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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 best-known of the prosimians, the [[lemur]]s, are located on the [[Madagascar|island of Madagascar]] and to a lesser extent on the [[Comoros|Comoros Islands]], isolated from the rest of the world. The New World monkeys, which include the familiar [[Capuchin monkey|capuchin]], [[howler monkey|howler]], and [[squirrel monkey]]s, live exclusively in the Americas. With the exception of humans, the rest of 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.
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 best-known of the prosimians, the [[lemur]]s, are located on the [[Madagascar|island of Madagascar]] and to a lesser extent on the [[Comoros|Comoros Islands]], isolated from the rest of the world. The New World monkeys, which include the familiar [[Capuchin monkey|capuchin]], [[howler monkey|howler]], and [[squirrel monkey]]s, live exclusively in the Americas. With the exception of humans, the rest of 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.


Primates are adapted for a tree-dwelling lifestyle. Anatomical adaptations support their reliance on vision, their dominant sensory system, rather than smell, which is the dominant sensory system in most mammals. In some primates, [[Trichromacy|three color vision]] has developed. Most primates also have [[thumb|opposable thumb]]s and some have [[Prehensility|prehensile]] tails. Many species are [[Sexual dimorphism|sexually dimorphic]], in that males and females have different physical traits, including body mass, canine tooth size, and coloration. Primates have slower development rates than other similarly sized mammals, and reach maturity later, but have longer life spans.
Primates are adapted for a tree-dwelling lifestyle. Anatomical adaptations support their reliance on vision, their dominant sensory system, rather than smell, which is the dominant sensory system in most mammals. In some primates, [[Trichromacy|three color vision]] has developed. Most primates also have [[thumb|opposable thumb]]s and some have [[Prehensility|prehensile]] tails. Many species are [[Sexual dimorphism|sexually dimorphic]], in that males and females have different physical traits, including body mass, canine tooth size, and coloration. Primates have slower development rates than other similarly sized mammals, and reach maturity later, but have longer life spans. A variety of locomotion techniques are used, including leaping from tree to tree, walking on two or four limbs, knuckle-walking and swinging between branches of trees (known as [[brachiation]]). Similarly, primates use a variety of social systems. Some species are solitary, others are [[Monogamy|monogamous]], while others live in groups that can include up to hundreds of members.


⚫
According to fossil evidence, primitive ancestors of primates already existed in the late [[Cretaceous]] period. [[Molecular clock]] studies suggest that the primate branch is even more ancient (originating at least in the mid-Cretaceous). Primates are now thought to be most closely related to [[Colugo|flying lemur]]s and, more distantly, to [[treeshrew]]s. Primates probably descended from [[Plesiadapiformes]].{{Fact|date=August 2008}}
Primates use a variety of locomotion techniques. These include leaping from tree to tree, walking on two or four limbs, knuckle-walking and swinging between branches of trees (known as [[brachiation]]). Similarly, primates use a variety of social systems. Some species are solitary, others are [[Monogamy|monogamous]], while others live in groups that can include up to hundreds of members. Among group living primates, in some species members of one gender typically remain with their natal group for their entire lives while members of the other transfer to a new group upon reaching sexual maturity. In other species both sexes leave their natal group to form or find new groups upon reaching sexual maturity.

⚫
According to fossil evidence, primitive ancestors of primates already existed in the late [[Cretaceous]] period. [[Molecular clock]] studies suggest that the primate branch is even more ancient (originating at least in the mid-Cretaceous). Primates are now thought to be most closely related to [[Colugo|flying lemur]]s and, more distantly, to [[treeshrew]]s. Primates probably descended from [[Plesiadapiformes]].




Revision as of 16:02, 9 August 2008

For other uses, see Primate (disambiguation).

Primates[1]
Temporal range: Late Cretaceous – recent
Olive Baboon, an Old World monkey
Scientific classification
Kingdom:
Animalia
Phylum:
Chordata
Class:
Mammalia
Infraclass:
Eutheria
Superorder:
Euarchontoglires
Order:
Primates

Linnaeus, 1758
Families
Range of the non-human primates

A primate is any member of the biological order Primates (Latin: "prime, first rank"[2]), the group that contains all the species commonly related to the lemurs, monkeys, and apes, with the last category including humans.[3] 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: prosimians, monkeys of the New World, and monkeys and apes of the Old World. The prosimians are species whose bodies most closely resemble that of the early proto-primates. The best-known of the prosimians, the lemurs, are located on the island of Madagascar and to a lesser extent on the Comoros Islands, isolated from the rest of the world. The New World monkeys, which include the familiar capuchin, howler, and squirrel monkeys, live exclusively in the Americas. With the exception of humans, the rest of the Old World monkeys and the apes inhabit Africa and southern and central Asia, although fossil evidence shows many species existed in Europe as well.

