Temporal range: Late Paleocene–recent
Olive baboon, Papio anubis Scientific classification Kingdom: Animalia Phylum: Chordata Class: Mammalia Infraclass: Eutheria Superorder: Euarchontoglires Order: Primates
- 16, See classification
Range of the non-human primates (green)
A primate ( // US dict: prī′·māt) is a mammal of the order Primates (// US dict: prī·mā′·tēz; Latin: "prime, first rank"), which contains prosimians and simians. Primates arose from ancestors that lived in the trees of tropical forests; many primate characteristics represent adaptations to life in this challenging three-dimensional environment. All but a few primate species remain at least partly arboreal.
With the exception of humans, who inhabit every continent,[a] most primates live in tropical or subtropical regions of the Americas, Africa and Asia. Primates range in size from Madame Berthe's mouse lemur, which weighs only 30 grams (1 oz), to the mountain gorilla, weighing 200 kilograms (440 lb). According to fossil evidence, the primitive ancestors of primates may have existed in the late Cretaceous period around 65 million years ago; the oldest known primate is the Late Paleocene Plesiadapis, c. 55–58 million years ago. Molecular clock studies suggest that the primate branch may be even older, originating in the mid-Cretaceous period around 85 mya.
Order Primates has traditionally been divided into two main groupings: prosimians and anthropoids (simians). Prosimians have characteristics more like those of the earliest primates, and include the lemurs of Madagascar, lorisiforms and tarsiers. Simians include the monkeys, apes and humans. More recently, taxonomists have preferred to split primates into the suborder Strepsirrhini, or curly-nosed primates, consisting of non-tarsier prosimians, and the suborder Haplorhini, or dry-nosed primates, consisting of tarsiers and the simians. Simians are divided into two groups: platyrrhine ("flat nosed") or New World monkeys of South and Central America and catarrhine (narrow nosed) monkeys and apes of Africa and southeastern Asia. New World monkeys include the capuchin, howler and squirrel monkeys; catarrhines consist of Old World monkeys (such as baboons and macaques), gibbons and great apes. Humans are the only extant catarrhines that have spread successfully outside of Africa, South Asia, and East Asia, although fossil evidence shows many other species were formerly present in Europe.
Considered generalist mammals, primates exhibit a wide range of characteristics. Some primates (including some great apes and baboons) are primarily terrestrial rather than arboreal, but all species possess adaptations for climbing trees. Locomotion techniques used include leaping from tree to tree, walking on two or four limbs, knuckle-walking, and swinging between branches of trees (known as brachiation). Primates are characterized by large brains relative to other mammals, as well as an increased reliance on stereoscopic vision at the expense of smell, the dominant sensory system in most mammals. These features are more developed in monkeys and apes and noticeably less so in lorises and lemurs. Three-color vision has developed in some primates. Most also have opposable thumbs and some have prehensile tails. Many species are sexually dimorphic, which means males and females have different physical traits, including body mass, canine tooth size, and coloration. Primates have slower rates of development than other similarly sized mammals and reach maturity later but have longer lifespans. Depending on the species, adults may live in solitude, in mated pairs, or in groups of up to hundreds of members.
Historical and modern terminology
The relationships among the different groups of primates were not clearly understood until relatively recently, so that commonly used terms are somewhat confused. For example, "ape" has been used either as an alternative for "monkey" or for any tailless, relatively humanlike primate.
Sir Wilfrid Le Gros Clark was one of the primatologists who developed the idea that there were trends in primate evolution, and that the living members of the order could be arranged in an ascending series leading to humans. Commonly used names for groups of primates, for example "prosimians", "monkeys", "lesser apes" and "great apes", reflect this arrangement. According to our current understanding of the evolutionary history of the primates, several of these groups are paraphyletic, meaning that although all the species in the group descend from a common ancestor, the group does not include all the descendants of that ancestor.
By contrast with approaches such as those of Le Gros Clark, modern classifications typically use groups which are monophyletic, since they include all the descendants of a common ancestor. The diagram below shows one currently accepted classification of the primates, with groups with commonly used names shown on the right.
Primates Haplorhini Simiiformes Catarrhini Hominoidea Hominidae Homininae Hominini
humans (genus Homo)
chimpanzees (genus Pan)
gorillas (tribe Gorillini)
orangutans (subfamily Ponginae)
gibbons (family Hylobatidae)
Old World monkeys (superfamily Cercopithecoidea)
New World monkeys (parvorder Platyrrhini)
tarsiers (infraorder Tarsiiformes)
lemurs (infraorder Lemuriformes)
lorises and allies (infraorder Lorisiformes)prosimiansmonkeysgreat apeshumanslesser apes
All the groups which have scientific names are monophyletic (i.e. they are clades), so that the scientific classification reflects evolutionary history. Some of the traditional groups shown on the right, which form an "ascending series", are paraphyletic:
- "prosimians" contain two monophyletic groups, the suborder Strepsirrhini (lemurs, lorises and allies), as well as the tarsiers, which are a sister group to the infraorder Simiiformes
- "monkeys" consist of two monophyletic groups, New World monkeys and Old World monkeys, but exclude hominoids (superfamily Hominoidea)
- "apes" as a whole, and the "great apes" in particular, are not monophyletic because they exclude humans.
Thus the two sets of groups, and hence names, do not match, which causes problems in relating scientific names to common names. Consider the superfamily Hominoidea. In terms of the common names on the right, this group consists of apes and humans, and there is no single common name for all the members of the group. One possibility is to create a new common name, in this case "hominoids". Another possibility is to expand the use of one of the traditional terms. For example, in a 2005 book, the vertebrate palaeontologist Benton wrote "The apes, Hominoidea, today include the gibbons and orang-utan ... the gorilla and chimpanzee ... and humans", thereby using "apes" to mean "hominoids". The group traditionally called "apes" must then be called the "nonhuman apes".
As of July 2011[update], there is no consensus as to which approach to follow, whether to accept traditional paraphyletic common names or whether to use monophyletic names, either new ones or adaptations of old ones. Both approaches will be found in biological sources, often in the same work. Thus although Benton defines "apes" to include humans, he also repeatedly uses "ape-like" to mean "like an ape rather than a human", and when discussing the reaction of others to a new fossil writes of "claims that Orrorin ... was an ape rather than a human".
Lagomorpha (rabbits, hares, pikas)
Order Primates is part of the clade Euarchontoglires, which is nested within the clade Eutheria of Class Mammalia. Recent molecular genetic research on primates, colugos, and treeshrews has shown that the two species of colugos are more closely related to primates than treeshrews, even though treeshrews were at one time considered primates. These three orders make up the clade Euarchonta. The combination of this clade with the clade Glires (composed of Rodentia and Lagomorpha) forms the clade Euarchontoglires. Variously, both Euarchonta and Euarchontoglires are ranked as superorders. Some scientists consider Dermoptera a suborder of Primates and call the "true" primates the suborder Euprimates.
The primate lineage is thought to go back at least 65 mya (million years ago), even though the oldest known primate from the fossil record is Plesiadapis (c. 55–58 mya) from the Late Paleocene. Other studies, including molecular clock studies, have estimated the origin of the primate branch to have been in the mid-Cretaceous period, around 85 mya.
By modern cladistic reckoning, Order Primates is monophyletic. Suborder Strepsirrhini, the curly-nosed or "wet-nosed" primates, is generally thought to have split off from the primitive primate line about 63 mya, although earlier dates are also supported. The seven strepsirhine families are the five related lemur families and the two remaining families that include the lorisids and the galagos. Older classification schemes wrap Lepilemuridae into Lemuridae and Galagidae into Lorisidae, yielding a four-one family distribution instead of five-two as presented here. During the Eocene, most of the northern continents were dominated by two groups, the adapiforms and the omomyids. The former are considered members of Strepsirrhini, but did not have a toothcomb like modern lemurs; recent analysis has suggested Darwinius masillae fits into this grouping. The latter was related closely to tarsiers, monkeys, and apes. It is unclear exactly how these two groups relate to extant primates. Omomyids perished about 30 mya, while adapids survived until about 10 mya.
According to genetic studies, the lemurs of Madagascar diverged from the lorisiforms approximately 75 mya. These studies, as well as chromosomal and molecular evidence, also show that lemurs are more closely related to each other than to other strepsirrhine primates. However, Madagascar split from Africa 160 mya and from India 90 mya. To account for these facts, it is thought that a founding lemur population of a few individuals reached Madagascar from Africa via a single rafting event between 50 and 80 million years ago. Other colonization options have been examined, such as multiple colonizations from Africa and India, but none are supported by the genetic and molecular evidence.
Until recently the aye-aye has been difficult to place within Strepsirrhini. Theories had been proposed that its family, Daubentoniidae, was either a lemuriform primate (meaning its ancestors split from the lemur line more recently than lemurs and lorises split) or a sister group to all the other strepsirrhines. In 2008, the aye-aye family was confirmed to be most closely related to the other Malagasy lemurs, likely having descended from the same ancestral population that colonized the island.
