome has gained two loci to house both the green allele and the red allele. Howler monkeys are perhaps the most folivorous of the New World monkeys. Fruits are not part of their diet, and the type of leaves they consume (young, nutritive, and digestible), are detectable only by a red-green signal. Field work exploring the dietary preferences of howler monkeys suggest that routine trichromacy was environmentally selected for.<ref name=Surridge2003>{{cite journal | author = Surridge, A. K., and D. Osorio | year = 2003 | title = Evolution and selection of trichromatic vision in primates | journal = Trends in Ecology and Evolution | volume = 18 | pages = 198–205 | doi = 10.1016/S0169-5347(03)00012-0}}</ref>
===Locomotion===
Various species of primates move by [[brachiation]], [[bipedalism]], [[leaping]], arboreal and terrestrial [[quadrupedalism]], [[climbing]] or [[knuckle walking]].
Most prosimians are arboreal quadrupedalists and climbers. Many are also terrestrial quadrupedalists, while some are leapers. Most monkeys are both arboreal and terrestrial quadrupedalists and climbers. Gibbons are brachiators. Chimps, orangutans, and gorillas all knuckle walk, and can move bipedally for short distances. Humans are the only fully bipedal species.
==Ethology==
{{see|Primatology}}
===Social systems===
[[Image:Jigokudani_hotspring_in_Nagano_Japan_001.jpg|thumb|right|[[Japanese Macaque]]s bathe together in Jigokudani Hot Spring]]
[[Richard Wrangham]] proposed that [[Social structure|social systems]] are best classified by the amount of movement by females occurring between groups.<ref>{{cite book |author=Wrangham, R. W. |authorlink=Richard Wrangham |chapter=Mutualism, kinship and social evolution |year=1982 |title=Current Problems in Sociobiology |publisher=Cambridge University Press |pages=269–289 |isbn=0521242037}}</ref> He proposed 4 categories:
*Female transfer systems - females move away from the group in which they were born. Females of a group will not be closely related whereas males will have remained with their natal groups and therefore the close association may be influential in affecting social behaviour. The groups formed are generally quite small. This organisation can be seen in chimpanzees, the males will cooperate in defending their territories.
*Male transfer systems - while the females remain in their natal groups, the males will emigrate as adolescents. Polygynous and multi-male societies are classed in this category. Group sizes are usually larger.
*Monogamous species - a male-female bond, sometimes accompanies by juvenile offspring. There is shared responsibility of parental care and territorial defence. The offspring will leave the parents territory during adolescence.
*Solitary species - often males who defend territories that include the home ranges of several females, found in the prosimians. Orangutans do not defend their territory but have the same organisation.
Primatologist Dr. [[Jane Goodall]], who studied in the [[Gombe Streams National Park]], noted [[fission-fusion society|fission-fusion societies]] in chimpanzees.<ref>{{cite journal | title=Noninvasive paternity assignment in Gombe chimpanzees | author=Constable, J. L. ''et al'' | journal=Molecular Ecology | year=2001 | volume=10 | issue=5 | pages=1279–1300 |doi=10.1046/j.1365-294X.2001.01262.x}}</ref> There is ''fission'' where the main group splits up to forage during the day, then ''fusion'' when the group returns at night to sleep as a group. This social structure can also be observed in [[Hamadryas Baboon]]s, [[Gelada]]s, [[spider monkey]]s and [[Bonobo]]s.
These social systems are affected by three main ecological factors; distribution of resources, group size and predation.<ref name=vertlife>{{cite book |author=Pough, F. W., Janis, C. M. & Heiser, J. B. |title=Vertebrate Life |chapter=Primate Societies |year=2005 |origyear=1979 |edition=7th edition |publisher= Pearson |pages=621-623 |isbn=0-13-127836-3}}</ref> Within a social group there is a balance between cooperation and competition. Cooperation comes in the form of [[allogrooming]]; whereby [[ectoparasite]]s are removed and wounds cleaned, food sharing and collective defence against predators or of a territory. Competition is demonstrated by aggression and may come about through availability of food, sleeping sites or mates. Aggression is often used in establishing [[Social hierarchy|social hierarchies]].<ref name=vertlife/>
===Cognition===
{{see|Great ape language|Primate empathy|Hominid intelligence|Chimpanzee intelligence|Animal intelligence}}
===Laughter===
[[Laughter]] may not be confined or unique to humans, despite Aristotle's observation that "only the human animal laughs". But some behavioural psychologists argue that self-awareness of one's situation, or the ability to identify with somebody else's predicament, are prerequisites for laughter, so animals do not laugh like humans do.