Primates are adapted for a tree-dwelling lifestyle. Anatomical adaptations support their reliance on vision, their dominant sensory system, rather than smell, which is the dominant sensory system in most mammals. In some primates, three color vision has developed. Most primates also have opposable thumbs and some have prehensile tails. Many species are sexually dimorphic, in that males and females have different physical traits, including body mass, canine tooth size, and coloration. Primates have slower development rates than other similarly sized mammals, and reach maturity later, but have longer life spans. A variety of locomotion techniques are used, including leaping from tree to tree, walking on two or four limbs, knuckle-walking and swinging between branches of trees (known as brachiation). Similarly, primates use a variety of social systems. Some species are solitary, others are monogamous, while others live in groups that can include up to hundreds of members.

According to fossil evidence, primitive ancestors of primates already existed in the late Cretaceous period. Molecular clock studies suggest that the primate branch is even more ancient (originating at least in the mid-Cretaceous). Primates are now thought to be most closely related to flying lemurs and, more distantly, to treeshrews. Primates probably descended from Plesiadapiformes.[citation needed]


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.[4] They are distinguished by the retention of the collar bone in the pectoral girdle; shoulder joints which allow high degrees of movement in all directions; five digits on the fore and hind limbs with opposable thumbs and big toes; fingernails; sensitive tactile pads on the ends of the digits; a trend towards reduced snout and sense of smell; 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 glands; typically one young per pregnancy; a long gestation and developmental period and a trend toward holding the torso upright leading to bipedalism.[4] Primates are frequently highly social, with flexible dominance hierarchies.[5] New World species form monogamous pair bonds, and show substantial paternal care of young unlike most Old World monkeys.[6]

Anatomy, physiology and morphology

Primate skull size and weight comparison

Primates have two forward-facing eyes on the front of the skull; binocular vision allows accurate distance perception, useful for the brachiating ancestors of humans.[4] There is a bony ridge located above the eye sockets which reinforces weaker bones in the face which are put under strain during chewing. Strepsirrhines have a postorbital bar, a bone which runs around the eye socket, to protect their eyes, which is in contrast to the higher primates, haplorrhines, which have evolved fully enclosed sockets.[7]

The primate skull has a large domed cranium which is particularly prominent in anthropoids. The cranium protects the large brain, a distinguishing characteristic of this group.[4] The endocranial volume (the volume within the skull) is three-fold greater in humans than in the greatest non-human primate, reflecting a larger brain size.[8] The mean human endocranial volume is 1201 cubic centimetres, it is 469 cm³ in gorillas, 400 cm³ in chimpanzees and 397 cm³ in orangutans.[8] The primary evolutionary trend of primates has been the elaboration of the brain, in particular the neocortex (a part of the cerebral cortex) which is involved with sensory perception, generation of motor commands, spatial reasoning, conscious thought and, in humans, language.[9] While other mammals rely heavily on their sense of smell, the arboreal life of primates has led to a tactile, visually dominant sensory system,[9] a reduction in the olfactory region of the brain and increasingly complex social behavior.[10]

An 1893 drawing of the hands and feet of primates

Primates generally have five digits on each limb (pentadactyly), with keratin nails on the anterior ends. The bottom sides of the hands and feet have senstive pads on the finger tips. Most have opposable thumbs, which are a characteristic primate feature, but are not limited to this order; opossums, for example, also have opposing thumbs.[4] Thumbs allow some species to use tools to perform some tasks. In primates, the combination of opposing thumbs, short fingernails (rather than claws) and long, inward-closing fingers is a relic of the ancestral practice of gripping branches, and has, in part, allowed some species to develop brachiation (swinging by the arms from tree limb to tree limb) as a significant means of transportation. Prosimians have clawlike nails on the second toes of their feet.[4]

The primate collar bone is retained as prominent element of the pectoral girdle; this allows the shoulder joint broad mobility.[5] 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 lower 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. Only the New World Atelidae family have prehensile tails.