Suborder Haplorhini, the simple-nosed or "dry-nosed" primates, is composed of two sister clades. Prosimian tarsiers in the family Tarsiidae (monotypic in its own infraorder Tarsiiformes), represent the most basal division, originating about 58 mya. The infraorder Simiiformes emerged about 40 mya, and contains two clades, both parvorders: Platyrrhini, which developed in South America, consisting of New World monkeys, and Catarrhini, which developed in Africa, consisting of Old World monkeys, humans and the other apes. A third clade, which included the eosimiids, developed in Asia but went extinct millions of years ago.
As in the case of lemurs, the origin of New World monkeys is unclear. Molecular studies of concatenated nuclear sequences have yielded a widely varying estimated date of divergence between platyrrhines and catarrhines, ranging from 33 to 70 mya, while studies based on mitochondrial sequences produce a narrower range of 35 to 43 mya. It is possible that anthropoid primates traversed the Atlantic ocean from Africa to South America during the Eocene by island hopping, facilitated by Atlantic Ocean ridges and a lowered sea level. Alternatively, a single rafting event may explain this transoceanic colonization. Due to continental drift, the Atlantic Ocean was not nearly as wide at the time as it is today. Research suggests that a small 1 kg (2.2 lb) primate could have survived 13 days on a raft of vegetation. Given estimated current and wind speeds, this would have provided enough time to make the voyage between the continents.
Apes and monkeys spread from Africa into Europe and Asia starting in the Miocene. Soon after, the lorises and tarsiers made the same journey. The first hominid fossils were discovered in Northern Africa and date back 5–8 mya. Old World monkeys disappeared from Europe about 1.8 mya. Molecular and fossil studies generally show that modern humans originated in Africa 100,000–200,000 years ago.
Although primates are well studied in comparison to other animal groups, several new species have been recently discovered, and genetic tests have revealed previously unrecognised species in known populations. Primate Taxonomy listed about 350 species of primates in 2001; the author, Colin Groves, increased that number to 376 for his contribution to the third edition of Mammal Species of the World (MSW3). However, publications since the taxonomy in MSW3 was compiled in 2003 have pushed the number to 424 species, or 658 including subspecies.
Classification of living primates
- Order Primates
- Suborder Strepsirrhini: non-tarsier prosimians
- Infraorder Lemuriformes
- Infraorder Lorisiformes
- Family Lorisidae: lorises, pottos and allies (9 species)
- Family Galagidae: galagos (19 species)
- Suborder Haplorhini: tarsiers, monkeys and apes
- Infraorder Tarsiiformes
- Family Tarsiidae: tarsiers (9 species)
- Infraorder Simiiformes
- Parvorder Platyrrhini: New World monkeys
- Family Callitrichidae: marmosets and tamarins (42 species)
- Family Cebidae: capuchins and squirrel monkeys (17 species)
- Family Aotidae: night or owl monkeys (douroucoulis) (10 species)
- Family Pitheciidae: titis, sakis and uakaris (42 species)
- Family Atelidae: howler, spider, woolly spider and woolly monkeys (28 species)
- Parvorder Catarrhini
- Parvorder Platyrrhini: New World monkeys
- Infraorder Tarsiiformes
- Suborder Strepsirrhini: non-tarsier prosimians
Order Primates was established by Carl Linnaeus in 1758, in the tenth edition of his book Systema Naturae, 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). 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), but these new taxa were not accepted.
Before Anderson and Jones introduced the classification of Strepsirhini and Haplorhini in 1984, (followed by McKenna and Bell's 1997 work Classification of Mammals: Above the species level), the Primates were divided into two superfamilies: Prosimii and Anthropoidea. Prosimii included all of the prosimians: Strepsirrhini plus the tarsiers. Anthropoidea contained all of the simians.
Primate hybrids usually arise in captivity, but there have also been examples in the wild. Hybridization occurs where two species' range overlap to form hybrid zones; hybrids may be created by humans when animals are placed in zoos or due to environmental pressures such as predation. Intergeneric hybridizations, hybrids of different genera, have also been found in the wild. Although they belong to genera that have been distinct for several million years, interbreeding still occurs between the gelada and the Hamadryas baboon.
- 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;
- nails on the fingers and toes (in most species);
- a flat nail on the hallux (in all extant species);
- sensitive tactile pads on the ends of the digits;
- orbits encircled in bone;
- a trend towards a reduced snout and flattened face, attributed to a reliance on vision at the expense of olfaction (most notably in haplorrhines, and less so in strepsirrhines);
- a complex visual system with stereoscopic vision, high visual acuity and color vision;
- a brain having a well-developed cerebellum with posterior lobe and a calcarine fissure;
- a large brain in comparison to body size, especially in simians;
- differentiation of an enlarged cerebral cortex;
- reduced number of teeth compared to primitive mammals;
- three kinds of teeth;
- a well-developed cecum;
- two pectoral mammary glands;
- typically one young per pregnancy;
- a pendulous penis and scrotal testes;
- a long gestation and developmental period; and
- a trend towards holding the torso upright leading to bipedalism.
Not all primates exhibit these anatomical traits, nor is every trait unique to primates. For example, other mammals have collar bones, three kinds of teeth and a pendulous penis, while spider monkeys have greatly reduced thumbs, ruffed lemurs have six mammary glands and strepsirrhines generally have longer snouts and a strong sense of smell.
In regard to behavior, primates are frequently highly social, with flexible dominance hierarchies. New World species form monogamous pair bonds, and show substantial paternal care of their young, unlike most Old World monkeys.
Anatomy, physiology, and morphology
Primates have forward-facing eyes on the front of the skull; binocular vision allows accurate distance perception, useful for the brachiating ancestors of all great apes. There is a bony ridge above the eye sockets; this ridge 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; in contrast, the higher primates, haplorrhines, have evolved fully enclosed sockets.
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. The endocranial volume (the volume within the skull) is three times greater in humans than in the greatest non-human primate, reflecting a larger brain size. The mean endocranial volume is 1201 cubic centimeters in humans, 469 cm3 in gorillas, 400 cm3 in chimpanzees and 397 cm3 in orangutans. 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. While other mammals rely heavily on their sense of smell, the arboreal life of primates has led to a tactile, visually dominant sensory system, a reduction in the olfactory region of the brain and increasingly complex social behavior.
Primates generally have five digits on each limb (pentadactyly), with keratin nails on the end of each finger. The bottom sides of the hands and feet have sensitive pads on the fingertips. Most have opposable thumbs, a characteristic primate feature; however, opposing thumbs are not limited to this order (opossums, for example, also have them). Thumbs allow some species to use tools. In primates, the combination of opposing thumbs, short fingernails (rather than claws) and long, inward-closing fingers is a relict 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 toe of each foot, called toilet-claws, which they use for grooming.
The primate collar bone 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 lower vertebrae greatly reduced, resulting in tail loss in some species). Old World monkeys are unlike apes in that most have tails. The only primate family with fully prehensile tails are the New World atelids, including the howler, spider, woolly spider and woolly monkeys (New World capuchins have partially prehensile tails).
Primates show an evolutionary trend towards a reduced snout. Technically, Old World monkeys are distinguished from New World monkeys by the structure of the nose, and from apes by the arrangement of their teeth. In New World monkeys the nostrils face sideways; in Old World monkeys, they face downwards. 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. In most strepsirhines, the lower incisors and canines form a toothcomb, which is used in grooming and sometimes foraging, and the first lower premolar is shaped like a canine. Old World monkeys have eight premolars, compared with twelve in New World monkeys. 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, although humans may have 4 or 5. 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, the moist tip of their nose and forward-facing lower front teeth.
The evolution of color vision in primates is unique among most eutherian mammals. While the remote vertebrate ancestors of the primates possessed three color vision (trichromaticism), the nocturnal, warm-blooded, mammalian ancestors lost one of three cones in the retina during the Mesozoic era. Fish, reptiles and birds are therefore trichromatic or tetrachromatic while all mammals, with the exception of some primates and marsupials, are dichromats or monochromats (totally color blind). Nocturnal primates, such as the night monkeys and bush babies, are often monochromatic. Catarrhines are routinely trichromatic due to a gene duplication of the red-green opsin gene at the base of their lineage, 30 to 40 million years ago. Platyrrhines, on the other hand, are trichromatic in a few cases only. Specifically, individual females must be heterozygous for two alleles of the opsin gene (red and green) located on the same locus of the X chromosome. Males, therefore, can only be dichromatic, while females can be either dichromatic or trichromatic. Color vision in strepsirrhines is not as well understood; however, research indicates a range of color vision similar to that found in platyrrhines.
Like catarrhines, howler monkeys (a family of platyrrhines) show routine trichromatism that has been traced to an evolutionarily recent gene duplication. Howler monkeys are one of the most specialized leaf-eaters of the New World monkeys; fruits are not a major part of their diet, and the type of leaves they prefer to consume (young, nutritive, and digestible) are detectable only by a red-green signal. Field work exploring the dietary preferences of howler monkeys suggests that routine trichromaticism was environmentally selected for.