[[Chimpanzee]]s, [[gorillas]], and [[orangutans]] show laughter-like vocalizations in response to physical contact, such as wrestling, play chasing, or tickling. This is documented in wild and captive chimpanzees. Chimpanzee laughter is not readily recognizable to humans as such, because it is generated by alternating inhalations and exhalations that sound more like breathing and panting. The differences between chimpanzee and human laughter may be the result of adaptations that have evolved to enable human speech. There are instances in which non-human primates have been reported to have expressed joy. One study analyzed and recorded sounds made by human babies and Bonobos (a species of chimpanzee) when they were tickled. It found although the Bonobo's laugh was a higher frequency, the laugh followed the same spectrographic pattern of human babies to include as similar facial expressions. Humans and chimpanzees share similar ticklish areas of the body such as the armpits and belly. The enjoyment of tickling in chimpanzees does not diminish with age.<ref>{{cite journal | archiveurl = http://web.archive.org/web/20070301065905/http://www.discover.com/issues/apr-03/features/featlaugh/ | archivedate=2007-03-01 | journal = Discover | author = Johnson, S. | title = Emotions and the brain | volume = 24 | issue = 4 | month = April | year = 2003}}</ref>
==Habitat and distribution==
Primates evolved from [[arboreal]] animals and many species live most their lives in trees. Some species are partially [[Terrestrial animal|terrestrial]], such as [[baboon]]s and the [[Patas Monkey]] and a few species are fully terrestrial, such as the [[Gelada]] and [[Human]]s. Non-human primates live in a diverse number of forested habitats in the tropical latitudes of Africa, India, Southeast Asia, and South America, including [[rain forest]]s, [[mangrove]] forests, and [[montane forest]]s. There are some examples of non-human primates that live outside of the tropics; the mountain-dwelling [[Japanese Macaque]] lives in the north of [[Honshu]] where there is snow-cover eight months of the year, while the [[Barbary macaque]] lives in the [[Atlas Mountains]] of Algeria and Morocco. Primates also have a considerable vertical range; the [[Black Snub-nosed Monkey]] has been found living in the [[Hengduan Shan|Hengduan Mountains]] at altitudes of 4,700 metres (15,400 ft),<ref name=Long94>{{cite journal | author = Long, Y. C., Kirkpatrick, R. C., Zhong, T., and Xiao, L. | year = 1994 | title = Report on the distribution, population, and ecology of the Yunnan snub-nosed monkey (Rhinopithecus bieti) | journal = Primates | volume = 35 | pages = 241–250 | doi = 10.1007/BF02382060}}</ref> [[Mountain Gorilla]]s can be found at 4,200 metres (13,200 ft) crossing the [[Virunga Mountains]]<ref name=Schaller>{{cite book | author = Schaller, G.B. | year = 1963 | title = The Mountain Gorilla: Ecology and Behavior | location = Chicago | publisher = Univ. Chicago Press |isbn=978-0226736358}}</ref> and the Gelada has been found at elevations of up to 5,000 meters (16,400 ft) in the [[Ethiopian Highlands]]. Although most species are generally shy of water, a few are good swimmers and are comfortable in swamps and watery areas, including the [[Proboscis Monkey]], [[De Brazza's Monkey]] and [[Allen's Swamp Monkey]], which has developed small webbing between its fingers. Some primates, such as the [[Rhesus Macaque]] and the [[Hanuman Langur]], can exploit human-modified environments and even live in cities.{{Fact|date=May 2008}}
===Conservation status===
{{seealso|Critically endangered species#Primates}}
[[Madagascar]], home to five endemic primate families, has experienced the greatest extinction of the recent past; since human settlement 1,500 years ago, at least eight classes and fifteen species have become extinct due to hunting and habitat destruction. Among the primates wiped out were [[Archaeoindris]], a lemur larger than a silverback gorilla and the families [[Palaeopropithecidae]] and [[Archaeolemuridae]].