Primates show a trend towards a reduced snout.[5] 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 the arrangement of their teeth.[10] In New World monkeys the nostrils face sideways, while in Old World monkeys, they face downwards.[10] There is a considerably varied dental pattern in primates and although some have lost most of their incisors, all retain at least one lower incisor.[10] In most strepsirhines, the lower incisors form a toothcomb which is thought to be used in grooming and possibly foraging.[10] Old World monkeys also have eight premolars, compared with twelve in New World monkeys.[10] The Old World species are divided into apes and monkeys depending on the number of cusps on their molars; apes have five, Old World monkeys have four.[10] The main hominid molar cusp (hypocone) evolved in early primate history, while the cusp of the corresponding primitive lower molar (paraconid) was lost. Prosimians are distinguished by their immobilized upper lips, moist tip to their nose and forward-facing lower front teeth.

Sexual dimorphism

Distinct sexual size dimorphism can be seen between the male Hamadryas Baboons (grey) and the female (brown)
Main article: Sexual dimorphism in non-human primates

Sexual dimorphism is often exhibited in simians; 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[11][12] and canine tooth size[13][14] along with pelage and skin color.[15] The dimorphism can been attributed to and affected by different factors; mating system,[16] size,[16] habitat and diet.[17]

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.[18] 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.[19] There is some evidence from the fossil record that suggests that there was convergent evolution of dimorphism, and some extinct hominids probably exceeded dimorphism of any living primate.

Color vision

Main article: 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 three color vision, our nocturnal, warm-blooded, mammalian ancestors lost one of three cones in the retina at the time of dinosaurs. Fish, reptiles and birds are therefore trichromatic while all mammals, with the exception of some primates and marsupials,[20] are strictly dichromats.

Primates achieve trichromacy through color receptors, with spectral peaks in the violet (short wave, 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 opsins (pigments) sensitive to 430 nanometre, 530 nm, and 560 nm wavelengths.[21] Platyrrhines, on the other hand are non-routinely trichromatic; only a small population of platyrrhines are trichromats.[22]

Howler monkeys have reinvented routine trichromatism through a recent gene duplication of the red-green opsin gene.[23] 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 a major part of their diet[24], 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.[22]

Locomotion

Various species of primates move by brachiation, bipedalism, leaping, arboreal and terrestrial quadrupedalism, climbing or knuckle-walking.

Several prosimians are primarily vertical clinger and leapers. These include many bushbabies[25], all Indriids[26] (i.e., sifakas, avahis and indris), sportive lemurs[27], and all tarsiers.[28]

Other prosimians are arboreal quadrupedalists and climbers. Some are also terrestrial quadrupedalists, while some are leapers. Most monkeys are both arboreal and terrestrial quadrupedalists and climbers. Gibbons, muriquis and spider monkeys all use brachiation extensively.[29] Woolly monkeys also sometimes brachiate.[30] Orangutans use a similar form of locomotion called quadramanous climbing, in which they use their arms and legs to carry their heavy bodies through the trees.[29] Chimpanzees and gorillas knuckle walk[29], and can move bipedally for short distances. Humans are the only fully bipedal species.

Behavior

Further information: Primatology

Social systems

Japanese Macaques bathe together in Jigokudani Hot Spring

Richard Wrangham proposed that social systems are best classified by the amount of movement by females occurring between groups.[31] 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. Among New World Monkeys, spider monkeys and muriquis use this system.[32]
  • 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. This system is common among the Ring-tailed Lemur, capuchin monkeys and many Old World Monkeys (the cercopithecines).[33]
  • 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. Gibbons essentially use this system, although "monogamy" in this context does not necessarily mean absolute sexual fidelity.[34]
  • 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 effectively have this organisation.[35]

Some other systems are known to occur as well. For example, with howler monkeys both the males and females typically transfer from their natal group upon reaching sexual maturity, resulting in groups where neither the males or females are typically related.[36] Some prosimians, colobine monkeys and callitrichid monkeys also use this system.[33]

Primatologist Dr. Jane Goodall, who studied in the Gombe Stream National Park, noted fission-fusion societies in chimpanzees.[37] 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[38], spider monkeys[39] and Bonobos.[40] Geladas have a similar social structure in which many smaller groups come together to form temporary herds of up to 600 monkeys.[41]