Sexual dimorphism, the variation between individuals of different sex in the same species, is often exhibited in simians, though to a greater degree in Old World species (apes and some monkeys) than New World species. Recent studies involve comparing DNA 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 and canine tooth size along with pelage and skin color. The dimorphism can be attributed to and affected by different factors, including mating system, size, habitat and diet.
Comparative analyses have generated a more complete understanding of the relationship between sexual selection, natural selection, and mating systems in primates. Studies have shown that dimorphism is the product of changes in both male and female traits. Ontogenetic scaling, where relative extension of a common growth trajectory occurs, may give some insight into the relationship between sexual dimorphism and growth patterns. Some evidence from the fossil record suggests that there was convergent evolution of dimorphism, and some extinct hominids probably had greater dimorphism than any living primate.
Primate species move by brachiation, bipedalism, leaping, arboreal and terrestrial quadrupedalism, climbing, knuckle-walking or by a combination of these methods. Several prosimians are primarily vertical clingers and leapers. These include many bushbabies, all indriids (i.e., sifakas, avahis and indris), sportive lemurs, and all tarsiers. Other prosimians are arboreal quadrupeds and climbers. Some are also terrestrial quadrupeds, while some are leapers. Most monkeys are both arboreal and terrestrial quadrupeds and climbers. Gibbons, muriquis and spider monkeys all brachiate extensively, with gibbons sometimes doing so in remarkably acrobatic fashion. Woolly monkeys also brachiate at times. 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. Chimpanzees and gorillas knuckle walk, and can move bipedally for short distances. Although numerous species, such as australopithecines and early hominids, have exhibited fully bipedal locomotion, humans are the only extant species with this trait.
- 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 this close association may be influential in social behavior. The groups formed are generally quite small. This organization can be seen in chimpanzees, where the males, who are typically related, will cooperate in defense of the group's territory. Among New World Monkeys, spider monkeys and muriquis use this system.
- 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 cercopithecine monkeys.
- Monogamous species – a male–female bond, sometimes accompanied by a juvenile offspring. There is shared responsibility of parental care and territorial defense. The offspring leaves the parents' territory during adolescence. Gibbons essentially use this system, although "monogamy" in this context does not necessarily mean absolute sexual fidelity.
- Solitary species – often males who defend territories that include the home ranges of several females. This type of organization is found in the prosimians such as the slow loris. Orangutans do not defend their territory but effectively have this organization.
Other systems are known to occur as well. For example, with howler monkeys both the males and females typically transfer from their natal group on reaching sexual maturity, resulting in groups in which neither the males nor females are typically related. Some prosimians, colobine monkeys and callitrichid monkeys use this system.
Primatologist Jane Goodall, who studied in the Gombe Stream National Park, noted fission-fusion societies in chimpanzees. 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 the Hamadryas baboon, spider monkeys and the bonobo. The gelada has a similar social structure in which many smaller groups come together to form temporary herds of up to 600 monkeys.
These social systems are affected by three main ecological factors: distribution of resources, group size and predation. Within a social group there is a balance between cooperation and competition. Cooperative behaviors include social grooming (removing skin parasites and cleaning wounds), food sharing, and collective defense against predators or of a territory. Aggressive behaviors often signal competition for availability of food, sleeping sites or mates. Aggression is also used in establishing dominance hierarchies.
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. Among the predators of these monkeys is the common chimpanzee.
The red-tailed monkey associates with several species, including the Western red colobus, blue monkey, Wolf's mona monkey, mantled guereza, black crested mangabey and Allen's swamp monkey. Several of these species are preyed upon by the common chimpanzee.
Cognition and communication
Primates have advanced cognitive abilities: some make tools and use them to acquire food and for social displays; some have sophisticated hunting strategies requiring cooperation, influence and rank; they are status conscious, manipulative and capable of deception; they can recognise kin and conspecifics; and they can learn to use symbols and understand aspects of human language including some relational syntax and concepts of number and numerical sequence. Research in primate cognition explores problem solving, memory, social interaction, a theory of mind, and numerical, spatial, and abstract concepts. Comparative studies show a trend towards higher intelligence going from prosimians to New World monkeys to Old World monkeys, and significantly higher average cognitive abilities in the great apes. However, there is a great deal of variation in each group (e.g., among New World monkeys, both spider and capuchin monkeys have scored highly by some measures), as well as in the results of different studies.
Lemurs, lorises, tarsiers, and New World monkeys rely on olfactory signals for many aspects of social and reproductive behavior. 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 behavior of these primates. In Old World monkeys and apes this ability is mostly vestigial, having regressed as trichromatic eyes evolved to become the main sensory organ. Primates also use vocalizations, gestures, and facial expressions to convey psychological state.
Primates have slower rates of development than other mammals. All primate infants are breastfed by their mothers (with the exception of some human cultures and various zoo raised primates which are fed formula) and rely on them for grooming and transportation. In some species, infants are protected and transported by males in the group, particularly males who may be their fathers. Other relatives of the infant, such as siblings and aunts, may participate in its care as well. Most primate mothers cease ovulation while breastfeeding an infant; once the infant is weaned the mother can reproduce again. This often leads to weaning conflict with infants who attempt to continue breastfeeding.
Primates have a longer juvenile period between weaning and sexual maturity than other mammals of similar size. During the juvenile period, primates are more susceptible than adults to predation and starvation; they gain experience in feeding and avoiding predators during this time  They learn social and fighting skills, often through playing.
Primates, especially females, have longer lifespans than other similarly sized mammals. Late in life, female catarrhine primates appear to undergo a cessation of reproductive function known as menopause; other groups are less studied.
Diet and feeding
Primates exploit a variety of food sources. It has been said that many characteristics of modern primates, including humans, derive from an early ancestor's practice of taking most of its food from the tropical canopy. Most primates include fruit in their diets to obtain easily digested carbohydrates and lipids for energy. However, they require other foods, such as leaves or insects, for amino acids, vitamins and minerals. Primates in the suborder Strepsirrhini (non-tarsier prosimians) are able to synthesize vitamin C, like most other mammals, while primates of the suborder Haplorrhini (tarsiers, monkeys and apes) have lost this ability, and require the vitamin in their diet.
Many primates have anatomical specializations that enable them to exploit particular foods, such as fruit, leaves, gum or insects. For example, leaf eaters such as howler monkeys, black-and-white colobuses and sportive lemurs have extended digestive tracts which enable them to absorb nutrients from leaves that can be difficult to digest. Marmosets, which are gum eaters, have strong 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. The aye-aye combines rodent-like teeth with a long, thin middle finger to fill the same ecological niche as a woodpecker. It taps on trees to find insect larvae, then gnaws holes in the wood and inserts its elongated middle finger to pull the larvae out. Some species have additional specializations. For example, the grey-cheeked mangabey has thick enamel on its teeth, enabling it to open hard fruits and seeds that other monkeys cannot.
The gelada is the only primate species that feeds primarily on grass. Tarsiers are the only extant obligate carnivorous primates, exclusively eating insects, crustaceans, small vertebrates and snakes (including venomous species). Capuchin monkeys, on the other hand, can exploit many different types of food, including fruit, leaves, flowers, buds, nectar, seeds, insects and other invertebrates, bird eggs, and small vertebrates such as birds, lizards, squirrels and bats. The common chimpanzee has a varied diet that includes predation on other primate species, such as the western red colobus monkey.
Habitat and distribution
Primates evolved from arboreal animals, and many species live most of their lives in trees. Most primate species live in tropical rain forests. The number of primate species within tropical areas has been shown to be positively correlated to the amount of rainfall and the amount of rain forest area. Accounting for 25% to 40% of the fruit-eating animals (by weight) within tropical rainforests, primates play an important ecological role by dispersing seeds of many tree species.
Some species are partially terrestrial, such as baboons and patas monkeys, and a few species are fully terrestrial, such as geladas 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 Honshū where there is snow-cover eight months of the year; the Barbary macaque lives in the Atlas Mountains of Algeria and Morocco. Primate habitats span a range of altitudes: the black snub-nosed monkey has been found living in the Hengduan Mountains at altitudes of 4,700 meters (15,400 ft), the mountain gorilla can be found at 4,200 meters (13,200 ft) crossing the Virunga Mountains, and the gelada has been found at elevations of up to 5,000 metres (16,000 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.
Interactions between humans and other primates
Close interactions between humans and non-human primates (NHPs) can create pathways for the transmission of zoonotic diseases. Viruses such as Herpesviridae (most notably Herpes B Virus), Poxviridae, measles, ebola, rabies, the Marburg virus and viral hepatitis can be transmitted to humans; in some cases the viruses produce potentially fatal diseases in both humans and non-human primates.
Only humans are recognized as persons and protected in law by the United Nations Universal Declaration of Human Rights.[b] The legal status of NHPs, on the other hand, is the subject of much debate, with organizations such as the Great Ape Project (GAP) campaigning to award at least some of them legal rights. In June 2008, Spain became the first country in the world to recognize the rights of some NHPs when its parliament's cross-party environmental committee urged the country to comply with GAP's recommendations, which are that chimpanzees, bonobos, orangutans, and gorillas not be used for animal experiments.