Although many species are still widespread in the wild, large-scale logging has destroyed expansive areas of land which provide many species' habitat. Among the most threatened primates include the [[Sumatran Orangutan]], the [[gorilla]], [[Golden Lion Tamarin]]s and the [[Silvery Gibbon]]. [[Miss Waldron's Red Colobus]] was the first primate to be declared extinct in the 21st century when a report from 1993-1999 could find no trace of the monkey. It was last sighted in 1973 and was officially declared extinct in 2000.
==Evolutionary history==
<center>
{{Clade
| label1=[[Euarchontoglires]]
| 1={{Clade
| label1=[[Glires]]
| 1={{Clade
| 1=[[Rodent]]ia (rodents)
| 2=[[Lagomorpha]] (rabbits, hares, pikas)}}
| label2=[[Euarchonta]]
| 2={{Clade
| 1=[[Treeshrew|Scandentia]] (treeshrews)
| 2={{Clade
| 1=[[Colugo|Dermoptera]] (colugos)
| 2={{Clade
| 1=†[[Plesiadapiformes]]
| 2=Primates}} }} }} }}
}}
</center>
The Primate [[order (biology)|order]] lies in a tight clustering of related orders (the [[Euarchontoglires]]) within the [[Eutheria]], a subclass of [[Mammalia]]. Recent molecular genetic research on primates, [[colugo]]s, and [[treeshrew]]s has shown that the two species of colugos ([[Dermoptera]]) are more closely related to the primates than the treeshrews of the order [[Scandentia]], even though the treeshrews were at one time considered primates. These three orders make up the [[Euarchonta]] [[clade]]. This clade combines with the [[Glires]] clade (made up of the [[Rodent]]ia and [[Lagomorpha]]) to form the Euarchontoglires clade. Variously, both Euarchonta and Euarchontoglires are ranked as superorders. Also, some scientists consider Dermoptera a suborder of Primates and call the "true" primates the suborder Euprimates (Hoffstetter, 1978).<ref>{{cite book | author = McKenna, Malcolm C. and Bell, Susan K.| year = 1997 | title = Classification of Mammals Above the Species Level| publisher = Columbia University Press, New York| pages = 329 |isbn=023111012X}}</ref>
===Evolution===
[[Image:Katta Lemur catta.jpg|upright|thumb|[[Ring-tailed Lemur]], a [[strepsirrhine]] primate]]
In modern, cladistic reckonings, the Primate order is also a true clade. The suborder [[Strepsirrhini]], the "[[rhinarium|wet-nosed]]" primates, split off from the primitive primate line about 63 [[mya (unit)|million years ago (mya)]]. The seven strepsirhine families are the four related [[lemur]] families and the three remaining families that include the [[loris]]es, the [[Aye-aye]], the [[galago]]s, and the [[potto]]s.<ref name=MSW3/> Older classification schemes wrap the [[Lepilemuridae]] into the [[Lemuridae]] and the [[Galagidae]] into the [[Lorisidae]], yielding a three-two family split instead of the four-three split as presented here.<ref name=MSW3/> Other lineages of lower primates inhabited Earth. During the [[Eocene]], most of the northern continents were dominated by two dominant groups, the [[adapid]]s and the [[omomyid]]s. The former is considered a member of Strepsirrhini, but it does not have a [[tooth-comb]] like modern lemurs. The latter was related closely to tarsiers, monkeys, and apes. Adapids survived until 10 mya; omomyids on the other hand perished 20 million years earlier.
The Aye-aye is difficult to place in Strepsirrhini.<ref name=MSW3/> Its family, Daubentoniidae, could be a lemuriform primate and its ancestors split from lemur line more recently than the lemurs and lorises split, about 50 mya. Otherwise it is sister to all of the other strepsirrhines, in which case in evolved away from the main strepsirrhine line between 50 and 63 mya.