These social systems are affected by three main ecological factors; distribution of resources, group size and predation.[6] Within a social group there is a balance between cooperation and competition. Cooperation comes in the form of allogrooming; whereby ectoparasites 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 hierarchies.[6]

Development through life stages

Primates have slower rates of development than other mammals.[42] All non-human primate infants rely on their mothers for nursing.[43] Infants are rely on their mothers for grooming and transportations.[43] In some species, infants are also protected and transported by males in the group, particularly males who may be their father.[43] Other relatives of the infant, such as siblings and aunts, may also participate in its care.[43] Most primate mothers cease ovulation while nursing an infant so the sooner the infant is weaned the sooner the mother can reproduce again.[44] This often leads to weaning conflict with the infant, who often resist weaning and attempt to continue nusring beyond the point where the mother wishes to cease.[44]

Primates have a longer juvenile period between weaning and sexual maturity for their size than other mammals.[45] During the juvenile period, primates are more susceptible than adults to predation and starvation.[45] During the juvenile period, primates gain experience in feeding and avoiding predators.[45] They also learn social and fighting skills, often through playing.[45]

In addition to reaching maturity later, primates have longer life spans than other similarly sized mammals.[46] Life spans are generally longer for female primates than for males.[46]

Diet and feeding

Primates species exploit a variety of food sources. Most primates include fruit in their diets to obtain easily digested carbohydrates and lipids for energy.[47] But they require other foods, such as leaves or insects, to obtain amino acids, vitamins and minerals. Many primates have anatomical specializations enabling them to exploit particular food specialties, such as fruit, leaves, gum or insects.[47] For example, leaf eaters such as howler monkeys, colobus monkeys and sportive lemurs, have extended degestive tracts to enable them to absorb nutrients from leaves that can be difficult to digest.[47] Marmosets, which are gum eaters, have stout incisor teeth, enabling them to open tree bark to get to the gum, and claws rather than nails, enabling them to cling to trees while feeding.[47] Some species have additional specializations. For example, grey-cheeked mangabeys have thick enamel on their teeth, enabling them to open hard fruits and seeds that other monkeys cannot.[47]

Some species are more or less specialized than described above. Geladas are the only primate species that feeds primarily on grass.[48] Capuchin monkeys, on the other hand, can exploit many different types of food, including fruit, leaves, flowers, buds, nectar, seeds, insects, other invertebrates, and small vertebrates such as birds, bird eggs, lizards, squirrels and bats.[49]

Cognition and communication

Further information: Great ape language, Primate empathy, Hominid intelligence, Chimpanzee intelligence, and Animal intelligence

Lemurs, lorises, tarsiers, and New World monkeys are reliant on olfactory signals for many aspects of social and reproductive behavior.[9] Specialized glands are used to mark territories with pheromones, which are detected by the vomeronasal organ; this process forms a large part of the communication behaviour of these primates.[9] In Old World monkeys and apes this ability is mostly vestigial, and regressed as color vision evolved to become the main sensory organ.[50] Primates also use vocalizations, gestures, and facial expressions to convey psychological state.[51]

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.

Chimpanzees, 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.[52]

Habitat and distribution

Rhesus Macaque in Indian city of Agra

Primates evolved from arboreal animals and many species live most their lives in trees. Some species are partially terrestrial, such as baboons and the Patas Monkey and a few species are fully terrestrial, such as the Gelada and Humans. Non-human primates live in a diverse number of forested habitats in the tropical latitudes of Africa, India, Southeast Asia, and South America, including rainforests, mangrove forests, and montane forests. 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 Mountains at altitudes of 4,700 metres (15,400 ft),[53] Mountain Gorillas can be found at 4,200 metres (13,200 ft) crossing the Virunga Mountains[54] 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 gray langurs, can exploit human-modified environments and even live in cities.[55][56]

Interspecific associations

Several species of primates are known to associate in the wild. Some of these associations have been extensively studied. In the Tai Forest of Africa, several species coordinate anti-predator behavior. These include the Diana Monkey, Campbell's Mona Monkey, Lesser Spot-nosed Monkey, Western Red Colobus, King Colobus and Sooty Mangabey, which coordinate anti-predator alarm calls.[57] Among the predators of these monkeys is the Chimpanzee.[58]