Many species of NHP are kept as pets by humans. GAP estimates that around 3,000 NHPs live as exotic pets in the United States, while the Humane Society of the United States puts the figure much higher, at around 15,000. The expanding Chinese middle class has increased demand for NHPs as exotic pets in recent years. Although NHP import for the pet trade was banned in the U.S. in 1975, smuggling still occurs along the United States – Mexico border, with prices ranging from US$3000 for monkeys to $30,000 for apes.
Primates are used as model organisms in laboratories and have been used in space missions. They serve as service animals for disabled humans. Capuchin monkeys can be trained to assist quadriplegic humans; their intelligence, memory, and manual dexterity make them ideal helpers.
NHPs are kept in zoos around the globe. Historically, zoos were primarily a form of entertainment, but more recently have shifted their focus to conservation, education and research. Many zoos now feature naturalistic exhibits and educational material for the public; in the United States many participate in the Species Survival Plan (SSP), developed by the Association of Zoos and Aquariums (AZA), to maximize genetic diversity through captive breeding. Zoos and other animal welfare supporters generally oppose animal rights initiatives and the GAP's insistence that all NHPs be released from captivity for two primary reasons. First, captive-born primates lack the knowledge and experience to survive in the wild if released. Second, zoos provide living space for primates and other animals threatened with extinction in the wild.
Role in scientific research
Thousands of non-human primates are used around the world in research because of their psychological and physiological similarity to humans. In particular, the brains and eyes of NHPs more closely parallel human anatomy than those of any other animals. NHPs are commonly used in preclinical trials, neuroscience, ophthalmology studies, and toxicity studies. Rhesus macaques are often used, as are other macaques, African green monkeys, chimpanzees, baboons, squirrel monkeys, and marmosets, both wild-caught and purpose-bred. In 2005, GAP reported that 1,280 of the 3,100 NHPs living in captivity in the United States were used for experiments. In 2004, the European Union used around 10,000 NHPs in such experiments; in 2005 in Great Britain, 4,652 experiments were conducted on 3,115 NHPs. Governments of many nations have strict care requirements of NHPs kept in captivity. In the US, federal guidelines extensively regulate aspects of NHP housing, feeding, enrichment, and breeding. European groups such as the European Coalition to End Animal Experiments are seeking a ban on all NHP use in experiments as part of the European Union's review of animal testing legislation.
The International Union for Conservation of Nature (IUCN) lists more than a third of primates as critically endangered or vulnerable. Trade is regulated, as all species are listed by CITES in Appendix II, except 50 species and subspecies listed in Appendix I, which gain full protection from trade. Common threats to primate species include deforestation, forest fragmentation, monkey drives (resulting from primate crop raiding), and primate hunting for use in medicines, as pets, and for food. Large-scale tropical forest clearing is widely regarded as the process that most threatens primates. More than 90% of primate species occur in tropical forests. The main cause of forest loss is clearing for agriculture, although commercial logging, subsistence harvesting of timber, mining, and dam construction also contribute to tropical forest destruction. In Indonesia large areas of lowland forest have been cleared to increase palm oil production, and 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.
Primates with a large body size (over 5 kg) are at increased extinction risk due to their greater profitability to poachers compared to smaller primates. They reach sexual maturity later and have a longer period between births. Populations therefore recover more slowly after being depleted by poaching or the pet trade. Data for some African cities show that half of all protein consumed in urban areas comes from the bushmeat trade. Endangered primates such as guenons and the drill are hunted at levels that far exceed sustainable levels. This is due to their large body size, ease of transport and profitability per animal. As farming encroaches on forest habitats, primates feed on the crops, causing the farmers large economic losses. Primate crop raiding gives locals a negative impression of primates, hindering conservation efforts.
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 of the larger 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.
In Asia, Hinduism, Buddhism, and Islam prohibit eating primate meat; however, primates are still hunted for food. Some smaller traditional religions allow the consumption of primate meat. The pet trade and traditional medicine also increase demand for illegal hunting. The rhesus macaque, a model organism, was protected after excessive trapping threatened its numbers in the 1960s; the program was so effective that they are now viewed as a pest throughout their range.
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. This increases the amount of forest vulnerable to edge effects such as farmland encroachment, lower levels of humidity and a change in plant life. Movement restriction results in a greater amount of inbreeding, which can cause deleterious effects leading to a population bottleneck, whereby a significant percentage of the population is lost.
There are 21 critically endangered primates, 7 of which have remained on the IUCN's "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 monkey, the Cross River gorilla and the Sumatran orangutan. Miss Waldron's red colobus was recently declared extinct when no trace of the subspecies could be found from 1993 to 1999. A few hunters have found and killed individuals since then, but the subspecies' prospects remain bleak.
- a Humans inhabit every continent if one includes the scientific and meteorological stations in Antarctica.
- b Article 6: Everyone has the right to recognition everywhere as a person before the law.
- ^ a b c d e f g h i Groves, C. (2005). Wilson, D. E., & Reeder, D. M, eds. ed. Mammal Species of the World (3rd ed.). Baltimore: Johns Hopkins University Press. pp. 111–184. OCLC 62265494. ISBN 0-801-88221-4. http://www.bucknell.edu/msw3/browse.asp?id=12100001.
- ^ "Primate". Merriam-Webster Online Dictionary. Merriam-Webster. http://www.merriam-webster.com/dictionary/primate. Retrieved 2008-07-21.
From Old French or 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. Linnaeus thought this the "highest" order of mammals
- ^ Goodman, M., Tagle, D. A., Fitch, D. H., Bailey, W., Czelusniak, J., Koop, B. F., Benson, P. & Slightom, J. L. (1990). "Primate evolution at the DNA level and a classification of hominoids". Journal of Molecular Evolution 30 (3): 260–266. doi:10.1007/BF02099995. PMID 2109087.
- ^ a b c d e f g "Primate". Encyclopædia Britannica Online. Encyclopædia Britannica, Inc.. 2008. http://www.britannica.com/EBchecked/topic/476264/primate. Retrieved 2008-07-21.
- ^ a b c Helen J Chatterjee, Simon Y.W. Ho , Ian Barnes & Colin Groves (2009). "Estimating the phylogeny and divergence times of primates using a supermatrix approach". BMC Evolutionary Biology 9: 259. doi:10.1186/1471-2148-9-259. PMC 2774700. PMID 19860891. http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=2774700
- ^ Anon. (1911), "Ape", Encyclopædia Britannica, XIX (11th ed.), New York: Encyclopædia Britannica, http://www.archive.org/stream/encyclopaediabri02chisrich#page/160/mode/2up, retrieved 2011-07-10
- ^ Dixson, A.F. (1981), The Natural History of the Gorilla, London: Weidenfeld & Nicholson, ISBN 978-0-297-77895-0
- ^ Definitions of paraphyly vary; for the one used here see e.g. Stace, Clive A. (2010a), "Classification by molecules: What’s in it for field botanists?", Watsonia 28: 103–122, http://www.watsonia.org.uk/Wats28p103.pdf, retrieved 2010-02-07 , p. 106
- ^ Definitions of monophyly vary; for the one used here see e.g. Mishler, Brent D (2009), "Species are not Uniquely Real Biological Entities", in Ayala, F.J. & Arp, R., Contemporary Debates in Philosophy of Biology, pp. 110–122, doi:10.1002/9781444314922.ch6, ISBN 978-1-4443-1492-2 , p. 114
- ^ a b Benton, Michael J. (2005), Vertebrate palaeontology, Wiley-Blackwell, ISBN 978-0-632-05637-8, http://books.google.com/books?id=SyJO3vpCk8AC, retrieved 2011-07-10 , p. 371
- ^ Janečka, J. E.; Miller, W., Pringle, T. H., Wiens, F., Zitzmann, A., Helgen, K. M., Springer, M. S. & Murphy, W. J. (2007). "Molecular and Genomic Data Identify the Closest Living Relative of Primates". Science 318 (5851): 792–794. Bibcode 2007Sci...318..792J. doi:10.1126/science.1147555. PMID 17975064. Archived from the original on June 21, 2008. http://web.archive.org/web/20080621092403/http://www.sciencemag.org/cgi/content/full/318/5851/792. Retrieved 2008-08-17.
- ^ Kavanagh, M. (1983). A Complete Guide to Monkeys, Apes and Other Primates. New York: Viking Press. pp. 18. ISBN 0670435430.
- ^ McKenna, M. C. and Bell, S. K. (1997). Classification of Mammals Above the Species Level. New York: Columbia University Press. pp. 329. ISBN 023111012X.
- ^ Williams, B.A.; Kay, R.F.; Kirk, E.C. (2010). "New perspectives on anthropoid origins". Proceedings of the National Academy of Sciences 107 (11): 4797–4804. Bibcode 2010PNAS..107.4797W. doi:10.1073/pnas.0908320107. PMC 2841917. PMID 20212104. http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=2841917.