[[Image:Tamarin portrait 2 edit3.jpg|left|thumb|[[Emperor Tamarin]], a [[New World monkey]]]]
The suborder [[Haplorrhini]], the "dry-nosed" primates, is composed of two sister clades.<ref name=MSW3/> The [[prosimian]] tarsiers in family [[Tarsiidae]] (monotypic in its own infraorder [[Tarsiiformes]]), represent the most primitive division at about 58 mya. The [[Simiiformes]] infraorder contains the two parvorders: the [[New World monkey]]s in one, and the [[Old World monkey]]s, [[human]]s and the other [[ape]]s in the other.<ref name=MSW3/> This division happened about 40 mya. However about 30 mya, three groups split from the main haplorrhine lineage. One group stayed in Asia and are closest in kin to the "dawn monkey" ''[[Eosimias]]''. The second stayed in Africa, where they developed into the Old World primates. The third rafted to South America to become the New World monkeys. Mysteriously the aboriginal Asian Haplorrhini vanished from record once Africa collided with Eurasia 24 mya. Apes and monkeys spread into Europe and Asia. Close behind came lorises and tarsiers, also African castaways. The first hominid fossils were discovered in Northern Africa and date back 7 mya. Modern humans did not appear until 0.2 mya, eventually becoming the most prevalent primate and mammal on Earth.
The discovery of new species happens at a rate of a few new species each year, and the evaluation of current populations as distinct species is in flux. [[Colin Groves]] listed about 350 species of primates in ''Primate Taxonomy'' in 2001.<ref name=PT>{{cite book |title=Primate Taxonomy |publisher=Smithsonian Institute Press |year=2001 |author=Groves, C. P. |isbn=1-56098-872-X}}</ref> The recently published third edition of ''Mammal Species of the World'' (MSW3) lists 376 species.<ref name=MSW3/> But even MSW3's list falls short of current understanding as its collection cutoff was in 2003, and a number publications since then have pushed the number of species up to 406. Notable new species not listed in MSW3 include the [[Bemaraha Woolly Lemur]] (''Avahi cleesei'') (named after [[United Kingdom|British]] [[actor]] and [[lemur]] enthusiast [[John Cleese]]) and the [[GoldenPalace.com Monkey]] (whose name was put up for auction).
==Classification==<!-- This section is linked from [[Primate]] -->
[[Image:Primates-drawing.jpg|thumb|right|A 1927 drawing of [[chimpanzee]]s, a [[gibbon]] (top right) and two [[orangutan]]s (center and bottom center)]]
[[Image:Human.svg|right|upright|thumb|''[[Homo sapiens]]'', a member of the order Primates]]
* '''ORDER PRIMATES'''
** '''Suborder [[Strepsirrhini]]''': non-tarsier prosimians
*** Infraorder [[Lemuriformes]]
**** Superfamily [[Cheirogaleidae|Cheirogaleoidea]]
***** Family [[Cheirogaleidae]]: dwarf lemurs and mouse-lemurs (31 species)
**** Superfamily [[Lemuroidea]]
***** Family [[Lemuridae]]: lemurs (19 species)
***** Family [[Lepilemuridae]]: sportive lemurs (22 species)
***** Family [[Indriidae]]: woolly lemurs and allies (18 species)
*** Infraorder [[Aye-aye|Chiromyiformes]]
**** Family [[Aye-aye|Daubentoniidae]]: Aye-aye (1 species)
*** Infraorder [[Lorisiformes]]
**** Family [[Lorisidae]]: lorises, pottos and allies (9 species)
**** Family [[Galagidae]]: galagos (19 species)
** '''Suborder [[Haplorrhini]]''': tarsiers, monkeys and apes
*** Infraorder [[tarsier|Tarsiiformes]]
**** Family [[tarsier|Tarsiidae]]: tarsiers (8 species)
*** Infraorder [[Simiiformes]]
**** Parvorder [[Platyrrhini]]: New World monkeys
***** Family [[Cebidae]]: marmosets, tamarins, capuchins and squirrel monkeys (56 species)
***** Family [[Aotidae]]: night or owl monkeys (douroucoulis) (7 species)
***** Family [[Pitheciidae]]: titis, sakis and uakaris (41 species)
***** Family [[Atelidae]]: howler, spider and woolly monkeys (24 species)
**** Parvorder [[Catarrhini]]
***** Superfamily [[Old World