
A primate is a member of the biological order Primates, the group that contains lemurs, the Aye-aye, lorisids, galagos, tarsiers, monkeys, and apes, with the last category including great apes. With the exception of humans, who inhabit every continent on Earth, most primates live in tropical or subtropical regions of the Americas, Africa and Asia. Primates range in size from the Pygmy Mouse Lemur weighing only 30 grams (1.1 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, and 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.
The Primates order has traditionally been divided into two main groupings: prosimians and simians. Prosimians have characteristics most like those of the earliest primates, and included the lemurs of Madagascar, lorisiforms, Aye-aye and tarsiers. Simians included the monkeys and apes. More recently, taxonomists have created the suborder Strepsirrhini, or "curly-nosed" primates, to include non-tarsier prosimians and the suborder Haplorrhini, or "dry-nosed" primates, to include tarsiers and the simians. Simians are divided into two groups: the platyrrhines ("flat nosed") or New World monkeys of South and Central America and the catarrhine ("narrow nosed") monkeys of Africa and southeastern Asia. The New World monkeys include the capuchin, howler and squirrel monkeys, and the catarrhines include the Old World monkeys (such as baboons and macaques) and the apes. Humans are the only catarrhines that have spread outside of Africa, South Asia, and East Asia, although fossil evidence shows many species once existed in Europe as well.
Considered generalist mammals, primates exhibit a wide range of characteristics. Some primates (including some great apes and baboons) do not live primarily in trees, 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 their 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 most significant 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. Some species live in solitude, others live in male–female pairs, and others live in groups of up to hundreds of members.
The primate lineage is thought go back to at least 65 mya, 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.
In modern cladistic reckonings, the Primates order is monophyletic. The suborder Strepsirrhini, the curly-nosed or "wet-nosed" primates, is generally thought to have split off from the primitive primate line about 63 mya (million years ago), although earlier dates are also supported. The seven strepsirhine families are the four related lemur families and the three remaining families that include the Aye-aye, the lorisids, and the galagos. 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. During the Eocene, most of the northern continents were dominated by two groups, the adapiforms and the omomyids. The former is considered a member of Strepsirrhini, but it does 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 at 160 mya and from India at 90 mya.
For lemurs to be more closely related to each other than other strepsirrhine primates, it is thought that a very small ancestral population came to Madagascar 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 lemur line more recently than the lemurs and lorises split) or a sister group to all the other strepsirrhines. In 2008, the Aye-aye family (Daubentoniidae) was confirmed to be a lemuriform and descended from the same ancestral lemur population that rafted to the island, it is contained within the Chiromyiformes infraorder, forming a sister clade to the lemurs.
The suborder Haplorrhini, the simple-nosed or "dry-nosed" primates, is composed of two sister clades. The prosimian tarsiers in family Tarsiidae (monotypic in its own infraorder Tarsiiformes), represent the most primitive division at about 58 mya. The Simiiformes infraorder emerged about 40 mya, and contains the two clades: the parvorder Platyrrhini that developed in South America and contains New World monkeys, and the parvorder Catarrhini that developed in Africa and contains the 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.
Like the lemurs, the New World monkeys have unclear origins. Molecular sequence studies based on concatenated sequences have yielded wide variations in the estimated date of the divergence between platyrrhines and catarrhines, ranging from 33 to 70 mya, while studies based on mitochondrial sequences suggest a more consistent date of 35 mya.
It has been postulated that there is a single origin for the anthropoids in Africa some migrated and subsequently speciation occurred. It is possible that the anthropoid primates traversed the Atlantic ocean during the Eocene, facilitated by Atlantic Ocean ridges and a lowered sea level, then island-hopped to South America. Once again, a rafting event may explain this transoceanic colonization. Due to continental drift, the young Atlantic Ocean was not nearly as wide as it is today, and 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. Some molecular and fossil studies generally show that modern humans originated in Africa 100–200 tya (thousand years ago).
Although primates are well studied in comparison to other animal groups, several new species have been recently discovered; genetic tests on some populations have revealed previously unrecognised species. 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, MSW3 falls short of current understanding as its collection was completed in 2003; a number of publications have since pushed the number to 424 species and 658 species and subspecies. 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).
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.
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 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 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 prehensile tails are the New World Atelids, including the howler, spider and woolly monkeys.
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 period. 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 clinger 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 use brachiation extensively. Woolly monkeys also sometimes brachiate. 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 the Australopithecines and early hominids, have exhibited fully bipedal locomotion, humans are the only extant species with this trait.
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 predated on by the Common Chimpanzee.
In South America, squirrel monkeys associate with capuchin monkeys. This may have more to do with foraging benefits to the squirrel monkeys rather than anti-predation benefits.
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.
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 non-human primate infants are breastfed by their mothers 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.