- ^ "Nova – Meet Your Ancestors". PBS. http://www.pbs.org/wgbh/nova/sciencenow/0303/02-mya-nf.html. Retrieved 2008-10-24.
- ^ "Plesiadapis" (PDF). North Dakota Geological Survey. https://www.dmr.nd.gov/ndfossil/Poster/PDF/Plesiadapis.pdf. Retrieved 2008-10-24.
- ^ Lee, M. (September 1999). "Molecular Clock Calibrations and Metazoan Divergence Dates". Journal of Molecular Evolution 49 (3): 385–391. doi:10.1007/PL00006562. PMID 10473780.
- ^ "Scientists Push Back Primate Origins From 65 Million To 85 Million Years Ago". Science Daily. http://www.sciencedaily.com/releases/2002/04/020418073440.htm. Retrieved 2008-10-24.
- ^ Tavaré, S., Marshall, C. R., Will, O., Soligo, C. & Martin R.D. (April 18, 2002). "Using the fossil record to estimate the age of the last common ancestor of extant primates". Nature 416 (6882): 726–729. doi:10.1038/416726a. PMID 11961552.
- ^ Klonisch, T., Froehlich, C., Tetens, F., Fischer, B. & Hombach-Klonisch, S. (2001). "Molecular Remodeling of Members of the Relaxin Family During Primate Evolution". Molecular Biology and Evolution 18 (3): 393–403. PMID 11230540. http://mbe.oxfordjournals.org/cgi/content/full/18/3/393. Retrieved 2008-08-22.
- ^ a b c d e Horvath, J. et al. (2008). "Development and Application of a Phylogenomic Toolkit: Resolving the Evolutionary History of Madagascar's Lemurs" (PDF). Genome Research 18 (3): 489–499. doi:10.1101/gr.7265208. PMC 2259113. PMID 18245770. http://www.biology.duke.edu/wraylab/papers/Horvath&al_2008.pdf. Retrieved 2008-08-22. [dead link]
- ^ a b Mittermeier, R., Ganzhorn, J., Konstant, W., Glander, K., Tattersall, I., Groves, C., Rylands, A., Hapke, A., Ratsimbazafy, J., Mayor, M., Louis, E., Rumpler, Y., Schwitzer, C. & Rasoloarison, R. (December 2008). "Lemur Diversity in Madagascar". International Journal of Primatology 29 (6): 1607–1656. doi:10.1007/s10764-008-9317-y.
- ^ a b c d Sellers, Bill (2000-10-20). "Primate Evolution" (PDF). University of Edinburgh. pp. 13–17. http://homepage.mac.com/wis/Personal/lectures/human-origins/PrimateEvolution.pdf. Retrieved 2008-10-23.
- ^ a b c d Hartwig, W. (2007). "Primate Evolution". In Campbell, C., Fuentes, A., MacKinnon, K., Panger, M. & Bearder, S.. Primates in Perspective. Oxford University Press. pp. 13–17. ISBN 978-0-19-517133-4.
- ^ Franzen, Jens L. et al. (2009). Hawks, John. ed. "Complete Primate Skeleton from the Middle Eocene of Messel in Germany: Morphology and Paleobiology". PLoS ONE 4 (5): e5723. Bibcode 2009PLoSO...4.5723F. doi:10.1371/journal.pone.0005723. PMC 2683573. PMID 19492084. http://www.plosone.org/doi/pone.0005723.
- ^ Ciochon, R. & Fleagle, J. (1987). Primate Evolution and Human Origins. Menlo Park, California: Benjamin/Cummings. pp. 72. ISBN 9780202011752.
- ^ a b Garbutt, N. (2007). Mammals of Madagascar, A Complete Guide. A&C Black Publishers. pp. 85–86. ISBN 978-0-300-12550-4.
- ^ a b Mittermeier, R.A.; et al. (2006). Lemurs of Madagascar (2nd ed.). Conservation International. pp. 23–26. ISBN 1-881173-88-7.
- ^ Shekelle, M. (2005). "Evolutionary Biology of Tarsiers". http://rmbr.nus.edu.sg/bejc/. Retrieved 2008-08-22.
- ^ Schmidt, T. et al. (2005). "Rapid electrostatic evolution at the binding site for cytochrome c on cytochrome c oxidase in anthropoid primates". Proceedings of the National Academy of Sciences of the United States of America 102 (18): 6379–6384. Bibcode 2005PNAS..102.6379S. doi:10.1073/pnas.0409714102. PMC 1088365. PMID 15851671. http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=1088365.
- ^ Marivaux, L. et al. (2005-06-14). "Anthropoid primates from the Oligocene of Pakistan (Bugti Hills): Data on early anthropoid evolution and biogeography". Proceedings of the National Academy of Sciences of the United States of America 102 (24): 8436–8441. Bibcode 2005PNAS..102.8436M. doi:10.1073/pnas.0503469102. PMC 1150860. PMID 15937103. http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=1150860.
- ^ Schrago, C.G. & Russo, C.A.M. (2003). "Timing the Origin of New World Monkeys" (PDF Reprint). Molecular Biology and Evolution 20 (10): 1620–1625. doi:10.1093/molbev/msg172. PMID 12832653. http://mbe.oxfordjournals.org/cgi/reprint/20/10/1620.pdf.
- ^ Houle, A. (1999). "The origin of platyrrhines: An evaluation of the Antarctic scenario and the floating island model". American Journal of Physical Anthropology 109 (4): 541–559. doi:10.1002/(SICI)1096-8644(199908)109:4<541::AID-AJPA9>3.0.CO;2-N. PMID 10423268.
- ^ Andrews, P. & Kelley, J. (2007). "Middle Miocene Dispersals of Apes". Folia Primatologica 78 (5–6): 328–343. doi:10.1159/000105148. PMID 17855786.
- ^ a b c d e f g h i j k l m n o p q r s t u Strier, K. (2007). Primate Behavioral Ecology (3rd ed.). Allyn & Bacon. pp. 7, 64, 71, 77, 182–185, 273–280, 284, 287–298. ISBN 0-205-44432-6.
- ^ Pough, F. W., Janis, C. M. & Heiser, J. B. (2005) . "Primate Evolution and the Emergence of Humans". Vertebrate Life (7th ed.). Pearson. pp. 650. ISBN 0-13-127836-3.
- ^ Groves, C. P. (2001). Primate Taxonomy. Smithsonian Institution Press. ISBN 1-56098-872-X.
- ^ a b Rylands, A. B. & Mittermeier, R. A. (2009). "The Diversity of the New World Primates (Platyrrhini)". In Garber, P. A., Estrada, A., Bicca-Marques, J. C., Heymann, E. W. & Strier, K. B.. South American Primates: Comparative Perspectives in the Study of Behavior, Ecology, and Conservation. Springer. ISBN 978-0-387-78704-6.
- ^ Linnaeus, C. (1758). Sistema naturae per regna tria Naturae, secundum classes, ordines, genera, species, cum characteribus differentiis, synonimis locis. Tomus I. Impensis direct. Laurentii Salvii, Holmia. pp. 20–32.
- ^ Linnaeus, C. (1735). Sistema naturae sive regna tria Naturae systematice proposita per classes, ordines, genera, & species. apud Theodorum Haak, Lugduni Batavorum. pp. s.p..
- ^ Blainville, H. (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. 268–269.
- ^ Thorington, R. W., Jr. & Anderson, S. (1984). "Primates". In Anderson, S. & Jones, J. K., Jr.. Orders and Families of Recent Mammals of the World. New York: John Wiley and Sons. pp. 187–217.
- ^ McKenna, M. C. & Bell, S. K. (1997). Classification of Mammals: Above the species level. New York: Columbia University Press. pp. 631. ISBN 0-231-11013-8.
- ^ Strier, K. (2007). Primate Behavioral Ecology (Third ed.). pp. 50–53. ISBN 0-295-44432-6.
- ^ Tenaza, R. (1984). "Songs of hybrid gibbons (Hylobates lar × H. muelleri)". American Journal of Primatology 8 (3): 249–253. doi:10.1002/ajp.1350080307.
- ^ a b Bernsteil, I. S. (1966). "Naturally occurring primate hybrid". Science 154 (3756): 1559–1560. Bibcode 1966Sci...154.1559B. doi:10.1126/science.154.3756.1559. PMID 4958933.
- ^ 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.
- ^ Jolly, C. J. et al. (1997). "Intergeneric Hybrid Baboons". International Journal of Primatology 18 (4): 597–627. doi:10.1023/A:1026367307470.
- ^ a b c d e f g h i j k l m n o p q Pough, F. W., Janis, C. M. & Heiser, J. B. (2005) . "Characteristics of Primates". Vertebrate Life (7th ed.). Pearson. pp. 630. ISBN 0-13-127836-3.
- ^ a b Soligo, C., Müller, A.E. (1999). "Nails and claws in primate evolution". Journal of Human Evolution 36 (1): 97–114. doi:10.1006/jhev.1998.0263. PMID 9924135.