monkey|Cercopithecoidea]]
****** Family [[Old World monkey|Cercopithecidae]]: Old World monkeys (135 species)
***** Superfamily [[Hominoidea]]
****** Family [[Hylobatidae]]: gibbons or "lesser apes" (13 species)
****** Family [[Hominidae]]: humans and other great apes (7 species)
The order Primates was established by [[Carl Linnaeus]] in 1758, in the tenth edition of his book [[Systema Naturae]],<ref>{{cite book | author = Linnaeus, Carolus | year = 1758 | title = Sistema naturae per regna tria Naturae, secundum classes, ordines, genera, species, cum characteribus differentiis, synonimis locis. Tomus I | publisher = Impensis direct. Laurentii Salvii, Holmia | pages = pp. 20-32}}</ref> for the genera ''[[Homo (genus)|Homo]]'' (humans), ''[[Simia]]'' (other apes and monkeys), ''[[Lemur (genus)|Lemur]]'' (prosimians) and ''[[Vespertilio]]'' (bats). In the first edition of the same book (1735), he had used the name [[Anthropomorpha]] for ''Homo'', ''Simia'' and ''[[Bradypus]]'' (sloths).<ref>{{cite book | author = Linnaeus, Carolus | year = 1735 | title = Sistema naturae sive regna tria Naturae systematice proposita per classes, ordines, genera, & species| publisher = apud Theodorum Haak, Lugduni Batavorum | pages = pp. s.p.}}</ref> In 1839, [[Henri-Marie Ducrotay de Blainville]], following Linnaeus and imitating his nomenclature, established the orders [[Secundates]] (including the suborders [[Chiroptera]], [[Insectivora]] and [[Carnivora]]), [[Tertiates]] (or [[Glires]]) and [[Quaternates]] (including [[Gravigrada]], [[Pachydermata]] and [[Ruminantia]]),<ref>{{cite book | author = Blainville, Henri-Marie Ducrotay de | year = 1839 | chapter = Nouvelle classification des Mammifères | title = Annales Françaises et Etrangères d’Anatomie et de Physiologie Appliquées à la Médicine et à l’Histoire Naturelle, 3| pages = pp. 268-269}}</ref> but these new taxa were not accepted.
In older classifications, the Primates were divided into two superfamilies: Prosimii and Anthropoidea. The Prosimii included all of the [[prosimian]]s: all of Strepsirrhini plus the tarsiers. The Anthropoidea contained all of the [[simian]]s.
===Hybrids===
Primate [[Hybrid (biology)|hybrids]] usually arise in captivity<ref name=songs>{{cite journal |title=Songs of hybrid gibbons (''Hylobates lar'' × ''H. muelleri'') |author=Tenaza, R. |year=1984 |journal=American Journal of Primatology |volume=8 |issue=3 |pages=249–253 |doi=10.1002/ajp.1350080307}}</ref> though there have been examples of primate hybrids in the wild.<ref name=natoccur>{{ cite journal |year=1966 |title=Naturally Occurring Primate Hybrid |author=Bernsteil, I. S. |journal=Science |volume=154 |issue=3756 |pages=1559–1560 |doi=10.1126/science.154.3756.1559 |pmid=4958933}}</ref> Hybridisation occurs where two species' territories overlap, these [[hybrid zone]]s may be created by humans when animals are placed in zoos or may be due to environmental pressures such as predation.<ref name=natoccur/> In ''The variation of animals and plants under domestication'' [[Charles Darwin]] noted: "Several members of the family of [[Lemur]]s have produced hybrids in the Zoological Gardens."<ref>{{cite book |author=Darwin, C |title=The variation of animals and plants under domestication |year=1868 |publisher=London: John Murray |edition=1st edition |volume=2 |pages=153 |url=http://darwin-online.org.uk/content/frameset?viewtype=side&itemID=F877.2&pageseq=167 |isbn=1421270730}}</ref>
Many [[gibbon]]s are hard to identify based on fur coloration and are identified either by song or genetics.<ref name=songs/> These morphological ambiguities have led to hybrids in zoos. Zoos often receive gibbons of unknown origin and therefore rely on morphological variation or labels that are impossible to verify to assign species and subspecies names so it is common for separate species of gibbons to be misidentified and housed together. Interspecific hybrids, hybrids within a genus, also occur in wild gibbons where the ranges overlap.<ref>{{cite journal |author=Sugawara, K. |title=Sociological study of a wild group of hybrid baboons between ''Papio anubis'' and ''P. hamadryas'' in the Awash Valley, Ethiopia |year=1979 |journal=Primates |volume=20 |issue=1 |doi=10.1007/BF02373827 |pages=21–56}}</ref>