Red-tailed Monkeys associate with several species, including the Western Red Colobus, Blue Monkey, Wolf's Mona Monkey, Mantled Guereza, Black Crested Mangabey and Allen's Swamp Monkey.[59] Several of these species are also predated on by Chimpanzees.[60]

In South America, but not in Central America, squirrel monkeys associate with capuchin monkeys.[61] This may have more to do with foraging benefits to the squirrel monkeys rather than anti-predation benefits.[61]

Conservation status

The IUCN lists more than a third of its primates as critically endangered, endangered or vulnerable. Deforestation, forest fragmentation, primate crop raiding, and primate hunting for use in medicines, as pets, and for food are cited as common threats to primate species. Large-scale tropical forest clearance is widely regarded as the process that most threatens primates.[62][63][64] More than 90% of primate species occur in tropical forests.[65][63] The main cause of forest loss is clearance for agriculture, although commercial logging, subsistence harvesting of timber, mining, and dam construction also contribute to tropical forest depletion.[65] In Indonesia large areas of lowland forest have been cleared to increase palm oil production, one analysis of satellite imagery concluded that during 1998 and 1999 there was a loss of 1,000 Sumatran Orangutans per year in the Leuser Ecosystem alone.[66]

Primates with a large body size (over 5 kg) have an increased extinction risk due to their increased profitability to poachers compared to smaller primates.[65] They also have a slow life history with an increased sexual maturity age and greater period between births. Populations therefore have a slower recovery time after the loss of members to poaching or the pet trade.[67] In some African cities estimates suggest that half of all protein consumed in urban areas comes from the bushmeat trade.[68] Endangered primates such as guenons and the Drill are hunted at levels that far exceed sustainable levels.[68] This is due to their large body size, ease of transport and profitability per animal.[68] As farming encroaches on forest habitats, primates feed on the crops causing the farmers large economic losses.[69] Primate crop raiding gives locals a negative impression of primates, hindering conservation efforts.[70]

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.[9] Among the primates wiped out were Archaeoindris, a lemur larger than a silverback gorilla and the families Palaeopropithecidae and Archaeolemuridae.[9]

In Asia, Hinduism, Buddhism, and Islam prohibit eating primate meat, despite this primates are still hunted for food.[65] Some smaller traditional religions allow the consumption of primate meat.[71][72] The pet trade and traditional medicine also increase demand for illegal hunting.[73][74][75] The expanding Chinese middle class have also increased demand for exotic pets in recent years.[75] The Rhesus Macaque, a model organism, was protected after overtrapping threatened numbers in the 1960s.[64] The program was so effective that the macaques are now seen as a pest throughout their range.[64]

In Central and South America forest fragmentation and hunting are the two main problems for primates. Large tracts of forest are now rare in Central America.[76][62] This increases the amount of forest vulnerable to edge effects such as farmland encroachment, lower levels of humidity and a change in plant life.Marsh, 2003; Turner, 1996). Movement restriction results in a greater amount of inbreeding, which can cause deleterious effects leading to a population bottleneck, whereby 50% of the population is lost.[77][78]

In the United States the main threat is the pet trade. Although primate import for the pet trade was banned in 1975, smuggling still occurs along the United States–Mexico border.[79] Prices range from $3000 for monkeys to $30,000 for apes.[79]

There are 21 critically endangered primates, 8 of these species have remained on the IUCNs "The World's 25 Most Endangered Primates" list since the year 2000: the Silky Sifaka, Delacour's Langur, the White-headed Langur, the Gray-shanked Douc, the Tonkin Snub-nosed Langur, the Hainan Gibbon, the Cross River Gorilla and the Sumatran Orangutan.[80] 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.[81] It was last sighted in 1978 and was officially declared extinct in 2000.[82]

Evolutionary history

Euarchontoglires
Glires

Rodentia (rodents)

Lagomorpha (rabbits, hares, pikas)

Euarchonta

Scandentia (treeshrews)

Dermoptera (colugos)

†Plesiadapiformes

Primates

The Primate order lies in a tight clustering of related orders (the Euarchontoglires) within the Eutheria, a subclass of Mammalia. Recent molecular genetic research on primates, colugos, and treeshrews has shown that the two species of colugos are more closely related to the primates than the treeshrews, 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 Rodentia 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.[83]