- ^ a b c d e f g h i j k l m Macdonald, David (2006). "Primates". The Encyclopedia of Mammals. The Brown Reference Group plc. pp. 290–307. ISBN 0-681-45659-0.
- ^ a b c White, T. & Kazlev, A. (2006-01-08). "Archonta: Primates". Palaeos. http://www.palaeos.com/Vertebrates/Units/480Archonta/480.400.html#Primates. Retrieved 2008-06-03.
- ^ a b c Pough, F. W., Janis, C. M. & Heiser, J. B. (2005) . "Primate Societies". Vertebrate Life (7th ed.). Pearson. pp. 621–623. ISBN 0-13-127836-3.
- ^ Campbell, B. G. & Loy, J. D. (2000). Humankind Emerging (8th edition). Allyn & Bacon. pp. 85. ISBN 0673523640.
- ^ a b Aiello, L. & Dean, C. (1990). An Introduction to Human Evolutionary Anatomy. Academic Press. pp. 193. ISBN 0120455900.
- ^ a b c d e f g Myers, P. (1999). ""Primates" (On-line)". Animal Diversity Web. http://animaldiversity.ummz.umich.edu/site/accounts/information/Primates.html. Retrieved 2008-06-03.
- ^ Ash, M. M., Nelson, S. J. & Wheeler, R. C. (2003). Wheeler's Dental Anatomy, Physiology, and Occlusion. W.B. Saunders. pp. 12. ISBN 9780721693828.
- ^ Arrese, C. A. et al. (2005). "Cone topography and spectral sensitivity in two potentially trichromatic marsupials, the quokka (Setonix brachyurus) and quenda (Isoodon obesulus)". Proceedings of Biological Science 272 (1565): 791–796. doi:10.1098/rspb.2004.3009. PMC 1599861. PMID 15888411. http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=1599861.
- ^ Bowmaker, J. K. & Astell, S. (1991). "Photosensitive and photostable pigments in the retinae of Old World monkeys" (pdf). Journal of Experimental Biology 156 (1): 1–19. ISSN 0022-0949. PMID 2051127. http://jeb.biologists.org/cgi/reprint/156/1/1.pdf. Retrieved 2008-06-16.
- ^ a b Surridge, A. K., and D. Osorio (2003). "Evolution and selection of trichromatic vision in primates". Trends in Ecology and Evolution 18 (4): 198–205. doi:10.1016/S0169-5347(03)00012-0.
- ^ Lucas, P. W. & Dominy, N. J. (2003). "Evolution and function of routine trichromatic vision in primates". Evolution 57 (11): 2636–43. doi:10.1554/03-168. PMID 14686538.
- ^ a b c d e Sussman, R. W. (2003). Primate Ecology and Social Structure, Volume 2: New World Monkeys (Revised First ed.). Needham Heights, MA: Pearson Custom Publishing & Prentice Hall. pp. 77–80, 132–133, 141–143. ISBN 0-536-74364-9.
- ^ Ralls, K. (1976). "Mammals in Which Females are Larger Than Males". The Quarterly Review of Biology 51 (2): 245–76. doi:10.1086/409310. PMID 785524.
- ^ Lindstedtand & Boyce; Boyce, Mark S. (1985). "Seasonality, Fasting Endurance, and Body Size in Mammals". The American Naturalist 125 (6): 873. doi:10.1086/284385.
- ^ Frisch, J. E. (1963). "Sex-differences in the canines of the gibbon (Hylobates lar)". Primates 4 (2): 1–10. doi:10.1007/BF01659148.
- ^ Kay, R. F. (1975). "The functional adaptations of primate molar teeth". American Journal of Physical Anthropology 43 (2): 195–215. doi:10.1002/ajpa.1330430207. PMID 810034.
- ^ Crook, J. H. (1972). "Sexual selection, dimorphism, and social organization in the primates". In Campbell, B. G.. Sexual selection and the descent of man. Aldine Transaction. pp. 246. ISBN 978-0202020051.
- ^ a b Cheverud, J. M., Dow, M. M. & Leutenegger, W. (1985). "The quantitative assessment of phylogenetic constraints in comparative analyses: Sexual dimorphism in body weight among primates". Evolution 39 (6): 1335–1351. doi:10.2307/2408790. JSTOR 2408790.
- ^ Leutenegger, W. & Cheverud, J. M. (1982). "Correlates of sexual dimorphism in primates: Ecological and size variables". International Journal of Primatology 3 (4): 387–402. doi:10.1007/BF02693740.
- ^ a b Plavcan, J. M. (2001). "Sexual dimorphism in primate evolution". American Journal of Physical Anthropology 33: 25–53. doi:10.1002/ajpa.10011. PMID 11786990.
- ^ O'Higgins, P. & Collard, M. (2002). "Sexual dimorphism and facial growth in papionine monkeys". Journal of Zoology 257 (2): 255–272. doi:10.1017/S0952836902000857.
- ^ Sussman, R. W. (1999). Primate Ecology and Social Structure Volume 1: Lorises, Lemurs and Tarsiers. Needham Heights, MA: Pearson Custom Publishing & Prentice Hall. pp. 78, 89–90, 108, 121–123, 233. ISBN 0-536-02256-9.
- ^ Wrangham, R. W. (1982). "Mutualism, kinship and social evolution". Current Problems in Sociobiology. Cambridge University Press. pp. 269–289. ISBN 0521242037.
- ^ Fiore, A. D. & Campbell, C. J. (2007). "The Atelines". In Campbell, C. J., Fuentes, A., MacKinnon, K. C., Panger, M. & Bearder, S. K.. Primates in Perspective. Oxford University Press. pp. 175. ISBN 978-0-19-517133-4.
- ^ Bartlett, T. Q. (2007). "The Hylobatidae". In Campbell, C. J., Fuentes, A., MacKinnon, K. C., Panger, M. & Bearder, S. K.. Primates in Perspective. Oxford University Press. pp. 283. ISBN 978-0-19-517133-4.
- ^ Knott, C. D. & Kahlenberg, S. M. (2007). "Orangutans in Perspective". In Campbell, C. J., Fuentes, A., MacKinnon, K. C., Panger, M. & Bearder, S. K.. Primates in Perspective. Oxford University Press. pp. 294. ISBN 978-0-19-517133-4.
- ^ Constable, J. L. et al. (2001). "Noninvasive paternity assignment in Gombe chimpanzees". Molecular Ecology 10 (5): 1279–1300. doi:10.1046/j.1365-294X.2001.01262.x. PMID 11380884.
- ^ a b c d e Rowe, N. (1996). The Pictorial Guide to the Living Primates. Pogonias Press. pp. 4, 139, 143, 154, 185, 223. ISBN 0-9648825-0-7.
- ^ Smuts, B.B., Cheney, D.L. Seyfarth, R.M., Wrangham, R.W., & Struhsaker, T.T. (Eds.) (1987). Primate Societies. Chicago: University of Chicago Press for articles on the structure and function of various primate societies.
- ^ Shultz, S. & Thomsett, S. (2007). "Interactions between African Crowned Eagles and Their Prey Community". In McGraw, W., Zuberbuhler, K. & Noe, R.. Monkeys of Tai Forest, An African Primate Community. Cambridge University Press. pp. 181. ISBN 0-521-81633-5.
- ^ a b Bshary, R. (2007). "Interactions between Red Colobus Monkeys and Chimpanzees". In McGraw, W., Zuberbuhler, K. & Noe, R.. Monkeys of Tai Forest, An African Primate Community. Cambridge University Press. pp. 155–170. ISBN 0-521-81633-5.
- ^ a b Stanford, C. (1998). Chimpanzee and Red Colobus : the ecology of predator and prey. Harvard University Press. pp. 130–138, 233. ISBN 0-674-00722-0.
- ^ a b Boinski, S. (2000). "Social Manipulation Within and Between Troops Mediates Primate Group Movement". In Boinski, S. and Garber, P.. On the Move : how and why animals travel in groups. University of Chicago Press. pp. 447–448. ISBN 0-226-06340-2.
- ^ Boesch, C. & Boesch, H. (1990). "Tool Use and Tool Making in Wild Chimpanzees". Folia Primatol 54 (1–2): 86–99. doi:10.1159/000156428. PMID 2157651.
- ^ Westergaard, G. C. et al. (1998). "Why some capuchin monkeys (Cebus apella) use probing tools (and others do not)". Journal of Comparative Psychology 112 (2): 207–211. doi:10.1037/0735-7036.112.2.207. PMID 9642788.
- ^ de Waal, F. B. M. & Davis, J. M. (2003). "Capuchin cognitive ecology: cooperation based on projected returns". Neuropsychologia 41 (2): 221–228. doi:10.1016/S0028-3932(02)00152-5. PMID 12459220.
- ^ Paar, L. A., Winslow, J. T., Hopkins, W. D. & de Waal, F. B. M. (2000). "Recognizing facial cues: Individual discrimination by chimpanzees (Pan troglodytes) and rhesus monkeys (Macaca mulatta)". Journal of Comparative Psychology 114 (1): 47–60. doi:10.1037/0735-7036.114.1.47. PMC 2018744. PMID 10739311. http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=2018744.