Intergeneric hybridizations, hybrids of a different genus, have been found in the wild. Though belonging to genera that have been distinct for several million years interbreeding still occurs between [[Gelada]]s (''Theropithecus gelada'') and [[Hamadryas Baboon]] (''Papio hamadryas'').<ref>{{cite journal |author=Jolly, C. J. ''et al'' |title=Intergeneric Hybrid Baboons |journal=International Journal of Primatology |year=1997 |volume=18 |issue=4 |doi=10.1023/A:1026367307470 |pages=597–627}}</ref>
==Non-human primate==
'''Non-human primates''' (NHP) are all [[species]] of [[animals]] under the order Primates that are not a member of the [[genus]] [[Homo]]. It is considered the proper term when referring to [[monkeys]] used in [[animal research]] or kept as [[exotic pet|pets]]. Governments of many nations have strict care requirements of NHPs that are kept in captivity. In the US, federal guidelines extensively regulate aspects of NHP housing, feeding, enrichment, and breeding.<ref name=usda_nhpresearch>{{cite web|title=Nonhuman Primates : Research Animals : Animal Welfare Information Center|url=http://awic.nal.usda.gov/nal_display/index.php?info_center=3&tax_level=3&tax_subject=169&topic_id=1078&level3_id=5345&level4_id=0&level5_id=0&placement_default=0|accessdate=2008-07-14}}</ref>
===Role in scientific research===
[[Image:Monkey5.jpg|right|thumb|Monkeys, imported for experimentation, in a crate. Credit: [[British Union for the Abolition of Vivisection|BUAV]]]]
NHPs are commonly used in scientific research as [[Non-human primate experiments|animal subjects]]. Thousands of NHPs are used around the world in scientific experiments. Given their close [[genetic]] similarity to [[humans]], they are excellent [[animal models]] with which to conduct experiments that are relevant to humans. Common research applications for NHPs include [[Clinical trial#Pre-clinical studies|preclinical trials]], [[neuroscience]], and [[ophthamology]] studies. The anatomy of NHPs, particularly their [[brain]] and [[eyes]], more closely parallel humans than any other group of animals. The most commonly used NHP for laboratory research is the [[rhesus macaque]]. [[Chimpanzee]]s, [[baboon]]s, [[marmoset]]s, and [[green monkey]]s are also commonly found in NHP research labs.<ref name=Conlee2004>{{cite journal
| author = Conlee, Kathleen M.; Hoffeld, Erika H.; Stephens, Martin L.
| year = 2004
| title = A Demographic Analysis of Primate Research in the United States
| journal = ATLA (Alternatives to Laboratory Animals)
| volume = 32
| issue = Sup 1
| pages = 315–322
| url = http://www.worldcongress.net/2002/proceedings/C2%20Conlee.pdf
}}</ref> In 2004 the [[European Union]] used around 10,000 animals; in 2005 in Great Britain alone there were 4,652 experiments conducted on 3,115 non-human primates.<ref>{{cite book |author= |title=Statistics of scientific procedures on living animals: Great Britain 2005 |year=2006 |month=July |publisher=[[The Stationary Office]] |isbn=0101687729 |url=http://www.official-documents.gov.uk/document/cm68/6877/6877.pdf |format=pdf |accessdate=2008-06-16}}</ref> As of 2004, 3,100 NHPs were living in captivity in the [[United States]], in [[zoo]]s, [[circus]]es, and [[Laboratory|laboratories]]; 1,280 of them being used in experiments.<ref name=GA/> European campaign groups such as the [[BUAV]] are seeking a ban on all NHP use in experiments as part of the [[European Union]]'s current review of existing law on animal experimentation.