Evolution

Ring-tailed Lemur, a strepsirrhine primate

In modern, cladistic reckonings, the Primate order is also a true clade. The suborder Strepsirrhini, the "wet-nosed" primates, split off from the primitive primate line about 63 million years ago (mya). The seven strepsirhine families are the four related lemur families and the three remaining families that include the lorises, the Aye-aye, the galagos, and the pottos.[1] 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.[1] Other lineages of lower primates inhabited Earth. During the Eocene, most of the northern continents were dominated by two dominant groups, the adapids and the omomyids. The former is considered a member of Strepsirrhini, but it does not have a toothcomb 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.[1] 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.

Emperor Tamarin, a New World monkey

The suborder Haplorrhini, the "dry-nosed" primates, is composed of two sister clades.[1] 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 monkeys in one, and the Old World monkeys, humans and the other apes in the other.[1] 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.[84] The recently published third edition of Mammal Species of the World (MSW3) lists 376 species.[1] 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 British actor and lemur enthusiast John Cleese) and the GoldenPalace.com Monkey (whose name was put up for auction).

Classification

A 1927 drawing of chimpanzees, a gibbon (top right) and two orangutans (center and bottom center)
Homo sapiens, a member of the order Primates

The order Primates was established by Carl Linnaeus in 1758, in the tenth edition of his book Systema Naturae,[85] for the genera Homo (humans), Simia (other apes and monkeys), 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).[86] 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),[87] 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 prosimians: all of Strepsirrhini plus the tarsiers. The Anthropoidea contained all of the simians.

Hybrids

Primate hybrids usually arise in captivity[88] though there have been examples of primate hybrids in the wild.[89] Hybridisation occurs where two species' territories overlap, these hybrid zones may be created by humans when animals are placed in zoos or may be due to environmental pressures such as predation.[89] In The variation of animals and plants under domestication Charles Darwin noted: "Several members of the family of Lemurs have produced hybrids in the Zoological Gardens."[90]

Many gibbons are hard to identify based on fur coloration and are identified either by song or genetics.[88] 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.[91]

Intergeneric hybridizations, hybrids of different genera, have been found in the wild. Though belonging to genera that have been distinct for several million years interbreeding still occurs between Geladas (Theropithecus gelada) and Hamadryas Baboon (Papio hamadryas).[92] Red-tailed Monkeys (Cercopithecus ascanius) are known to hybridize with Blue Monkeys (C. mitis) in several locations in the wild in Africa.[59] Hybrids between Müller's Bornean Gibbons (Hylobates muelleri) and Agile Gibbons (Hylobates agilis) have been reported in areas of Borneo.[93] And in India, Gray langurs (Semnopithecus sp) are known to hybridize with Nilgiri Langurs (Trachypithecus johnii).[56]

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 primates used in animal research or kept as 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.[94]

Role in scientific research

File:Caged monkeys.jpg
Monkeys, imported for experimentation, in a crate.

NHPs are commonly used in scientific research as 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 preclinical trials, neuroscience, and ophthalmology 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. Chimpanzees, baboons, marmosets, and green monkeys are also commonly found in NHP research labs.[95] 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.[96] As of 2004, 3,100 NHPs were living in captivity in the United States, in zoos, circuses, and laboratories; 1,280 of them being used in experiments.[97] 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 persons and protected in law by the United Nations Universal Declaration of Human Rights.[98] The status of NHPs is a point of debate, particularly from organizations such as the Great Ape Project[97] and activists such as Ignaas Spruit[99] (director of the Pro-Primates organization), which argue for increased legal rights of NHPs.

See also

References

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  87. ^ Blainville, Henri-Marie Ducrotay de (1839). "Nouvelle classification des Mammifères". Annales Françaises et Etrangères d’Anatomie et de Physiologie Appliquées à la Médicine et à l’Histoire Naturelle, 3. pp. pp. 268-269. {{cite book}}: |pages= has extra text (help)
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  91. ^ Sugawara, K. (1979). "Sociological study of a wild group of hybrid baboons between Papio anubis and P. hamadryas in the Awash Valley, Ethiopia". Primates. 20 (1): 21–56. doi:10.1007/BF02373827.
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  98. ^ Article 6: Everyone has the right to recognition everywhere as a person before the law.
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