- ^ Paar, L. A. & de Waal, F. B. M. (1999). "Visual kin recognition in chimpanzees". Nature 399 (6737): 647–648. Bibcode 1999Natur.399..647P. doi:10.1038/21345. PMID 10385114.
- ^ Fujita, K., Watanabe, K., Widarto, T. H. & Suryobroto, B. (1997). "Discrimination of macaques by macaques: The case of sulawesi species". Primates 38 (3): 233–245. doi:10.1007/BF02381612.
- ^ Call, J. (2001). "Object permanence in orangutans (Pongo pygmaeus), chimpanzees (Pan troglodytes), and children (Homo sapiens)". Journal of Comparative Psychology 115 (2): 159–171. doi:10.1037/0735-7036.115.2.159. PMID 11459163.
- ^ Itakura, S. & Tanaka, M. (1998). "Use of experimenter-given cues during object-choice tasks by chimpanzees (Pan troglodytes), an orangutan (Pongo pygmaeus), and human infants (Homo sapiens)". Journal of Comparative Psychology 112 (2): 119–126. doi:10.1037/0735-7036.112.2.119. PMID 9642782.
- ^ Gouteux, S., Thinus-Blanc, C. & Vauclair, J. (2001). "Rhesus monkeys use geometric and nongeometric information during a reorientation task". Journal of Experimental Psychology: General 130 (3): 505–519. doi:10.1037/0096-3418.104.22.1685.
- ^ Tomasello, M. & Call, J. (1997). Primate Cognition. Oxford University Press US. ISBN 9780195106244.
- ^ a b c Deaner, R. O.; van Schaik, C. P.; Johnson, V.E. (2006). "Do some taxa have better domain-general cognition than others? A metaanalysis of nonhuman primate studies". Evolutionary Psychology 4: 149–196. http://www.epjournal.net/filestore/ep04149196.pdf. Retrieved 2011-07-04.
- ^ a b c Reader, S. M.; Hager, Y.; Laland, K. N. (2011-04-12). "The evolution of primate general and cultural intelligence". Philosophical Transactions of the Royal Society B: Biological Sciences 366 (1567): 1017–1027. doi:10.1098/rstb.2010.0342. http://lalandlab.st-andrews.ac.uk/pdf/Publication163.pdf. Retrieved 2011-07-04.
- ^ Liman, E. R. & Innan, H. (2003). "Relaxed selective pressure on an essential component of pheromone transduction in primate evolution" (pdf). Proceedings of the National Academy of Sciences of the United States of America 100 (6): 3328–3332. Bibcode 2003PNAS..100.3328L. doi:10.1073/pnas.0636123100. PMC 152292. PMID 12631698. http://www.pnas.org/cgi/reprint/100/6/3328.pdf. Retrieved 2008-07-23.
- ^ Egnor, R., Miller, C. & Hauser, M.D. (2004). "Nonhuman Primate Communication" (pdf). Encyclopedia of Language and Linguistics (2nd ed.). Elsevier. ISBN 0080442994. http://www.wjh.harvard.edu/~mnkylab/publications/animalcommunication/PrimateComm_ElsevierEncy.pdf. Retrieved 2008-07-23.
- ^ Walker ML, Herndon JG (2008). "Menopause in nonhuman primates?". Biology of Reproduction 79 (3): 398–406. doi:10.1095/biolreprod.108.068536. PMC 2553520. PMID 18495681. http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=2553520.
- ^ Milton, C. (1993). "Diet and Primate Evolution". Scientific American 269: 70–77. Bibcode 1993SciAm.269...70O.
- ^ Pollock, J. I. & Mullin, R. J. (1986). "Vitamin C biosynthesis in prosimians: Evidence for the anthropoid affinity of Tarsius". American Journal of Physical Anthropology 73 (1): 65–70. doi:10.1002/ajpa.1330730106. PMID 3113259. http://www3.interscience.wiley.com/journal/110488482/abstract?CRETRY=1&SRETRY=0.
- ^ Milliken, G. W., Ward, J. P. & Erickson, C. J. (1991). "Independent digit control in foraging by the aye-aye (Daubentonia madagascariensis)". Folia Primatologica 56 (4): 219–224. doi:10.1159/000156551. PMID 1937286.
- ^ Hiller, C. (2000). "Theropithecus gelada". Animal Diversity Web. http://animaldiversity.ummz.umich.edu/site/accounts/information/Theropithecus_gelada.html. Retrieved 2008-08-08.
- ^ Wright, P., Simmons, E. & Gursky, S. (2003). "Introduction". In Wright, P., Simmons, E. & Gursky, S.. Tarsiers Past, Present and Future. Rutgers University Press. pp. 1. ISBN 0-8135-3236-1.
- ^ Reed, K. & Fleagle, J. (August 15. 1995). "Geographic and climatic control of primate diversity". Proceedings of the National Academy of Sciences of the United States of America 92 (17): 7874–7876. Bibcode 1995PNAS...92.7874R. doi:10.1073/pnas.92.17.7874. PMC 41248. PMID 7644506. http://www.pnas.org/content/92/17/7874.full.pdf+html.
- ^ Chapman, C. & Russo, S. (2007). "Primate Seed Dispersal". In Campbell, C. J., Fuentes, A., MacKinnon, K. C., Panger, M. & Bearder, S. K.. Primates in Perspective. Oxford University Press. pp. 510. ISBN 978-0-19-517133-4.
- ^ Long, Y. C., Kirkpatrick, R. C., Zhong, T., and Xiao, L. (1994). "Report on the distribution, population, and ecology of the Yunnan snub-nosed monkey (Rhinopithecus bieti)". Primates 35 (2): 241–250. doi:10.1007/BF02382060.
- ^ Schaller, G. B. (1963). The Mountain Gorilla: Ecology and Behavior. Chicago: University Chicago Press. ISBN 978-0226736358.
- ^ Wolfe, L. D. & Fuentes, A. (2007). "Ethnoprimatology". In Campbell, C. J., Fuentes, A., MacKinnon, K. C., Panger, M. & Bearder, S. K.. Primates in Perspective. Oxford University Press. pp. 692. ISBN 978-0-19-517133-4.
- ^ Renquist, D. M. & Whitney, R. A. (1987). "Zoonoses Acquired from Pet Primates". Veterinary Clinics of North America, Small Animal Practice 17 (1): 219–240. http://pin.primate.wisc.edu/aboutp/pets/zoonoses.html. Retrieved 2008-08-11.
- ^ a b Cavalieri, P. & Singer, P.. "Declaration on Great Apes". Great Ape Project. http://www.greatapeproject.org/declaration.php. Retrieved 2008-06-16.
- ^ Glendinning, L. (2008-06-26). "Spanish parliament approves 'human rights' for apes". The Guardian. http://www.guardian.co.uk/world/2008/jun/26/humanrights.animalwelfare?gusrc=rss&feed=networkfront. Retrieved 2008-11-10.
- ^ Singer, P. (2008-07-18). "Of great apes and men". The Guardian. http://www.guardian.co.uk/commentisfree/2008/jul/18/animalwelfare.animalbehaviour. Retrieved 2008-11-10.
- ^ Mager, B. (2006-07-06). "The HSUS Applauds the U.S. Senate for Unanimously Passing the Captive Primate Safety Act". Humane Society of the United States. http://www.hsus.org/press_and_publications/press_releases/the_hsus_applauds_senate_cpsa.html. Retrieved 2008-11-10.
- ^ a b Workman, C. (2004). "Primate conservation in Vietnam: toward a holistic environmental narrative". American Anthropologist 106 (2): 346–352. doi:10.1525/aa.2004.106.2.346.
- ^ "IPPL News: The US Pet Monkey Trade". International Primate Protection League. 2003. http://www.aesop-project.org/US_Pet_Monkey_Trade.htm. Retrieved 2008-08-04.
- ^ Bushnell, D. (1958). "The Beginnings of Research in Space Biology at the Air Force Missile Development Center, 1946–1952". History of Research in Space Biology and Biodynamics. NASA. http://history.nasa.gov/afspbio/part1.htm. Retrieved 2008-08-18.
- ^ Blumenthal, D. (1987-06-17). "Monkeys as Helpers To Quadriplegics At Home". The New York Times. http://query.nytimes.com/gst/fullpage.html?sec=health&res=9B0DE5D81231F934A25755C0A961948260. Retrieved 2008-10-08.
- ^ a b "The supply and use of primates in the EU". European Biomedical Research Association. 1996. http://www.ebra.org/ebrabulletin-the-supply-and-use-of-primates-in-the-eu_17.htm. Retrieved 2008-08-18.
- ^ Chen, F. C. & Li, W. H. (2001). "Genomic divergences between humans and other hominoids and the effective population size of the common ancestor of humans and chimpanzees". American Journal of Human Genetics 68 (2): 444–456. doi:10.1086/318206. PMC 1235277. PMID 11170892. http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=1235277.