===Legal status===
Within the order [[Primates]], only humans are recognized as [[person]]s and protected in law by the [[United Nations]] [[Universal Declaration of Human Rights]].<ref>'''Article 6''': Everyone has the right to recognition everywhere as a person before the law.</ref> The status of NHPs has generated much debate, particularly through the [[Great Ape Project]] which argues for the personhood of the non-human members of the family [[Hominidae]].<ref name=GA>{{cite web |author=Cavalieri, P. & Singer, P. |url=http://www.greatapeproject.org/declaration.php |title=Declaration on Great Apes |publisher=[[Great Ape Project]] |accessdate=2008-06-16}}</ref> In 1995 Ignaas Spruit, director of Leiden (Netherlands) based Pro-Primates organization, went farther, as he proposed that some rights should be recognized to all NHPs.<ref>{{cite book | author = Spruit, I. | year = 1995 | chapter = On Declaring Non-human Primate Rights: An Approach to Primate Protection | editor = Corbey, R. & Theunissen, B. | title = Ape, Man, Apeman: Changing Views since 1600 | publisher = Department of Prehistory, Leiden University | pages = pp. 377-383 |isbn=9073368057}}</ref> American anthropologist [[Earnest Albert Hooton]], enlarged the famous quote by [[Terence]], ''"Homo sum: humani nil a me alienum puto"'' and used to say ''"Primas sum: primatum nil a me alienum puto"'', which he himself translated as “I am a primate; nothing about primates is outside of my [[bailiwick]]” and has become a logo accepted by [[primatologists]] worldwide.<ref>{{cite book | author = Hooton, E. | authorlink=Earnest Hooton | year = 1955 | chapter = The Importance of Primate Studies in Anthropology | editor = Gavan, J.A. | title = The Non-Human Primates and Human Evolution | publisher = Wayne University Press, Detroit | pages = pp. 1-10 |oclc=557632}}</ref>
==See also==
*[[List of placental mammals in Order Primates|List of primates]]
*[[List of prehistoric mammals#Order Primates|List of prehistoric primates]]
*[[Arboreal theory]]
*[[Primatology]]
==References==
{{reflist|colwidth=30em}}
==External links==
{{Sisterlinks|Primate}}
{{Wikispecies|Primates}}
{{Wikibookspar|Dichotomous Key|Primates}}
* [http://pin.primate.wisc.edu Primate Info Net]
<!-- * [http://www.buav.org/campaigns/primates/index.html Primates in scientific experimentation]{{deadlink}} -->
* [http://www.pri.kyoto-u.ac.jp/ Primate Research Institute, Kyoto University]
<!-- * [http://images.google.com/imgres?imgurl=http://webdrive.service.emory.edu/groups/research/chimpanzee-cognition/CCL/ethogram/spec_laugh.jpg&imgrefurl=http://webdrive.service.emory.edu/groups/research/chimpanzee-cognition/CCL/ethogram.htm&h=382&w=784&sz=94&hl=en&start=6&tbnid=SzTd6o7BvU2mUM:&tbnh=70&tbnw=143&prev=/images%3Fq%3DChimpanzee%2Blaughter%26svnum%3D10%26hl%3Den%26lr%3D%26rls%3DHPIA,HPIA:2006-24,HPIA:en%26sa%3DN Chimpanzee Facial Expression & Vocalizations]{{deadlink}} -->
* [http://primate-brain.org High-Resolution Cytoarchitectural Primate Brain Atlases]
* [http://www.euprim-net.eu EUPRIM-Net: European Primate Network]
* [http://realaudio.service.emory.edu/ramgen/YERKES/PARR/laugh.rm A chimpanzee laughter sample]
* [http://digital.library.wisc.edu/1711.dl/PCLNatHist PrimateImages: Natural History Collection]
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