- ^ Conlee, K. M., Hoffeld, E. H. & Stephens, M. L. (2004). "A Demographic Analysis of Primate Research in the United States" (pdf). ATLA (Alternatives to Laboratory Animals) 32 (Sup 1): 315–322. Archived from the original on June 25, 2008. http://web.archive.org/web/20080625161118/http://www.worldcongress.net/2002/proceedings/C2%20Conlee.pdf. Retrieved 2008-10-08.
- ^ presented to Parliament by the Secretary of State for the Home Department by Command of Her Majesty, July 2006. (July 2006) (pdf). Statistics of scientific procedures on living animals: Great Britain 2005. The Stationery Office. ISBN 0101687729. http://www.official-documents.gov.uk/document/cm68/6877/6877.pdf. Retrieved 2008-06-16.
- ^ "Nonhuman Primates: Research Animals". Animal Welfare Information Center. United States Department of Agriculture. 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. Retrieved 2008-07-14.
- ^ "Directive 86/609". European Coalition to End Animal Experiments. http://www.eceae.org/a1_directive.php. Retrieved 2008-10-08.
- ^ IFAW (2005) (pdf). Born to be wild: Primates are not pets. International Fund for Animal Welfare. http://www.ifaw.org/Publications/Program_Publications/Wildlife_Trade/Campaign_Scientific_Publications/asset_upload_file812_49478.pdf. Retrieved 2011-02-26.
- ^ CITES (2010-10-14). "Appendices I, II and III". Convention on International Trade in Endangered Species of Wild Fauna and Flora. http://www.cites.org/eng/app/appendices.shtml. Retrieved 2011-02-26.
- ^ Grubb, P. (1998). "The Sierra Leone monkey drives". Mammals of Ghana, Sierra Leone, and the Gambia. St. Ives: Trendrine. pp. 214–219. ISBN 0951256246.
- ^ a b Chapman, C. A. & Peres, C. A. (2001). "Primate conservation in the new millennium: the role of scientists". Evolutionary Anthropology 10: 16–33. doi:10.1002/1520-6505(2001)10:1<16::AID-EVAN1010>3.0.CO;2-O.
- ^ a b Mittermeier, R. A. & Cheney, D. L. (1987). "Conservation of primates and their habitats". In Smuts, B. B., Cheney, D. L., Seyfarth, R. M., Wrangham, R. W. & Struhsaker, T. T.. Primate Societies. Chicago: University of Chicago Press. pp. 477–490.
- ^ a b Southwick, C. H. & Siddiqi, M. F. (2001). "Status, conservation and management of primates in India" (pdf). Envis Bulletin: Wildlife and Protected Areas 1 (1): 81–91. Archived from the original on October 12, 2003. http://web.archive.org/web/20031012162005/http://www.wii.gov.in/envis/primates/downloads/page81statusofprimates.pdf. Retrieved 2008-08-04.
- ^ a b c d Cowlishaw, G. & Dunbar, R. (2000). Primate Conservation Biology. Chicago: University of Chicago Press. ISBN 9780226116372.
- ^ Van Schaik, C. P., Monk, K. A. & Robertson, J. M. Y. (2001). "Dramatic decline in orangutan numbers in the Leuser Ecosystem, northern Sumatra". Oryx 35 (1): 14–25. doi:10.1046/j.1365-3008.2001.00150.x.
- ^ Purvis, A., Gittleman, J. L., Cowlishaw, G. & Mace, G. M. (2000). "Predicting extinction risk in declining species". Proceedings of the Royal Society of London 267 (1456): 1947–1952. doi:10.1098/rspb.2000.1234. PMC 1690772. PMID 11075706. http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=1690772.
- ^ a b c Fa, J. E., Juste, J., Perez de Val, J. & Castroviejo, J. (1995). "Impact of market hunting on mammal species in Equatorial Guinea". Conservation Biology 9 (5): 1107–1115. doi:10.1046/j.1523-1739.1995.9051107.x.
- ^ Hill, C. M. (1997). "Crop-raiding by wild vertebrates: The farmer's perspective in an agricultural community in western Uganda". International Journal of Pest Management 43 (1): 77–84. doi:10.1080/096708797229022.
- ^ Hill, C. M. (2002). "Primate conservation and local communities: Ethical issues and debates". American Anthropologist 104 (4): 1184–1194. doi:10.1525/aa.2002.104.4.1184.
- ^ Choudhury, A. (2001). "Primates in Northeast India: an overview of their distribution and conservation status" (pdf). Envis Bulletin: Wildlife and Protected Areas 1 (1): 92–101. Archived from the original on November 16, 2004. http://web.archive.org/web/20041116211303/http://www.wii.gov.in/envis/primates/downloads/page92primatesne.pdf. Retrieved 2008-08-04.
- ^ Kumara, H. N. & Singh, M. (2004). "Distribution and abundance of primates in rainforests of the Western Ghats, Karnataka, India and the conservation of Macaca silenus". International Journal of Primatology 25 (5): 1001–1018. doi:10.1023/B:IJOP.0000043348.06255.7f.
- ^ Nijman, V. (2004). "Conservation of the Javan gibbon Hylobates moloch: population estimates, local extinction, and conservation priorities" (pdf). The Raffles Bulletin of Zoology 52 (1): 271–280. http://rmbr.nus.edu.sg/rbz/biblio/52/52rbz271-280.pdf. Retrieved 2008-08-04.
- ^ O'Brien, T. G., Kinnaird, M. F., Nurcahyo, A., Iqbal, M. & Rusmanto, M. (2004). "Abundance and distribution of sympatric gibbons in a threatened Sumatran rain forest". International Journal of Primatology 25 (2): 267–284. doi:10.1023/B:IJOP.0000019152.83883.1c.
- ^ Estrada, A., Coates-Estrada, R. & Meritt, D. (1994). "Non-flying mammals and landscape changes in the tropical forest region of Los Tuxtlas, Mexico". Ecography 17 (3): 229–241. doi:10.1111/j.1600-0587.1994.tb00098.x.
- ^ Marsh, L. K. (2003). "The nature of fragmentation.". In Marsh, L. K.. Primates in Fragments: Ecology and Conservation. New York: Kluwer Academic/Plenum Publishers. pp. 1–10. ISBN 0306476967.
- ^ Turner, I. M. (1996). "Species loss in fragments of tropical rain forest: a review of the evidence". Journal of Applied Ecology 33 (2): 200–209. doi:10.2307/2404743. JSTOR 2404743.
- ^ Chiarello, A.G. (2003). "Primates of the Brazilian Atlantic forest: the influence of forest fragmentation on survival". In Marsh, L. K.. Primates in Fragments: Ecology and Conservation. New York: Kluwer Academic/Plenum Publishers. pp. 99–121. ISBN 9780306476969.
- ^ Pope, T.R. (1996). "Socioecology, population fragmentation, and patterns of genetic loss in endangered primates". In Avise, J. & Hamrick, J.. Conservation Genetics: Case Histories from Nature. Norwell: Kluwer Academic Publishers. pp. 119–159. ISBN 9780412055812.
- ^ Mittermeier, R.A.; Wallis, J.; Rylands, A.B. et al., eds (2009) (PDF). Primates in Peril: The World's 25 Most Endangered Primates 2008–2010. Illustrated by S.D. Nash. Arlington, VA.: IUCN/SSC Primate Specialist Group (PSG), International Primatological Society (IPS), and Conservation International (CI). pp. 23–26. ISBN 978-1-934151-34-1. http://www.primate-sg.org/PDF/Primates.in.Peril.2008-2010.pdf.
- ^ Oates, J. F., Abedi-Lartey, M., McGraw, W. S., Struhsaker, T. T. & Whitesides, G. H. (2000). "Extinction of a West African Red Colobus Monkey". The Journal of the Society for Conservation Biology 14 (5): 1526–1532. doi:10.1046/j.1523-1739.2000.99230.x.
- ^ McGraw, W. S. (2005). "Update on the Search for Miss Waldron's Red Colobus Monkey". International Journal of Primatology 26 (3): 605–619. doi:10.1007/s10764-005-4368-9.
- ^ "The Universal Declaration of Human Rights". United Nations. 1948. http://www.un.org/Overview/rights.html. Retrieved 2008-12-02.
- David J. Chivers, Bernard A. Wood & Alan Bilsborough, ed (1984). Food Acquisition and Processing in Primates. New York & London: Plenum Press. ISBN 0-306-41701-4
- Primate Info Net
- Primates at Animal Diversity Web
- Primate Research Institute, Kyoto University
- High-Resolution Cytoarchitectural Primate Brain Atlases
- EUPRIM-Net: European Primate Network
- PrimateImages: Natural History Collection
- Tree of Life web project
Extant mammal orders by infraclassKingdom Animalia · Phylum Chordata · Subphylum Vertebrata · (unranked) Amniota Australosphenida Metatheria
Extant primate families by suborder Strepsirrhini Haplorhini
Wikimedia Foundation. 2010.